DEEP-SEA RESEARCH Supplement toVolume 3 Papers in Marine Biology and Oceanography Dedicated to HENRY BRYANT B I G E L O W By His Former Students and Associates on the occasion of The Twenty-fifth Anniversary of the Founding of The Woods Hole Oceanographic Institution 1955 Distributed with the compliments of the Woods Hole Oceanographic Institution PERGAMON PRESS London & New York I 955 / 3 > MBL/WHOI Library '/I-- /^.c^y^iM — DEEPS i: A RESEAIUIl Supplement to Volume 3 PAPERS IN MARINE BIOLOGY AND OCEANOGRAPHY _ nj ^ i— J CD i ■ o // is .\i;i;gcslc. •>!(■ Slioiihl upi\i-: •'■ ''■ ■ ., H./.\- '.■■'') P«/>. /V/r//. Biol, uiu! ' '•. w//»/>/ /,) I <•/ \ DEEP-SEA RESEARCH ^ Supplement to Volume 3 Papers in Marine Biology / and Oceanography Dedicated to HENRY BRYANT B I G E L O W By His Former Students and Associates on the occasion of The Twenty-fifth Anniversary of the Founding of The Woods Hole Oceanographic Institution 1955 PERGAMON PRESS London & New York 1955 Published by Pergamon Press Ltd., 4 & 5 Fitzroy Square, London, W.L and Pergamon Press Inc., 122 East 55th Street, New York 22, N. Y. Printed in Great Britain by Page Bros. (Norwich) Ltd., Norwich Publication of this volume has been aided by a gram tVom the Woods Hole Oceano- GRAPHic Institution. Individual papers have been supported by the Johannes Schmidt Foundation for Marine Research, by the Agnes Anderson Fund of the University of Washington, by Rutgers University and by the University of Florida. Contributions have been received from the following individuals who wish to honour HENRY BRYANT BIGELOW Charles Francis Adams Jr. Mabel B. Agassiz W. R. G. Atkins Eric G. Ball Charlotte H. Bartol Robert P. Bellows Margaret D. Bigelow Trygve Braarud Mary and Lewis Bremer Anton Fr. Bruun W. Sohier Bryant Dean F. Bumpus Nancy and Samuel Cabot Cornelia L. Carey J. N. Carruthers Fenner a. Chace Jr. Ruth and Richard Chute George L. Clarke Stanley Cobb L. O. Colbert John W. Condon Charles A. Coolidge Leslie H. N. Cooper Elisabeth Deichmann Tilly Edinger Gardner Emmons Marie and Charles J. Fish Alexander Forbes Horace S. Ford Michael Graham H. B. Hachey Jan Hahn Julia A. Hallowell Alistair C. Hardy Frank A. Howard Georgina and Llewellyn Howland A. G. Huntsman Columbus O'D. Iselin Bostwick H. Ketchum Mary and Alfred Kidder II Ellen Knudsen Poul M. Kramp Milford R. Lawrence Benjamin B. Leavitt Lois C. Lillick Charles P. Lyman Elizabeth C. Lyman Jessie Bell Mackenzie Arnaud C. Marts Shefna M. Marshall Ernst Mayr Dorothea F. Merriman William Jason Mixter A. W. H. Needler Margaret S. Nicodemus A. P. Orr Frances L. Parker Louisa B. Parker Hans Pettersson Helen F. Phillips John E. G. Raymont Alfred C. Redfield LaWRASON RlGGS Gordon A. Riley Alfred S. Romer Johan T. Ruud Barbara and William E. Schevill Ingeborg Schmidt Waldo L. Schmitt William C. Schroedfr Mary Sears O. E. Sette Harlow Shapley Virginia and George C. Shattuck Henry L. Shattuck Edward H. Smith Virginia Walker Smith Floyd M. Soule Ragner Sparck Athelstan F. Spilhaus EiNAR Steemann Nielsen Henry C. Stetson Raymond Stevens GuDRUN and Harald U. Sverdrup A. Vedel Taning Thomas G. Thompson Alice Thorndike GUNNAR THORSON A. Knyvett Totton Theodor Von Brand George Wald Lionel A. Walford Talbot H. Waterman Edmond E. Watson Francis C. Welch George C. Whiteley Jr. Florence H. Whitman H. B. Whittington Susan J. Williams R. S. Wimpenny 0. WiNGE Mary A. Winsor Alfred H. Woodcock Donald J. Zinn VI CONTENTS page FOREWORD xi BIBLIOGRAPHY OF HENRY BRYANT BIGELOW xviii Michael Graham: Effect of trawling on animals of the sea bed I A. C. Hardy: A further example of the patchiness of plankton distribution 7 Mary Alys Plunkett and Norris W. Rakestraw: Dissolved organic matter in the sea - 12 Selman a. Waksman: Marine bacteria: recollections and problems 15 Thomas G. Thompson and Tsaihwa J. Chow : The strontium-calcium atom ratio in carbonate-secreting marine organisms 20 Jean Brouardel and Louis Fage: Variation, en mer, de la teneur en oxygene dissous au proche voisinage des sediments 40 Fred B Phleger: Foraminiferal faunas in cores offshore from the Mississippi Delta 45 Pamela G. Jenkins: Seasonal changes in the phytoplankton as indicated by spectrophotometric chlorophyll estimations 1952-53 58 H. B. Hachey: Water replacements and their significance to a fishery 68 Daniel and Mary Merriman: Sir C. Wyville Thomson's correspondence on the " Challenger " fishes 74 Dean F. Bumpus and E. Lowe Pierce: Hydrography and the distribution o^ chaetognaths over the continental shelf off North Carolina 92 S. M. Marshall and A. P. Orr: Experimental feeding of the copepod Ca/fl«w.sy7/;mfl/-c/7/rw5 (Gunner) on phytoplankton cultures labelled with radio- active carbon (^^C) ' •*-' Alfred C. Redfield: The hydrography of the Gulf of Venezuela 115 H. Boschma: The specific characters of the coral Stylaster roseus 134 vii Contents {com.) Page C. E. Lucas: External metabolites in the sea 139 P. L. Kramp: a revision of Ernst Haeckel's determinations of a collection of Medusae belonging to the Zoological Museum of Copenhagen 149 J. N. Carruthers: Some very simple devices for various oceanographical uses 169 J. E. G. Raymont: The fauna of an inter-tidal mud flat 178 Frances L. Parker: Distribution of planktonic Foraminifera in some Mediter- ranean sediments 204 L. H. N. Cooper: Hypotheses connecting fluctuations in Arctic climate with biological productivity of the English Channel . 212 Gordon A. Riley: Review of the oceanography of Long Island Sound 224 A. K. Totton: Development and metamorphosis of the larva of Agalma elegans (Sars) (Siphonophora Physonectae) 239 Charles J. Fish: Observations on the biology of Microsetella norvegica 242 R. S. Wimpenny: The production of Lisa ramada (Risso) in Lake Mariut, Egypt 250 JoHAN T. Ruud: The mortality rates of Antarctic fin whale stocks 257 Alfred Sherwood Romer: Fish origins — fresh or salt water? 261 E. Steemann Nielsen: The production of antibiotics by plankton algae and its effect upon bacterial activities in the sea 281 John H. Welsh : On the nature and action of coelenterate toxins 287 Henry C. Stetson: Patterns of deposition at the continental margin 298 Ruth D. Turner : Scaphopods of the Atlantis dredgings in the Western Atlantic with a catalogue of the Scaphopod types in the Museum of Com- parative Zoology 309 A. G. Huntsman: Effect of freshets on Passamoquoddy plankton 321 R. B. Montgomery: Characteristics of surface water at Weather Ship J. 331 Hans Pettersson: Manganese nodules and oceanic radium 335 BosTWiCK H. Ketchum and D. Jean Keen: The accumulation of river water over the continental shelf between Cape Cod and Chesapeake Bay 346 viii Contents (cont.) Page William C. Schroeder: Report on ihe results of exploratory otter-trawling along the continental shelf and slope between Nova Scotia and Virginia during the summers of 1952 and 1953 358 Georg WiJST : Stromgeschwindigkeiten im Tiefen- und Bodenwasser des Atlantischen Ozeans auf Grund dynamischer der Meteor- Profile der Deutschen Atlantischen Expedition 1925 27 373 Wm. C. Herrington : U.S. participation in conservation of international fishery resources 398 Donald R. Griffin; Hearing and acoustic orientation in marine animals 406 Anton Fr. Bruun and A. Kiilerich: Characteristics of the water-masses of the Philippine, Kermadec, and Tonga Trenches 413 Talbot H. Waterman: Polarization of scattered sunlight in deep water 426 Walter Hansen : Elevation of sea level caused by wind in a rectangular channel 435 A. Vedel Taning: On the breeding areas of the Swordfish {Xiphias) 438 W. T. Edmonson: Factors effecting productivity in fertilized salt water 451 A. Defant: Die Ausbreitung des Mittelmeerwassers in Nordatlantischen Ozean 465 Lionel A. Walford: New directions in fishery research 471 C. O'D. Iselin: Coastal currents and the fisheries 474 Trygve Braarud: Electron microscopy in oceanographic research 479 John B. Tait: Long-term trends and changes in the hydrography oi the Faroe-Shetland Channel region . 48- IX XI FOREWORD The papers collected in this volume have been prepared by the students, disciples and friends of Henry Bryant Bigelow, as a testimonial not only of our personal affection but to honor him for his many contributions to the advancement of oceano- graphy. A half century ago when Alexander Agassiz set sail for the Maldive Islands with Henry Bigelow as his assistant, oceanography in America was an interest promoted from time to time through individual initiative and, when in hne with their primary duties, by appropriate governmental agencies. Today, as his latest monograph on fishes comes off the press, oceanography is a fully recognized division of science, complete with standard textbook and special journals. Its work is instru- mented by half a dozen full-scale laboratories and research vessels operated by university departments or independently, and distributed equably along our coast. More important, it is a science in which a new viewpoint has developed and new vistas have opened. Of course, this has been the work of many men, but in the United States Henry Bigelovv' more than any other has provided the wise leadership which has insured success. In the preface to his book, " Wind Waves at Sea, Breakers and Surf" (written with W. T. Edmondson as a contribution to the World War II effort), it is stated: " We wish it expressly understood that we have made no contributions to the theory of waves. But we would not have dared to undertake the task, if we had not observed the behavior of waves at sea, from large craft and from small, in various parts of the world, under various conditions of wind and weather; or if we had not had an opportunity to watch the development of breakers — and cope with the smaller sizes — off beaches of various shapes, off rocky coastlines, and over submerged ledges ". This insistence on personal experience as a necessary prerequisite of scientific judg- ment (or any other judgment for that matter) is characteristic. By good fortune Henry Bigelow was born into a segment of New England society in which the tradition of plain living and high thinking was graced by the fruits of Yankee enterprise. Young men were expected to receive the best of educa- tion, any natural taste for outdoor life was encouraged and intellectual ambitions were not frowned upon. Summers at Cohasset, on Massachusetts Bay, gave him an instinctive knowledge of seamanship and the things of the sea. Hunting in autumn took him to all parts of the coast and the uplands as well. In winter the mountains were explored on snowshoes, and in later life on ski; the mountains are, in fact, the true love of this oceanographcr. And in the spring there are trout in the New England brooks. Thus he became the best-informed naturalist that one could hope to go afield with. His outdoor life is a routine, fixed by the seasons, and followed with the same insistence on knowing all that is to be known of the matter, which marks his more professional effort. Students of the " Physical Oceanography of the Gulf of Maine " note that the only period not covered by observations at sea coincide with the xii Foreword partridge season. One would no more expect him to be " at work " at that time than to find him missing on the day his course began at Harvard. Henry Bigelow graduated from Harvard College, cum laude, in 1901. As a graduate student in zoology his doctoral dissertation, prepared under Professor E. L. Mark's guidance, was on the nuclear cycle of Gonionemus murbachi. Although he did not pursue cytological studies further, this was a valued experience, for from Mark he first learned the exacting requirements of scientific work. Students among us will find here the source of the discipline to which Henry Bigelow subjected us to our immediate chagrin but ultimate profit. It was inevitable that Henry Bigelow should become a naturaHst of some sort but it was not at all clear during his student days that he would become an oceano- grapher or even a marine biologist. His first publication was on the birds of the northeast coast of Labrador, where he had gone in company with Reginald A. Daly and Merritt L. Fernald on the Brown-Harvard Expedition in the summer before his graduation. A later one was on hybrid ducks. A study of hearing in gold- fish under the guidance of Professor G. H. Parker gave him acquaintance with experimental procedures. The die was cast, however, by the opportunity to accom- pany Alexander Agassiz on the voyage to the Maldive Islands, and, later on expedi- tions to the Eastern Tropical Pacific and to the West Indies. His duty was to care for the medusae and siphonophores collected on these voyages; and thus he gained a first-hand experience and competence in the classical disciplines of taxonomy and zoogeography which occupied the first decades of his mature career. Perhaps more important, he was introduced to the universal problems of oceanography and met first-hand the detailed tasks of scientific research at sea. In 1912 the United States' Bureau of Fisheries and the Museum of Comparative Zoology jointly undertook a general oceanographic exploration of the Gulf of Maine which continued under Henry Bigelow's direction through 1924 when the field work was terminated. These explorations resulted in the publication of three superb monographs; on the fishes, the plankton, and the hydrography of the Gulf. The preparation of the first of these, on the fishes, was far advanced when interrupted by the untimely death of W. W. Welsh who had given special attention to this phase of the work, and was completed by Bigelow at the request of the Bureau. The others are entirely his own work not only in planning and direction but in the execution at sea, in fair weather and foul, in spite of seasickness and with ships and gear far from adequate. It is difficult to appreciate today how primitive were the resources available for this work. Thus during 1912 and 1913 reversing thermometers were accurate only to ± 0.15° C; the shortage of water bottles required repeated casts for all but the shoalest stations. Limited means were, however, more than compensated by the challenge of the unknown. He wrote: *' Few hving zoologists have been as fortunate as were we on setting sail on the Grampus from Gloucester on our first oceanographic cruise in the Gulf of Maine on July 9, 1912, for a veritable mare incognitum lay before us, so far as its floating Ufe was concerned, though the bottom fauna can be described as fairly well-known. Not but what an extensive list of pelagic crustaceans, coelenterates and other plank- tonic animals had been recorded thence, but everything was yet to be learned as to what groups or species would prove predominant in the pelagic fauna; their relative Foreword xiii importance in the natural economy of the Gulf, their geographic and bathymetric variations; their seasonal successions, migrations, and annual fluctuations; their temperature affinities, whether arctic, boreal, or tropic; and whether they were oceanic or creatures of the coastal zone. We even had no idea (incredible though it may seem at this place and day) what we should probably catch when wc first lowered our tow nets into deeper strata of Massachusetts Bay, for, so far as we could learn, tows had never previously been tried more than a few fathoms below its surface." The outcome is that the Gulf of Maine is perhaps the best-known body of water of comparable size in the world; certainly the region most thoroughly explored by individual effort. Except for certain aspects which he did not examine deeply, i.e., submarine geology and sea-water chemistry, a quarter century of subsequent study added only trivial details to the picture. During this period Henry Bigelow was the trusted advisor of the government on fisheries, a trust well earned by his first-hand knowledge of fish and fishermen, and by his incisive, direct, and ever-practical approach to human problems and his understanding of the role which science can play in their solution. A number of men later to hold important posts in the fisheries service, Herrington, Nesbit, Schroeder. Sette, and Walford, were among his students at Harvard. In 1917-19 Henry Bigelow served as Special Expert to the U.S. Shipping Board and during 1918 the work on the Gulf of Maine was interrupted while he did his trick as navigation officer on the U.S. Army transport Amphion. Of more interest in connection with scientific developments were his connections with the International Ice Patrol, established in 1913 as a result of tragic loss of life and property due to the collision of the steamship Titanic with an iceberg. Operation of the patrol became the duty of the U.S. Coast Guard, while the scientific studies necessary for its intelhgent prosecution were directed by an interdepartmental board composed of the heads of the interested agencies. Henry Bigelow was the special consultant to the Commandant of the Coast Guard for the work of this board. During the early years of the patrol observations on plankton as well as surface tem- peratures and salinities, were used to trace the drift of water carrying icebergs into the shipping lanes; later the techniques of dynamic oceanography were introduced to estimate, on the spot, the velocity of the movement. As in the case of the fishery experts a succession of officers of the Coast Guard, Smith, Ricketts, Hoyle, and Graves, came to Cambridge to receive indoctrination in oceanography from Pro- fessor Bigelow. Largely as the result of his wisdom in guiding the scientific studies on which the work of the ice patrol is based, the hydrography of the northern seas is well understood and the patrol has been enabled to discharge its duties with intelli- gence and success. The study of the Gulf of Maine naturally led to intimate contact with Canadians working in adjacent, and often overlapping waters. One fruit of this was a close and continuing friendship with Professor A. G. Huntsman, for many years chairman of the Biological Board of Canada; another was Bigelow's association with the North American Committee on Fishery Investigations, in which Canada, Newfound- land, France, and the United States were associated. He attended the meetings of the committee regularly between 1921 and 1933 and served as chairman at all but a few of them. During this period Henry Bigelow formed associations with the European xiv Foreword leaders in oceanography, marine biology, and fisheries; such men as Johannes Schmidt, Johan Hjort, D'Arcy Thompson, Martin Knudsen, Henry Maurice and many others still living. The esteem and affection which he won from these colleagues is nicely shown by the records of the meeting of the International Council for the Exploration of the Sea, which he attended in March 1931, as a representative of the North American Committee on Fishery Investigations and where he reported on the newly-founded Woods Hole Oceanographic Institution. " The president . . . wished to take opportunity of his being actually present to express to him the satisfaction which his visit had caused to the Council. Dr. Bige- Low . . . had attended many council meetings and had so impressed his personality on the members and experts that the Consultative Committee had passed a recom- mendation ... so important that it ought to be specially treated. In effect it contained a standing invitation to the representatives of the Woods Hole Oceanographic Institution and the North American Council on Fisheries Investigations and he might add to Dr. Bigelow personally, whatever his future might be, to attend all meetings of the Council. The Council hoped in future to have many opportunities to consult them, to learn from them and to link up its own investigations with the work done on the western side of the Atlantic." When in 1927 a committee of the National Academy of Sciences engaged Henry Bigelow as its secretary to prepare a report on the share of the United States in a world-wide program of oceanographic research, no one could have been found so well equipped by personal experience or general ability for the task. The greater part of this report, reviewing the scope, problems, and applications of oceanography, has been made public in a book entitled " Oceanography " pubHshed under his name in 1931. It is in the unpublished sections of this report, however, in which are set forth the principles that should determine the type of organization which would best remedy the then-present handicaps to the development of oceanography, that his genius for striking directly at the heart of any question and his power of exposition are superbly displayed. It is no wonder that this report was received with confidence, or that it led to the establishment of a new institution at Woods Hole and to sub- stantial benefits to oceanography and marine biology through gifts to the Scripps Institution, the University of Washington, and the Bermuda Biological Station, And it was inevitable that the author of this report should have been asked to direct the newly-established Woods Hole Oceanographic Institution. The principle of the ripeness of time, as applied to the appearance of prophets, is well illustrated by the history of oceanography at this period. Not only did a man emerge who had prepared himself, perhaps unwittingly, for leadership at a time when men of influence had sensed that something should be done to improve the status of marine science in America, but new ideas were in the air wafted across the ocean from a multitude of general scientific advances. Henry Bigelow, though trained in the classical tradition, was sensitive to these breezes, bold enough to grasp their implication, and wise enough to act on their meaning. The following paragraphs, excerpted from a paper pubhshed in Science in 1930, entitled " A developing viewpoint in Oceanography ", express in Bigelow's own words the creed which was to guide his thinking. " Oceanography has of late entered a new intellectual phase, to explain which a word of retrospect is necessary . . . Students of the history of science may well date Foreword XV the birth of modern oceanography from December 21, 1872. the day when the Challenger set sail from Plymouth, England, on her memorable voyage. . . . One great deep-sea expedition led to another, and more was learned about the sea during the last thirty years of the nineteenth century than had been during the preceding three thousand. But after a time, as so often happens when some scientific discipline takes a sudden spurt, this fact-finding began to lose something of its freshness. " Students began, in short, to feel that the mere accumulation of facts from the sea, when there is an inexhaustible supply, may actually become a bit sterile, just as catching fish is to a sportsman where fish are too plentiful. So it was natural that when persistence in the old methods no longer yielded startling discoveries, signs could be seen of the approach of a period of stagnation . And oceanography would probably be in a moribund state in America today, just as the art of sailing a square-rigger is, but for the birth of the new idea that what is really interesting in sea science is the fitting of these facts to gether, and that enough facts had accumu- lated to make the time ripe for an attempt to lift the veil that had obscured (and still obscures) any real understanding of the marvelously complex and equally marvel- ously regulated cycle of events that take place within the sea. " The foundation for this conscious alteration in view-point, from the descriptive to the explanatory, was a growing realization . . . that the further development of sea science the keynote must be physical, chemical and biological unity. . . . " When one picks up a fish one may be said, allegorically, to hold one of the knots in an endless web of netting of which the countless other knots represent other facts, whether of marine chemistry, physics or geology, or other animals and plants. And just as one cannot make a fish-net until one has tied all the knots in their proper positions, so one cannot hope to comprehend this web until one can see its internodes in their true relationship. This is today the conscious aim of oceanographers." Newcomers may feel surprise that this viewpoint had novelty, for it is still our guiding principle. But therein lies meaning. The task of assembling a staff for the new Oceanographic Institution at Woods Hole was not an easy one for there was little raw material with which to work. There were a few young men with some experience at sea, and by combing the museums of the country doubtlessly he could have assembled a respectable group of experts on special groups of marine organisms. A primary objective, however, was to give impetus to oceanographic studies in the universities, and there was the " developing viewpoint " to be fostered. He chose the bolder course of educating a new generation drawn from the universities; physical chemists, meteorologists, physiologists, bac- teriologists, whoever could be persuaded that scope for their skills could be found in studies at sea. And so the practice grew that each should make at least one short voyage at sea each season. Daily the director made his rounds, instilling little by little something of his viewpoint and wisdom on the opportunities that lay beyond the tide fine. Boldness was encouraged for we were told that an oceanographer, like a turtle, made progress only by sticking his neck out. After ten years as Director of the Woods Hole Oceanographic Institution. Henry Bigelow asked to be relieved. The painstaking labour of creating and guiding the development of the Institution had been a sacrifice to the general welfare, made at the expense of his own natural preference for first-hand scientific investigation. He continued to guide the Institution, at first as President of the Corporation and. xvi Foreword now, as Chairman of the Board of Trustees, but was thereafter able to devote full time to his interests at Harvard University. Harvard University appears to have been a bit slow in recognizing the merits of Henry Bigelow. Tradition has it that, during his first year of service as assistant in the invertebrate zoology laboratory, he discovered a student who had depicted a tunicate embellished with a complete set of neatly labelled mammalian viscera, based rather on " natural logic " than on direct observation. This intellectual dis- honesty, or stupidity, so enraged him that the unfortunate student was told off in words so monosyllabic and unambiguous that the young assistant was never again to be permitted to have contact with the students in Harvard College. While the inference is perhaps apocryphal there is no doubt that the incident is authentic. However this may be, so far as Harvard College was concerned Henry Bigelow remained relatively obscure, serving as curator and lecturer in the Museum of Comparative Zoology up to the time of the events leading to the establishment of the Oceanographic Institution. This may have resulted from the almost complete lack of intercourse between the Museum and the Department of Zoology during the period when these institutions existed under the same roof. At all events, it was a most fortunate state of affairs, for had he been burdened with the ordinary academic routine his achievements during this period would have been impossible. It is a tribute to the liberal policies of the University at that time that such great talent was enabled to fruit without distraction. In 1931 Henry Bigelow became a full professor at Harvard and inaugurated a course in biological oceanography. Some years later this course passed into the hands of one of his disciples. Dr. George L. Clarke, and Bigelow took over instruc- tion in invertebrate zoology. This was a task he could put his heart into for he felt that nowhere else is the wonderful diversity of form with which organisms are en- dowed so well displayed as among the invertebrates. His obligation as a professor to the students in Harvard College was to him most sacred. It is too bad they could not have had more of him. Those who benefitted most from his talents as a teacher (and taskmaster) were the succession of graduate students, both men and women, who had the privilege of working at his side in the Museum, and those whom he in- fluenced by indirection at the Oceanographic Institution or wherever else he came in contact with thinking people. As the years passed the counsel of Henry Bigelow, early recognized for its worth in the Museum of Comparative Zoology, became more and more influential in the Department of Biology and in Harvard University at large. On reaching the ordinary age of retirement the University asked him to continue in service, a very real honour, until the mandatory age of seventy was reached. In 1946 he was granted an honorary degree by Harvard and in the same year by the University of Oslo, similar recognition having been made by Yale University some years earlier. Among other formal honours he is the recipient of the Johannes Schmidt Medal, the Agassiz Medal awarded by the National Academy of Sciences for contributions to Oceanography, and the Bowie Medal of the American Geophysical Union in recognition of accomplishment through cooperative effort in the advancement of the geophysical sciences. He has also been elected to membership in the National Academy, the American Academy of Arts and Sciences, and the Philosophical Society, and is affiUated with the Norske Videnskaps Academy, the Royal Foreword xvii Geographical Society of London, the Zoological Society of London, and the Marine Biological Society of the United Kingdom. The crowning glory of these later years is not, however, the honours which come his way. It is the stream of publications which flowed from his pen, always in association with William C. Schroeder, about fishes. This flow shows no attenua- tion with time; it reached, in fact, its spate after retirement from academic duty, as a glance at the appended bibliography will show. The handsome monographs on the Fishes of the North-western Atlantic show where his heart really lies, for he was free to follow its guidance once he had played his part in putting American Oceano- graphy on its feet. Henry Bryant Bigelow, we greet you on this the twenty-fifth anniversary of the opening of the laboratory of the Woods Hole Oceanographic Institution. " We '" are the authors of the papers appended hereto. If they are diverse in subject matter it is because you are catholic in the interests we share in common. If they appear disconnected it is because we have individually failed to achieve the physical, chemical, and biological unity to which you have encouraged us to aspire. " We " are also many others who have been unable to contribute a paper, for reasons which you can comprehend, but who none the less wish to join in this testimonial. You have broadened the vision, sharpened the perception, fortified the deter- mination, simplified the outlook, improved the standards, and corrected the folly of each of us. We continue to come to you for counsel. You have always been your own excuse for being, and to all of us it is a joy to be with you. We hope that what is written will not offend your Yankee reticence. We know it will not inflate your pride, for that is built of something tough. You are an "* indivi- dual " and we have hailed you as something of a prophet. That is a combination from which legends grow. Your legend can afford to be correct; it is not necessary to exaggerate the truth. We have followed your precepts in setting down what can be learned about you from the written word and the observed fact. In the inferences drawn and the judgments passed we hope we have not stuck the neck out too far. We will be happy if this volume pleases you. BIBLIOGRAPHY OF HENRY BRYANT BIGELOW ;i902 (1904 (1904 [1905 :1907 1907 (1909 :1909 1909 U911 (1911 ;1911 ;i9ii (1912 (1912 (1913) (1913) (1913) (1914) (1914) :1914) (1914) (1915) (1915) (1916) (1917) Birds of the northeastern coast of Labrador. The Auk, 19, 24-3 1 . Medusae from the Maldive Islands. Bull. Mus. Comp. ZooL, Harvard Coll., 39 (9), 245-269, 9 pis. The sense of hearing in the goldfish, Carassius auratus L. Amer. Natur., 38 (448), 275-284. The shoal-water deposits of the Bermuda Banks. Proc. Amer. Acad. Arts and Sci., 40 (15), 559-592. On hybrids between the mallard (Anas boschas) and certain other ducks. The Auk, 24 (4), 382-388. Studies on the nuclear cycle of Gonionenms murbachii A. G. Mayer. Bull. Mus. Comp. ZooL, Harvard Coll., 48 (4), 287-399, 8 pis. Coelenterates from Labrador and Newfoundland, collected by Mr. Owen Bryant from July to October, 1908. Proc. U.S. Nat. Mus., 37 (1706), 301-320, Pis. 30-32. Cruise of the U.S. Fisheries Schooner Grampus in the Gulf Stream during July, 1908, with description of a new Medusa (Bythotiaridae). Bull. Mus. Comp. ZooL, Harvard Coll., 52 (12), 195-210, 1 pi. Report on the scientific results of the Expedition to the Eastern Tropical Pacific, in charge of Alexander Agassiz, by the U.S. Fish Commission Steamer Albatross, from October, 1904, to March, 1905, Lieut. Commander L. M. Garrett, U.S.N., Commanding. XVL The Medusae. Mem. Mus. Comp. ZooL, Harvard Coll., 37, 243 pp., 48 pis. Biscayan plankton collected during a cruise of H.M.S. Research, 1900. XIIL The Siphono- phora. Trans. Linn. Soc, London, (2nd ser., ZooL), 10 (10), 337-358, PI. 28. Report on the scientific results of the Expedition to the Eastern Tropical Pacific, in charge of Alexander Agassiz, by the U.S. Fish Commission Steamer Albatross, from October, 1904, to March, 1905, Lieut. Commander L. M. Garrett, U.S.N. , Commanding. XXIIL The Siphonophorae. Mem. Mus. Comp. ZooL, Harvard Coll., 38 (2), 173^02, 32 pis. The work of the Michael Sars in the North Atlantic in 1910. (A review.) Science, n.s., 34, 7-10. Fishes and Medusae of the intermediate depths. A note on the work of the Michael Sars. Nature, 86, 483. Reports on the scientific results of the Expedition to the Eastern Tropical Pacific, in charge of Alexander Agassiz, by the U.S. Fish Commission Steamer Albatross, from October, 1904, to March, 1905, Lieut. Commander L. M. Garrett, Commanding. XXVI. The Ctenophores. Bull. Mus. Comp. ZooL, Harvard Coll., 54 (12), 369^04, 2 pis. Scientific results of the Philippine cruise of the Fisheries Steamer Albatross, 1907-1910. 22. 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Medusae of the Templeton Crocker and Eastern Pacific " Zaca " Expeditions. 19.^6 1938. Zoolovica. S.Y. Zool. Soc. 25 (3). 281 321, text tigs 1 20. (1952). Thomas Barbour. Biof^n: Mem.. Nat. .had. Sci.. 21, \} 45. BroELow. H. B. and Edmondson, W. T. (1947), Wind waves at sea, breakers and surf. U.S. Nt;vy Hydro^r. Office Puh. 602, xii plus 177 pp.. 57 text figs.. 24 pis. (Also translated into Russian in 1951 by B. B. SHTOKCiANA.) BiGKLOW, H. B. and Farfante, I. P. (1948), Fishes of the Western North Atlantic. Chap. I. Lance- lets. Mem.. Sears Found. Mar. Res.. 1, 1-28. 3 text figs. BiGELow, H. B. and Isflin, C. (1927). Oceanographic reconnaissance o\' the northern sector of ihc Labrador current. Science. 65 ( 1 69 1 ), 55 1 552. BiciFLOW, H. B. and Lfslii , Maurinf ( 1930). Reconnaissance of the waters and plankton of Monterev Bay, July 1928. Bull. Mus. Comp. Zool.. Harvard Coll.. 70 (5). 429 481, 43 text tigs. BiGFLOW, H. B., LiLLiCK, LoFS C. and Sears. Mary (1940). Phytoplankton and planktonic protozoa of the offshore waters of the (iulf of Maine. I. Numerical distribution. Trans. Amer. ^/7. Soc, n.s., 31 (3). 149 191, 10 text tigs. ^ BiGELOW, H. B. and Schroedfr, W. C. (1927), Notes on northwest Atlantic sharks and skates. Bull. Mus. Comp. Zool.. Harvard Coll.. 68 (5). 239-251. BiGtLow, H. B. and Schrcfijfr. W. C. ( 1929), A rare Braniid fish (Taractes princeps Johnson) in the northwestern Atlantic. Bull. Mus. Comp. Zool., Harvard Coll., 69 (2), 41-50. ! pi. BiGELOW. H. B. and Schrofdfr, W. C. (19.34), Canadian Atlantic Fauna. 12. Chordata. I2d. Marsipobranchii (Lampreys). 12e. Elasmobranchii (Sharks and rays). 121". Holocephali (Chimaeroids). Univ. Toronto Press for Biol. Bd., Canada. 38 pp., 35 text figs. ,\\ BiGtLOW, H. B. and ScHRotDbR, W. C. ( 1935), Two rare fishes, Notacanthus phas^anorus Goode and Lvcichthvs latifrons (Steenstrup and Hallgrimsson), from the Nova Scotian banks. Proc. Boston Soc. Nat. Hist.. 41 (2). 13-18, PI. 3. BiGELOW, H. B. and Schroeder, W. C. (1936), Supplemental notes on fishes of the Gulf of Maine. Bull. U.S. Bur. Fish., 48 {Bull No. 20), 319-343. BiGELOW, H. B. and Schroeder, W. C. (1937), A record of Centrolophus ni^er (Gmelin) from the western Atlantic. Copeia. 1937 (1). 61. BiGELOW, H. B. and Schroeder, W. C. (1939), Notes on the fauna above mud bottoms in deep water in the Gulf of Maine. Biol. Bull., 46 (3). 305-324, 8 text figs. BiGELOW, H. B. and Schroeder, W. C. (1940), Notes on New England fishes — Carcharodon carcharias (Linnaci:,)- Copeia, 1940(2), 139. BiGELOW, H. B. and Schroeder, W. C. (1940), Some deep sea fishes from the North Atlantic. Copeia, 1940(4), 231-238. BiGELOW, H. B. and Schroeder, W. C. (1940), Sharks of the genus Mustelus in the western Atlantic. Proc. Bo.ston Soc. Nat. Hist., 41 (8), 417^38, Pis. 14-19. BiGELOW, H. B. and Schroeder, W. C. (1941), Cephalurus, a new genus of Scyliorhinid shark with redescription of the genotype Catulus cephalus Gilbert. 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A new genus and species of Acanthobatid skate from the Gulf of Mexico. J. Washington Acad. Sci., 41 (3). 1 10-113, 1 text fig. BiGELOW. H. B. and Schroeder. W. C. (1951), Three new skates and a new chimaerid fish from the Gulf of Mexico. J. Washington Acad. Sci.. 41 (12), 383-392. 4 text figs. BiGELOW, H. B. and Schroeder, W. C. (1952), A new species of the cyclostome genus Paramyxine from the Gulf of Mexico. Breviora. No. 8, 10 pp., 6 text figs. BiGELOW, H. B. and Schroeder, W. C. (1953), Fishes of the Gulf of Maine. First Revision. Fi.sh and Wildlife Service, Fish. Bull.. 53 (F/.s/). Bidl. 74), 1-577, 288 text figs. BiGELOW, H. B. and Schroeder, W. C. (1953), Fishes of the Western North Atlantic. Ch. 1. Saw- fishes, Guitarfishes, Skates and Rays. Ch. 2. Chimaeroids. Mem., Sears Found. Mar. Res., 1 (2). 588 pp.. 127 text figs. BiGELOW, H. B. and Schroeder, W. C. ( 1954), Deep water elasmobranchs and chimaeroids from the northwestern Atlantic slope. Bull. Mus. Cotnp. Zool., Harvard Coll., 1 12 (2), 37-87, 7 text figs. BiGELOW, H. B. and Schroeder. W. C. (1954). A new family, a new genus and two new species of Batoid fishes from the Gulf of Mexico. Breviora. No. 24. 1 6 pp., 4 text figs. BiGELOW, H. B. and Schroeder. W. C. ( 1955). Occurrence off the Middle and North Atlantic United States of the offshore hake Merluccius alhidus (Mitchill) 1818. and of the blue whiting Gadus {Micromesistius) Poutas.sou (Risso) 1826. Bull. Mus. Comp. Zool.. Harvard Coll., 113 (2). 205-226. 3 text figs. BiGELOW. H. B.. Schroeder. W. C. and Springer. Stewart (1943), A new species of Carcharinus from the western Atlantic. Proc. New England Zool. Club. 11, 69-74. BiGELOW. H. B., Schroeder, W. C. and Springer. Stewart (1953). New and little known sharks from the Atlantic and from the Gulf of Mexico. Bull. Mus. Comp. Zool.. Harvard Coll., 109(3), 213 276, 10 text figs. BiGELOW. H. B. and Sears, Mary (1935), Studies of the waters of the continental shelf. Cape Cod to Chesapeake Bay. II. Salinity. Pap. Phys. Oceanogr. Meteorol., 4(1), 1-94, 55 text figs. BiGELOW, H. B. and Sears, Mary (1937), H 2. Siphonophorae. Rept. Danish Oceanogr. Exped., 1908-10, to the Mediterranean and Adjacent Seas, 2 (Biol.), 144 pp., 83 text figs. BiGELOW, H. B. and Sears, Mary (1939), Studies of the waters of the continental shelf. Cape Cod to Cape Hatteras. HI. A volumetric study of the zooplankton. Mem. Mus. Comp. Zool., Harvard Coll., 54 (4), 183-378, 42 text figs. BiGELOW, H. B. and Welsh, W. W. (1925), Fishes of the Gulf of Maine. Bull. U.S. Bur. Fish., 40 ( 1 ), 1-567, 278 text figs. Barbour, Thomas and Bigelow, H. B. (1944), A new giant Ceriatid fish. Proc. New England Zool. C/m6.23, 9-15, Pis. 4-6. Papers in Marine Biology and Oceanography. Suppl. to vol. 3 of Deep-Sea Research, pp. 1-6. Effect of trawling on animals of the sea bed By Michael Graham Fisheries Laboratory, Lowestoft, England Summary — Damage to fish food species trawled over in the main area of the North Sea plaice, cannot be serious; otherwise there would be a noticeable diflFcrence where trawling is impossible, as close to light vessels or among the under-water sand dunes. Direct attack, covering the ground some five or six times over on the average, did break full-grown Heart Urchins, Echinocardium cordatum, and possibly swimming or paddlcr crabs (Portunus dcpuraior), but appeared not to damage Ophiura albida, nor any of the fragile-shelled plaice food animals: razor shells, Mactra or Tellina. Those forms were not very abundant, but all the 15 specimens taken of fragile animals (other than urchins and paddlers) were undamaged. Such large urchins as were damaged were not plaice food. Doubtless Sabellaria habitations (ross) would be broken and laid low, but they would probably soon be reconstructed. Trawling, even with a tickler chain, seems again to escape the so viable indictment. INTRODUCTION Many trawlermen are sure that their work alters the bed of the sea. Some say that it improves it, increasing the growth of animal forms growing up from the sea bed by clearing out old structures. Others say that trawling, especially with tickler chains, harms the food of fishes by breaking protective shells and structures. The complaint of damage is an old one. In 1376 the Commons petitioned the King of England " that the great and long iron of the wondyrchoun runs so heavily and hardly over the ground when fishing that it destroys the flowers of the land below water there ". In the 19th century the beam trawl came in for similar criticism, which Dr. W, C. Macintosh, acting for the Royal Commission of 1863, disposed of. However, trawls have become heavier and tickler chains more common, in certain fisheries almost universal. One such fishery is the English one for plaice in the southern North Sea; and in 1938 it seemed worth while to devote a little research- vessel time to finding out whether tickler chains had any marked effect. Doubtless trawling with heavy tickler chains breaks up and flattens structures made of sand-tubes by Sabellaria, and similar comparatively fragile highly projecting structures, when these are not so strong as to prevent trawling because of the frequency of tearing. But Sabellaria tenements seem to be annual growths, run up fairly quickly, so that it may not be assumed that much permanent damage is done. The fishing skipper of the George Bligh advised that it would be foolish and not really relevant to experiment on Sabellaria grounds, which he thought to be limited compared with the whole plaice area; so the decision was made to confine the enquiry to the usual clean, sandy ground frequented by plaice. METHODS The first line of approach was to assume that there exists a small sanctuary about three-quarters of a mile in radius centred on each light vessel, within which trawling would be very rare, for fear of fouling the lightship's moorings. If trawling had made B 1 2 Michael Graham an appreciable effect on the benthos of the plaice area, the community within that sanctuary should be appreciably different from the community outside. The sanctuaries of two light vessels, the Haaks and the Terschelling, were examined with the following gear : Petersen's grab ; Naturalist's dredge ; Agassiz trawl ; otter trawl with shrimp- Table I. — Dredging before and after trawling BEFORE George Bligh, Cruise K (1938), Sta. 16, 10th July, 1938, Naturalist's dredge (6 foot) lined shrimp-netting, Egmont SE'ly 15 miles, Dhan on 90 fms. wire in Lat. 52° 44i' N. Long. 4° 16' E., 13 fathoms, sand, 1015 to 2010 hours, tides various, sea 4, wind SW'ly 4, sky cloudy, 14 hauls driving with wind and tide except where otherwise noted. Dhan bearing and distance N to NxW \ mile (magnetic variation 9° W) S to SJE \ mile S 1} miles E to SE^E * mile W to SWxW * mile NE 200 yds. to \ mile (? direction reversed) NE X E to SxWiW i mile NExE i mile to ESE i mile NExE i mile to 200 yards NExE i mile '* tide getting less " NExE i mile to ENE i mile NExE to SW i mile NExE to SW \ mile " with help of engines " buoy to NE'ward no distance AFTER 42 traverses with the trawl as nearly as possible to the tracks of Sta. 16, Hauls 7-13; Cruise K 1938, Sta. 20, 13th July, 1938, otherwise as for Sta. 16 but time 1415 to 1920 hours, tide " 1400, BuflFs and Dhan NNW, so pro- ceeded to buoy ", sea 2, wind variable 2, sky cloudy, 7 hauls driving with wind and tide. Haul Time Dhan bearing and distance (assumed 10 mins. duration) 1 1418-1428 NExN to ESE i mile 2 1447-1505 NNE to Simile 3 1524-1538 NExE to S i mile 4 1599-1612 NExE to SiE i mile 6 no record — " foul haul " 6 1 704- 1712 ENE 300 yards to SSW 200 yards 7 1730-1745 NExE to SxE i mile 8 1810-1826 ENE i mile to SSE 200 yards 9 1 842-1900 " ? washed it all out " net cover on the codend. Control hauls were made 5 miles away from the Haaks only. That work was done on the 6th, 7th, 8th and 9th of July, 1938. {George Bligh K 1938, Stas. 2-10, 11-12, 13-14). Another sanctuary is provided by the under-water sand dunes of the extreme south-eastern North Sea. These are dangerous to otter trawls, because of the trawl doors getting down into the small valleys between the ridges and the net then fouling iaul Time (assumed 10 mins. duration) 1 1038-1048 2 1110-1120 3 1145-1155 4 1235-1245 5 1314-1324 6 1356-1406 7 1440-1450 8 1519-1529 9 1551-1601 10 1623-1633 11 1654-1704 12 1725-1755 13 10 minutes 14 10 minutes Effect of trawling on animals of the sea bed 3 the intervening crest. The beam trawls formerly used could generally get along safely, presumably because the beam took the blow of the sand-crest. By 1939, beam trawls were almost extinct, so the " bank " area provided another sanctuary. Had trawling had an appreciable effect, the benthos should be different as one left the bank area. On July 23rd and 24th, 1939, this was investigated near the northern boundary, as located by echo-sounding, using Petersen's grab and the Naturalists' dredge. Two lines of stations were completed {George B/igh L 1939, Sta. 14, Hauls 0-9, Sta. 15, Hauls 0-6). Another line of approach was to try to damage animals of the sea-bed with a trawl armed with a heavy tickler-chain. The idea was to choose a bed of some animal with a fragile shell, as fragile as possible, and then tow over it as precisely as possible. Fig. I. Dredge hauls before and after trawling Tracks of hauls before using trawl and tickler chain are shown as whole arrows with the serial number in a circle; hauls after, as broken arrows with unringed serial numbers, related to the position of a buoy's anchor. " Before " hauls 7 to 10 showed a bed of Heart Urchins, which was also sampled by " after " hauls 2 to 8 with moderate success, the wind having changed. The swinging positions of the buoy have been taken into account. Its area of 180 yards radius limited trawling, probably to something like the shaped area indicated. In the waist, the area would be trawled over on, the average, about 5 times. Heart Urchins were broken, but not Mactra. Range judged visually was perhaps accurate to within 20 per cent; but the buoy's position (on 90 fms. wire in 13 fms.) would alter with the tide. A trawl, which probably fished at something over 20 yards width, was furnished with half-inch chain extending for 100 feet between the otter-boards, shorter and therefore mostly ahead of the 120 foot ground-rope. This was towed up and down on the 11th, 12th, and 13th July, 1938, crossing the ground 42 times; but owing to the error in the buoy's position due to tide and the imprecision of trawling, these cannot be thought of as 42 exactly super- imposed hauls 20 yards wide. Instead we must think of an area like that shown in Fig. 1, representing fishing as close to the buoy's moorings as the ship dare go. The waist of the constricted area is 1 50 yards wide, which allows for 7 to 8 strips side by 4 Michael Graham side; so, on the average, the ground would be swept about 5-6 times. It was possible to be more precise about the examination of the benthos bed before and after the trawling, by choosing the same phase of the tide. The animal tested was Echinocardium cordatum, the Heart Urchin, which was the most vulnerable animal we encountered — certainly not a plaice food at the size (from memory, about an inch and one-half in diameter), but no better animal was found. It had the further disadvantage that dredge catches would fall from 39 in a 10 minute haul to 3 when the tide slackened. This leads to an investigation of where exactly the dredge hauls lay (Table I). The dredging ground (52° 44|' N., 4° 16' E.) was some 13 miles southerly of the Haaks Light Vessel, and because many of the hauls were made while the George Bligh was driving with wind and tide it is possible to plot the approximate tidal pattern (Sta. 16, Hauls 1-13, but discarding No. 6, for which the record is suspect). Assuming as a first approximation that the Dhan buoy did not swing during a 10 minute dredge haul, the course of the haul can be plotted out from the bearing and distance at the beginning and end of the haul, taking an arbitrary position for the buoy. Then, as a second approximation, the buoy is assumed to be on that bearing and at a distance of 180 yards (90 fms of wire in 13 fms of depth) from its anchor. The position of dredge haul is then replotted from the assumed true position of the buoy. Most positions of the buoy were found to lie near a N.E. and S.W. zone (magnetic), not very different from that derived from tidal information on admiralty chart 2 182 A for a position 52° 36' N., 4° 10' E. The information given in Table I has been plotted on Fig, 1, from which it is seen that there is reasonable overlap of Sta. 16, Hauls 7-13 with Sta. 20, Hauls 2-8. Neither set lay perfectly in the trawlable area; but both lay partly within it. Sta. 16, Hauls 7-13, and Sta. 20, Hauls 1-8, may be said to do so. Some benthos material was preserved from these investigations ; but none is to be found 17 years later. It is thought that the jars were buried for safety of staff in 1940, and discarded as not in good order in 1945. Data are therefore in the form of incom- plete identifications written in the log-books at the time. They are so given here= During the attempt to break Echinocardium it was found possible to distinguish whole, broken or even fragmentary specimens, as " dead ", meaning " long dead " (having no spines), " recently dead " (having lost some spines), and " broken " (having all the proper spines). By counting fragments containing apical pores, it was possible to estimate the number of whole urchins. RESULTS The sanctuaries from trawling showed no marked difference in benthos from outside, so there seems no reason to burden this paper with details of the results. The only point of interest was a bed of Echinocardium within three-quarters of a mile of the Haaks Light Vessel, but because this did not extend around the Light Vessel, we can hardly attribute significance to it, especially because later on we found a localized bed 13 miles away in the fishing area. The fauna also included " flat stars " (Astropecten), Asterias rubens, Portunus depurator, Eupagurus, and a few Mactra, Venus, and razor-shells. These showed no marked differences three-quarters of a mile and 5 miles from the Haaks Light Vessel, and the species and quantities three- quarters of a mile from the Terschelling Light Vessel seemed similar to those close to the Haaks. Effect of trawling on animals of the sea bed 5 Similarly, there was no appreciable change in fauna taken in grab and dredge on stations 3 miles apart running out of the protected " banks " area and in to it again. The fauna was similar to that noted above. On the other approach — the direct attack with the tickler chain and trawl — the results are shown in Table II. It will be remembered that Hauls 7-13 of the first Table II. — Dredi^e catches before and after traw/ing Heart Urchins * Paddlerst Other fragile forms Live Recently Dead Dead Live Recently Dead Dead Mactra, Razor-shells. Tel- lina. or Ophiura alhida. ) unbroken exccp* where _o JD c ' c % % % % Ca (C) PHYLUM PORIFERA CLASS CALCAREA ORDER HOMOCOELA Leucosolenia eleanor Calif. 1-29 trace 3-24 20-2 — Leucosolenia sp. Maine 13-6 099 23-3 25 ; 3-33 Sponge (unidentified) Beaufort Sea 1 3 -5 0098 34-4 22-8 3-34 ORDER HETEROCOELA Leuconia heathi Calif. 100 trace 2-28 30-3 — Rhabdodermella nuttingi Calif. 29-6 0149 35-8 12-7 2-30 CLASS DEMOSPONGIAE SUBCLASS TETRAXONIDA, ORDER EPIPOLASIDA Tethya aurantia Calif. 06 trace 6-30 42-3 — SUBCLASS CORNACUSPONGIDA ORDER POECILOSCLERINA Esperiopsis originalis Calif. 08 trace 2-12 38-2 — Ophlitaspogia pennata Calif. 005 trace 1-97 40-8 — ORDER HAPLOSCLERINA Haliclona permolUs Calif. 07 trace 6-57 42-2 — ORDER KERATOSA Euspongia sp. Unknown 005 trace 12-6 85-0 — (D) PHYLUM COELENTERATA* CLASS HYDROZOA, ORDER MILLEPORINA Millepora tenera Ifalik Atoll 381 0-686 41-4 3-15 8-22 ORDER STYLAS' ERINA Allopora calif ornica Calif. 36-5 0-778 38-5 8-46 9-74 A. campyleca paragea Gulf of Alaska 35-7 0-768 40-6 4-47 9-83 A. porphyra Calif. 38-8 0-580 41-1 2-91 6-83 A. venusta Calif. 370 0-808 40-4 5-38 9-97 Cryptohelia trophostega Bering Sea 37-8 0-930 411 3-42 11-2 Distichopora violacea Marshall Is. 37-6 0-640 41-2 3-54 7-77 Errinopora zarhyncha Aleutian Is. 37-5 0-224 43-1 2-50 2-73 Stylaster elegans Marshall Is. 37-4 0-852 41-3 3-84 10-4 S. sanguineus Hawaii 37-7 0-843 41-2 3-88 10-2 CLASS ANTHOZOA, SUBCLASS AL CYONAR lA ORDER STOLONIFERA Tubipora sp. Unknown 311 0-225 41-8 2-38 3-30 ORDER ALCYNACEA Eunephthya rubiformis Beaufort Sea 4-72 0-039 12-4 30-4 3-78 ORDER PENNATULACEA Stylatula elongata Calif. 22-7 0-128 28-5 33 2-64 ORDER GORGO NACEA Psammogorgia arbuscuLa : Calif. 21-4 0-136 30-0 37-5 2-90 ORDER COENOTHECALIA Heliopora coerulea I Ifalik Atoll 37-3 0-618 40 660 7-57 * Some of the specimens were contributed by the Hopkins Marine Station from the collection of the late Professor W. K. Fisher. The strontium-calcium atom ratio in carbonate-secreting marine organisms 31 Table I (cont.) Locality Carbon Calcium Strontium dioxide o / 0/ /o / /cj Organic matter /o Atom ratio ^ y 1000 Ca SUBCLASS ZOANTHARIA, ORDER MADREPORARIA Acropora sp. Astrangia sp. Balanophyllia elegans Caryophyllia sp. Meandrina sinuosa PociUopora sp. Porites sp. Ifalik Atoll Calif. Calif. Calif. Bermuda Ifalik Atoll Ifalik Atoll 37-3 370 32-3 37-4 36-5 37-9 37-3 (E) PHYLUM ANNELIDA, CLASS POLYCHAETA FAMILY CIRRATULIDAE Dodecaceria pacifica B.C., Canada* 32-5 D.fistulicola Calif. 35 4 FAMILY SERPULIDAE Salmacina tribranchiata \ Calif. Serpula vermicu/aris B.C., Canada S. vermicularis Calif. Spirorbis sp. N. H. 33-2 32-4 321 29-2 0-810 0-868 0-666 0-850 0-802 0-735 0-790 0-574 0-530 0-596 0-288 0-289 0-247 41 41 37 41 41 41 40 40-6 40-3 36-5 40-7 42-5 34-2 3-85 3-43 8-84 2-90 3-25 3-13 5-38 5-38 5-20 6-35 4-51 4-60 15-0 ORDER THORACICA Floridaf N. H. N. H. Wash. Beaufort Sea Florida Florida Wash. Calif. Calif. Wash. Wash. B.C., Canada Calif. N. C. Calif. Calif. 370 37-6 34-1 39-1 36-1 36-7 36-6 37-0 37-4 35-6 34-2 37-4 36-1 37-2 32-4 36-8 35-6 0-428 0-382 0-336 0-324 0-337 0-381 0-345 0-358 0-371 0-334 0-282 0-364 0-337 0-38! 0-328 0-344 0-381 41 41 39 42 41 41 41 41 41 39 36 41 41 41 37 40 41 5 3 7 1 3 4 2 3 2 7 12 2 3 1 8 4 4 •00 •27 -76 -80 -24 -80 -39 -12 -88 -18 -3 -72 -80 -88 -33 -48 -07 (F) PHYLUM ARTHROPODA, SUBPHYLUM MANDIBULATA, SUPERCLASS CRUSTACEA CLASS CIRRIPEDIA, Balanus amphitrite B. balanoides B. balanoides B. cariosus B. crenatus B. eburneus B. eburneus B. glandula B. glandula B. nubilis B. nubilis Balanus sp. Balanus sp. B. tintinnabulum Chthamalus fragilis Mitella polymerus Tetraclita squamosa 9-92 10-7 9-41 10-4 10-1 8-85 9-67 8-07 6-84 8-24 4-06 4-12 3-86 5-28 4-64 CLASS MALACOSTRACA, SUBCLASS EUCARIDA, ORDER Cancer antennarius C. antennariust C. borealis C. magister C. productus C. productus ** Hemigrapsus nudus Pugettia producta Calif. Calif. N. H. Wash. Calif. Calif. Wash. Calif. (G) PHYLUM MOLLUSCA CLASS AMPHINEURA, ORDER PO 27-2 28-0 24 24-0 23-3 0-44 26-0 21-2 0-363 0-371 0-322 0-351 0-308 trace 0-349 0-278 30-2 32-5 29 3 281 30-2 8-6 29-3 25-7 I DECAPODA 32-7 20 27-2 28-1 30-0 85-5 28-7 33-3 LYPLACOPHORA Cryptochiton stelleri C. stelleri Cyanoplax hartwegii Nuttallina californica Tonicella lineata Calif. Wash. Calif. Calif. Calif. 38-2 37-1 38-3 38-2 37-6 0-736 0-751 0-716 0-662 0-640 40-3 41-4 41-3 41 2 41 4 6-10 3-66 5-70 4-69 4-83 50 78 27 74 32 42 53 29 3-77 4-45 4-27 4-67 4-62 4-27 4-88 612 6 05 613 669 604 6-13 600 8-80 9-25 8-55 7-91 7-79 * The specimens from British Columbia were provided by Mr. Cyril Berkeley of the Pacihc Biological Station. /^ c v t The Florida specimens were provided by Mr. C. b. Yent.sch. t Claw ** Moulting 32 Thomas G. Thompson and Tsaihwa J. Chow Table I (cont.) Carbon Organic ' Atom ratio Calcium Strontium dioxide matter Sr 7o /o Ca X 1000 (G) PHYLUM MOLLUSCA (continued) CLASS AMPHINEURA, ORDER POLYPLACOPHORA Ischnochiton heathiana I. mertensii Ischnochiton sp. Katharina tunicata Mopah'a ciliata M. lignosa M.muscosa M. muscosa M. wosnessenskii Chiton (unidentified) Calif. Calif. Wash. CaUf. Calif. Calif. Calif. Wash. Calif. Beaufort Sea 38-7 38-6 38-8 38-7 37-2 37-9 37-9 37-8 37-6 37-0 (H) PHYLUM MOLLUSCA, CLASS PELECYPODA ORDER FILIBRANCHIA FAMILY ANOMIIDAE Pododesmus macroschisma Calif. FAMILY MYTILIDAE Botula falcata Modiolus capax M. modiolus M. modiolus M. modiolus M. modiolus Modiolus sp. Mytilus edulis M. edulis M. edulis M. edulis M. edulis M. edulis M. edulis M. edulis M. californianus M. californianus FAMILY Pecten hindsii P. hericius Calif. Calif. N. H. N. H. N. H. N. H. Wash. N. H. N. H. Maine Maine Wash. B.C., Canada Beaufort Sea Calif. Calif. Calif. PECTINIDAE Wash. Wash. ORDER EULAMELLIBRANCHIA FAMILY OSTREIDAE Crassostrea virginica C. virginica Ostrea gigas O. gigas O. lurida N. H. N. C. B.C., Canada B.C., Canada Calif. FAMILY SPONDYLIDAE Spondylus sp. I unknown FAMILY CHAMIDAE Chama pellucida Calif. FAMILY TELLINIDAE Macoma irus Calif. M. nasuta Calif. M. secta Calif. Tellina sp. B.C., Canada 38-6 38-6 37-5 37-8 38-8 37-5 37-8 38 37 37 38 37 38 38 36 36 390 38-5 38-2 37-8 33-7 37-8 34-6 36-2 38-6 38 36-8 0-636 0-636 0-680 0-687 0-626 0-664 0-608 0-709 0-606 0-680 0-103 0196 0-103 0125 0-116 0-112 0-132 0-099 0-176 110 128 116 106 154 117 118 0-086 0086 0111 0107 092 0-107 0-097 0100 0-085 0-128 0-142 •4 •3 •8 ■6 -1 -2 41' 41 41- 39- 41' 41- 41 42-2 41-4 41-5 42-6 38-9 41-1 41-8 42-7 41-3 4M 42-1 42-5 42-0 42-1 42-4 42-2 41-4 41-2 42-2 43-1 43-1 41-8 42-1 42-4 41-8 37-6 42-5 42-5 41-6 42-3 3-63 3-95 3-06 3-91 •83 -26 -74 -31 4- 4- 4- 3' 3-88 4 00 1-75 •67 10 ■97 39 •58 20 96 10 32 30 52 24 42 53 87 54 70 1-91 1-75 216 2-34 13-3 1-71 1-68 1-67 2-47 38-7 0153 40-8 2-42 1-81 38-0 .0-215 38-9 2-27 2-59 38-9 0-248 42-5 2-17 2-91 37-1 0173 42-1 2-50 214 7-52 7-55 8-02 8-12 7-69 801 7-32 8-57 7-37 8-41 1-22 2-32 1-25 1-52 1-37 1-37 1-60 119 216 1-33 1-53 1-43 1-26 1-82 1-46 1-48 101 102 1-33 1-29 1-25 1-29 1-28 1-26 101 1-54 1-76 The strontium-calcium atom ratio in carbonate-secreting marine organisms 33 Table I (conf.) Carbon Organic Atom ratio Locality Calcium Strontium dioxide matter ^~ ' 1000 Vo o - o o / o o/ Co FAMILY SOLENIDAE Ensis directiis N. H. 370 01 64 41-9 3-47 2 03 Siliqua costata N. H. 38-4 0182 42-4 2-46 217 S. patula Wash. 38-4 0-207 42-0 2-88 2-46 Solen sicarius CaHf. 38-1 0116 42-9 1-66 1-39 FAMILY MACTRIDAE Mesodesma deauratum N. H. 37-6 0164 42-0 3-31 2 00 Schizothaerus nuttallii Wash. 37-8 0195 41-6 4-00 2-36 S. nuttallii Calif. 391 0177 42-6 1-79 207 Spisida solidissima N. H. 38-2 0-216 42-0 2-58 2-58 FAMILY PLEUROPHORIDAE Cyprina islandica \ N. H. FAMILY CLINOCARDIIDAE Clinocardium nuttallii C. nuttallii C. nuttallii Wash. Wash. Calif. FAMILY VENERIDAE Compsomyax subdiaphcma Irus lamellifer Protothaca staminea P. staminea P. tenerrima Saxidomus giganteus S. muttallii Tivela stultorum Venus mercenaria Wash. Calif. Wash. Calif. Wash. Wash. Calif. Calif. Canadian Atlantic FAMILY Per ip lama sp. PERIPLOMATIDAE I B.C., Canada FAMILY LYONSIIDAE Mvtilimeria nuttallii Calif. FAMILY MYIDAE Mya arenaria M. arenaria M. arenaria M. arenaria Maine Wash. Oregon Wash. FAMILY Panope generosa Saxicava sp. SAXICAVIDAE Wash. Beaufort Sea FAMILY PHOLADIDAE Pholadidea penita Calif. P. ovoidea Calif. Zirfaea crispata Z. pilsbryi N. H. Calif. 39-2 37-6 38-5 38-0 38-6 37-6 39-4 38-2 37-3 38-6 37-8 38-8 38-6 37-9 36-6 38-6 38-7 38-8 38-3 38-2 37-9 37-0 38-7 38-3 39-0 0135 0-245 0-188 0-185 0-180 0134 0-179 0-149 0-175 0-192 0-125 0-120 0148 0-183 0-188 0-246 0-238 0-181 0181 0-207 0-196 0-147 0-192 0-148 0-230 42-7 41-9 41-8 41-8 42-2 41-6 41-8 42-0 42-0 42-3 42-2 42-0 42-5 42-0 39-8 42-2 42-3 42-3 42-2 42-3 40-5 41-2 42-2 4I-2 42-5 (I) PHYLUM MOLLUSCA, CLASS GASTROPODA SUBCLASS PROSOBRANCHIA ORDER ASPIDOBRANCHIA, FAMILY FISSURELLIDAE Diodora aspera i Calif. Fissurella volcano j Calif. Megathura crenulata ' Calif. Megatebennus bimaculatus \ Calif. C SUBORDER ZYGOBRANCHIA 38-5 39-5 38-0 390 0-114 0-108 0-104 0-117 41-5 42-6 42-8 42-1 1-93 2-98 3-68 3-55 07 20 30 45 94 23 63 ■34 2-38 2-60 7-63 2-44 2-28 2-48 2-22 2 03 4-98 3-63 2-36 3-63 2-30 3-54 1-86 2-03 2-74 1-57 2-98 2-23 2-22 2-13 1-63 2-08 1-78 214 2-27 1-52 1-41 1-75 2 20 2-35 2-91 2-81 2-12 216 2-48 2-36 1-82 2-27 1-77 2-70 1-35 1-25 1-25 1-38 34 Thomas. G. Thompson and Tsaihwa J. Chow Table I (cont.) Calcium Strontium Carbon dioxide Organic matter % Atom ratio Sr Ca 1000 FAMILY HALIOTIDAE Haliotis cracherodii Calif. H. rufescens Calif. SUBORDER PATEL .lace; FAMILY ACMAEIDAE Acmaea digitalis Calif. A. digitalis Calif. A. insessa Calif. A. limatula Calif. A. mitra Calif. A. mitra Wash. A. pelt a Calif. A. persona Calif. A. scabra Calif. A. t. scutum N. H. A. t. scutum Calif. Lottia gigantea Calif. SUBORDER TROCHACEA FAMILY TROCHIDAE Calliostoma canaliculatum ' Calif. C. costatum Calif. C. gloriosum Tegula brunnea T. funebralis T. montereyi Wash. Calif. Calif. Calif. FAMILY TURBINIDAE Astraea inaequalis I Calif. ORDER PECTINIBRANCHIA SUBORDER TAENIOGLOSSA FAMILY EPITONIIDAE Epitonium groenlandicum \ Atlantic Epitonium sp. | Calif. FAMILY VERMETIDAE Petaloconchus montereyensis i CaUf. FAMILY LITTORINIDAE Littorina litorea Maine L. litorea N. H. L. palliata ( = L. obtusata) N. H. L. planaxis Calif. L. rudis (= L. saxatilis) N. H. L. rudis N. H. L. scutulata Calif. FAMILY CALYPTRAEIDAE Crepidula adunca ' Calif. C.fornicata | Mass. C. nummaria ] Calif. FAMILY Polinices Heros P. draconis Natica sp. NATICIDAE Mass. ' Wash. Beaufort Sea SUBORDER STENOGLOSSA FAMILY OLIVIDAE Oliva litterata \ Florida Olivella biplicata \ Calif. 37-1 38-8 38-5 37-6 37-8 38 37 38 37 37 37 38 37 37 37-2 370 36-3 37-5 37-4 38-2 36-7 370 36-8 35-6 38-4 38-7 37-2 37-8 38-6 37-5 38-4 37-7 38 38-5 37-7 38-8 37-8 39-6 38-9 0123 0130 0199 0-204 0183 01 68 0161 01 62 0151 0-201 0-180 0180 01 64 0-170 0-129 0-122 0-131 0-124 0-113 0-121 0-120 0-118 0-120 0144 0107 0107 0-142 0-155 0-140 0-125 0-121 0140 0-149 0-153 0-121 0-137 0-108 0145 0113 42-2 41-7 41-0 41-2 41-7 40-6 41-9 41-5 41 41-9 41-2 40-2 42-5 42-8 41-0 42-4 41-8 41-7 42-5 42 42-3 42-3 42-2 42-5 41-8 42-8 42-4 3-87 3-82 42-6 2-51 42-1 2-79 42-4 2-84 42-0 3 03 42-4 2-61 42-3 2-48 40-4 3-39 42-0 3-16 42-5 1-97 41-8 3-28 41-7 3-65 42-2 2-71 4-95 2-96 4-73 4-06 3-64 3-99 5-37 3 07 4-80 710 27 84 72 62 50 39 77 3-22 2-37 2-36 2-96 2-28 3-54 1-61 1-84 1-52 1-54 2-37 2-48 2-21 201 1-98 1-94 1-84 2-45 2-21 2-14 2 00 2-08 1-59 1-51 -65 -51 •38 1-45 1-50 1-46 1-49 1-85 1-27 1-26 74 87 66 53 1-44 1-70 1-79 1-82 1-47 1-61 1-31 1-67 1-33 The strontium-calcium atom ratio in carbonate-secreting marine organisms Table I (cont.) 35 Locality Calcium Strontium % % Carbon dioxide /a Orf^anic matter % Atom ratio Sr Ca ^ 1000 FAMILY FUSINIDAE Fusinus monksae i Calif. FAMILY NASSARIIDAE Nassa obsoleta ' N. H. FAMILY MURICIDAE Acanthina spirata Calif. Murex pume Florida M. triolatus Calif. Thais canaliculata Calif. T. emarginata Calif. T. lamellosa Wash. T. lapillus N. H. T. lapillus N. H. 35-5 36-4 38-6 39-9 380 38-1 37-6 38-8 38-5 36-5 ons 0151 0131 0172 0131 0128 0138 0129 0126 0128 40- 1 40-9 42-3 41-7 42-6 42-8 42-3 42-8 42-7 39-4 4-27 6-46 2-66 2-30 1-68 2-26 3 03 1-45 113 3-68 1-48 1-89 55 97 58 54 68 1-52 1-50 1-60 SUBCLASS OPISTHOBRANCHIA, ORDER NUDIBRANCHIA Anisodoris nobilis Calif. Archidoris montereyensis I Calif. Triopha grandis I Calif. (J) PHYLUM MOLLUSCA CLASS SCAPHOPODA Dentalium entale i Wash. 2-60 316 0-25 38-2 063 062 trace 0196 3-25 4-68 67-2 78 -91-3- -) 42-3 1-81 11 9 2-35 CLASS CEPHALOPODA, SUBCLASS DIBRANCHIATA, ORDER DECAPODA Loligo opalescens Calif. I trace trace trace 99 5 Sepia sp. unknown 35-8 (K) PHYLUM BRYOZOA, CLASS ECTOPROCTA ORDER CYCLOSTOMATA Idmonea sp. Crista sp. Bryozoa (unidentified) Bryozoa (unidentified) Bryozoa (unidentified) f Calif Calif. Beaufort Sea Beaufort Sea Beaufort Sea ORDER CHEILOSTOMATA Bugula californica , Calif. Hippodiplosia insculpta Phidolopora pacifica Calif. Calif. 35-6 35-7 34-3 30-6 25-3 18-2 29-4 33-7 0-293 0-307 0-282 0-256 0-209 0181 0-124 0-247 0-221 401 38-1 38-2 40-2 35 31-8 23-7 35-8 400 7-35 7-90 6-28 7-30 16-5 23-2 36-0 13-8 6-37 3-74 3-94 3-61 3-41 3-12 3-27 312 3-84 3 00 (L) PHYLUM BRACHIOPODA, CLASS ARTICULATA, ORDER TESTICARDINES Hemithyris psittacea Beaufort Sea 37-7 0113 42-2 3 02 1 H. psittacea* Beaufort Sea Terebratalia transversa Wash. Terebratulina unguicala Calif. Brachiopoda (unidentified) Calif. 37-7 38-9 38 37-8 38-7 0113 0102 0130 0113 0108 42-2 42-8 41-3 42-5 42-7 3 02 1-94 2 02 1-8! 1-79 1-37 I 20 1-57 1-37 1-28 (M) PHYLUM ECHINODERMATA CLASS CRINOIDEA I Antedon sp. ! Calif. i CLASS ASTEROIDEA ORDER FORCIPULATA 25-8 0-145 38- 13-3 2-56 Asterias forbesi A. vulgaris A. vulgaris Leptasterias aequalis L. pusilla Mediaster aequalis Mass. N. H. Mass. Calif. Calif. Calif. 18-6 0113 24-9 39-4 2-78 22-5 0141 27-9 33-6 2-86 20-2 0-128 28-6 34-4 2-89 20-9 0-127 32-7 31-0 2-78 25-5 0146 37-7 204 2-61 27-3 01 58 36-1 190 2-60 * Remains. 36 Thomas G. Thompson and Tsaihwa J. Chow Table I (cont.) Carbon Organic Atom ratio Locality Calcium Strontium dioxide matter — 1000 % % /o % Ca CLASS ASTEROIDEA (continued) ORDER FORCIPULATA Pisaster brevispinus Calif. 22-7 0-131 33-6 28-5 2-64 P. giganteus Calif. 17-2 0101 31-9 37-2 2-69 P. ochraceiis Calif. 24-6 0148 34-0 24-4 2-76 Pycnopodia helianthoides Calif. 200 0114 31-8 33-5 2-60 ORDER SPINULOSA Henricia leviuscula Wash. 23-1 0136 30-3 28-6 2-69 H. leviuscula Calif. 25-2 0-155 33-3 25-8 2-81 H. sanguinolenta N. H. 18-4 0112 28-6 38-2 2-78 Henricia sp. Beaufort Sea 20-6 0125 29-0 34-0 2-78 Solaster papposus Beaufort Sea 23-2 0-131 31-0 28-8 2-60 Patiria miniata Calif. 26-8 0162 36-1 21-2 2-77 ORDER PHANERO ZONIA Hippasteria spinosa Wash. 22-7 0-137 30-8 29-6 2-76 Luidia sp. Calif. 24-1 0-142 34-1 19-3 2-69 CLASS OPHIUROIDE A ORDER EURYALAE Gorgonocephalus sp. Beaufort Sea 26-6 0158 33-4 21-8 2-72 ORDER OPHIURA] ^ Amphipholis squamata ^ Calif. 24-7 0-146 33-2 23-5 2-70 Ophiopholis aculeata Maine 24-5 0146 32-6 24-4 2-72 O. aculeata N. H. 25-9 0-149 34-5 19-7 2-63 Ophiothrix spiculata Calif. 20-4 0-118 28-9 33-4 2-64 Ophioplocus esmarki Calif. 27-1 0-165 36-9 17-0 2-79 Ophiura sarsii Wash. 31-2 0185 39-2 11-0 2-71 0. sarsii Beaufort Sea 30-2 0-175 38-0 13-5 2-65 CLASS ECHINOIDE/i k. ORDER CENTRECHINOIDA Strongylocentrotus drobachiensis N. H. 34-8 0-214 41 5-68 2-81 S. drobachiensis Wash. 35-5 0-214 42 3-33 2-76 S. drobachiensis Wash. 35-6 0-210 42-0 3-05 2-70 S. fragilis Calif. 32-7 0-188 39-6 9-28 2-63 S. franciscanus Wash. 36-4 0-218 42-8 2-97 2-74 S. pallidus Beaufort Sea 32-2 0-191 40-0 810 2-71 S. purpuratus Wash. 35-4 0-215 43-8 2-18 2-78 S. purpuratus Calif. 33-2 0-210 41-4 6-96 2-89 Heterocentrotus trigonarius Ifalik Atoll 31-6 0180 43-9 1-20 2-60 ORDER CLYPEAST ROIDA Dendraster excentricus Calif. 33-5 0-205 43-5 2-85 2-79 Echinarachnius parma N. H. 31-8 0-178 41-4 5-62 2-56 ORDER SPATANGi DIDEA Brisaster sp. Wash. 30-5 0164 40-1 8-77 2-46 CLASS HOLOTHURC >IDEA ORDER DENDROCHIROTIDA Cucumaria curata Calif. 0-8 trace 7-5 84-0 — Psolus chitonoides Wash. 29-1 0-173 38-3 12-2 2-72 P. peroni Beaufort Sea 25-2 0-153 32-1 25-8 2-78 P. sp. Calif 28-6 0-170 34-5 18-0 2-72 (N) PHYLUM CHORDATA CLASS ASCII )IACEA Polyclinum planum \ Calif. 0-30 trace 13-8 70-0 — Synoicum par-fustis Calif. 0-38 trace 13-5 69-1 — The strontium-calcium atom ratio in carbonate-secreting marine organisms 37 Table II. — The occurrence of calcium and strontium in substances other than marine organisms Localilv Carbon Organic Atom ratio Calcium Strontium dioxide matter ' Sr Ca X 1000 Walrus (Odobenus rosmarus) ivory Alaska 21 trace 2-96 36-8 Fresh water clam (unidentified) Wash. 380 085 41-4 5-53 102 Fresh water clam (unidentified) Wash. 36-4 0072 41-3 6 02 0-90 Potamobius sp. Wash. 19-8 0077 21-7 45-5 1-78 Deep sea sediments* Indian Ocean (Swedish Deep Sea Expedi- 29-4 0125 37-7 2-95 1-94 tion 1948- 1949) Globigerina oozet Pacific Ocean 37-5 0122 41-8 1-46 1-49 Coquina rock (cemented shells) Florida 37-8 0153 41-7 1-57 1-85 Limestone deposits Wash. 38-1 0052 42-5 0-68 0-63 Strontianite deposits Wash. 3-56 52-5 31-5 0-26 6,750 Sea water Over-all 8-90 * Specimen was provided by Mr. Taivo Laevastu. t Specimen was provided by Dr. Howard R. Gould. Table III. — Summary of results arranged in accordance with the phylogenetic classification of marine organisms Carbon Organic Mean atom ratio Classification Calcium Strontium dioxide matter ^'' 1000 mean % mean % mean "„ mean % Ca Marine Algae, Corallinaceae 27-4 0193 34-2 18-5 3-20 Protozoa, Foraminifera 32-1 0-216 40-6 7-10 3-07 Porifera, Calcarea 11-8 069 19-8 22-2 2-99 Demosponiae 06 trace 5-91 49-7 — Coelenterata, Hydrozoa 37-4 0-711 41-0 4-16 S-69 Anthozoa Alcyonaria 23-2 0-229 30-5 22-0 4-04 Zoantharia 36-5 0-789 40-7 4-40 9-86 Annelida, Polychaeta 32-5 0-421 39-1 6-84 5-87 Arthropoda. Cirripedia 36-3 0-354 40-8 4-65 4-45 Decapoda 24-6 0-328 29-3 27-8 6-17 Mollusca. Amphineura 38 0-669 41-2 4-30 8 06 Pelecypoda 37-9 0-154 41-8 3-18 1-85 Gastropoda Prosobranchia 37-8 0-139 41-8 314 1-68 Opisthobranchia 2-38 0-062 3-97 72-6 10 Scaphopoda 38-2 0-196 42-3 1-81 2-35 Cephalopoda 35-8 0-293 40-1 7-35 3-74 Bryozoa, Ectoprocta 30-4 0-228 35-4 14-7 3-41 Brachiopoda. Articulata 38-4 0-113 42-3 212 1-36 Echinodermata. Crinoidea 25-8 0145 38-2 13-3 2-56 Asteroidea 22-4 01 34 31-8 29-3 2-73 Ophiuroidea 26-3 0153 34-6 20-5 2-69 Echinoidea 33-6 0-199 41-8 5 00 2-70 Holothuroidea 27-8 0-166 35-0 18-3 2-74 Chordata, Ascidiacea 0-34 trace 13-7 69-6 ' 38 Thomas G. Thompson and Tsaihwa J. Chow Table IV. — Strontium-calcium atom ratio of marine invertebrates coltectea at different tide levels Mid-tidal Level Low Inter-tidal Level Atom ratio Atom ratio ^^ X 1000 Ca Sr Ca X 1000 Mytilus calif ornianus 101 Littorina scutalata 1 -44 Pisaster ochraceus 2-76 Diodora aspera Rhabdodermella nuttingi Henricia leviscula 1-35 2-30 2-81 Mitella polymerus 4-27 Balam4s glandula 5 06 Cancer ant ennarius 6-12 Nuttallina calif arnica 7-91 Tetraclita squamosa Pugettia producta Cryptochiton stelleri Balanophyllia elegans Anisodoris nobilis 4-88 6 00 8-80 9-41 11 Table V. — Strontium-calcium atom ratio of Echinodermata in relation to their habitats Atom ratio Habitat Specimen Sr Ca X 1000 Wharf Piling Pisaster giganteus 2-69 Mid-tidal Level Rocky Shore P. ochraceus 2-76 Low Inter-tidal Rocky Reef Henricia leviscula 2-81 Burrowing Strongylocentrotus purpuratus 2-89 Sandy Flat Dendraster excentricus 2-79 Sandy-mud Substratum Ophioplocus esmarki 2-79 Inter-tidal Zone P solus sp. 2-72 Deep water Strongylocentrotus fragilis 2-63 Table VI. — Strontium-calcium atom ratio in relation to the mineralogical character of calcium carbonate in marine organisms Atom ratio Atom ratio Atom ratio Calcite ^'' X 1000 Ca Calcite-Aragonite Mixture J'' X 1000 Ca Aragonite 4f- X 1000 Ca Algae, Corallinaceae 3-20 Mollusca, Coelenterata, Protozoa, Pelecypoda l-94t Hydrozoa 9-491 Foraminifera 3-07 Gastropoda Zoantharia 9-86 Porifera, Calcarea 2-99 Prosobranchia 1-68 Mollusca, Coelenterata, Amphineura 8 06 Alcyonaria 316* Gastropoda Arthropoda, Nudibranchia 10 Cirripedia 4-45 Scaphopoda 2-35 Mollusca, Cephalopoda 3-74 Pelecypoda Anomiidae 1-22 Ostreidae 1-22 Pectinidae 1-31 Bryozoa, Ectoprocta 3-41 Brachiopoda, Articulata 1-36 Echinodermata 2-71 * Does not include the aragonite Heliopora. t Does not include the calcite Anomiidae, Ostreidae and Pectinidae. t Does not include the calcite Errinopora. The strontium-calcium atom ratio in carbonate-secreting marine organisms 39 REFERENCES AsARi, Tamiya (1950), Geochemical distribution of strontium. VII. Strontium contents of shells /. Chem. Soc, Japan, 71, 156-158. B0GGILD, O. B. (1930), The shell structure of the molluscs. Kl. Danske Videnskab. Selskab, Skrifter, Naturvidenskab. math. Afdel., (9) 2, 231-326. Chave, E. K. (1954), Aspects of the biogeochemistry of magnesium. I. Calcareous marine organisms. J. GeoL, 62, 266-283. Chow, T. J. and Thompson, T. G. (1955 a). Flame photometric determination of strontium in sea water. Anal. Chem., 27, 18-21. Chow, T. J. and Thompson, T. G. (1955 b). Flame photometric determination of calcium in sea water and marine organisms. Anal. Chem. 11, 910-913. Clarke, F. W. and Wheeler, W. C. (1922), Inorganic constituents of marine invertebrates. Prof. pap. U.S. Geol. Sur., No. 124, 1-62. DiEULAFAiT, L. (1877), La strontiane, sa diffusion dans la nature minerale. C.R. Acad. Sci., Paris, 84, 1303-1305. Forchhammer, G. (1852), Beitrage zur Bildungsgeschichte des Dolomits. N. Jb. Min. Geol. Paldont., p. 854. Fox, H. M. and Ramage, H. (1931), A spectrograph ic analysis of animal tissues. Proc. Rov. Soc, London, B, 108, 157-173. KuLP, J. L., Truekain, Karl and Boyd, D. N. (1952), Strontium content of limestones and fossils, Bull. Geol. Soc. Amer., 63, 701-716. Lowenstam, H. a. (1954 a), Environmental relations of modification compositions of certain carbonate secreting marine invertebrates. Proc. Nat. Acad. Sci., U.S.A., 40, 39^8. Lowenstam, H. A. (1954 b). Factors affecting the aragonite-calcite ratios in carbonate secreting marine organisms. J. Geol., 62, 284-321. McCance, R. a. and Masters, M. (1937), The chemical composition and the acid base balance of Archidoris britannica. J. Mar. Biol. Ass., U.K., 22, 273-279. MoRETTi, Giuseppe (1813), Sur le sulfate de strontiane trouve dans le corps marins petrifes, et sur deverses combinaisons de cette terre avec quelques acides. Ann. Chem., Paris, (1) 86, 262-273. Noll, W. (1934), Geochemie des Strontiums. Chem. Erde, 8, 507-600. Odum, H. T. (1951 a). Note on the strontium content of sea water, celestite Radiolaria, and stron- tianite snail shells. Science, 114, 211-213. Odum, H. T. (1951 b). The stability of the world strontium cycle. Science, 114, 407-41 1. Rankama, K. and Sahama, T. G. (1949). Geochemistry, p. 226, The University of Chicago Press, Chicago. SCHMELCK, L. (1901), Chemical examination of shells of Mollusca and dried Echinodermata. Norweg. N. Atl. Exper., 1876-1878, Zoo/ogv, 7: (28), 129. Trueman, E. R. (1944), Strontium in molluscan shells. Nature, 135, 142. TsucHiYA, Yasuhiko (1944), Distribution of strontium in calcareous organisms. I. Strontium content of reef-forming corals. J. Agr. Chem. Soc, Japan, 20, 653-654. TsucHiYA, Yasuhiko (1948), Studies on the method for the preparation of strontium from corals and calcareous algae. /. Fac. Agric, Kyushu Univ., Japan, 9, 65-68. Vinogradov, A. P. (1953), The Elementary Chemical Composition of Marine Organisms. Sears Foundation for Marine Research, Memoir IT. Yale University, New Haven. Vinogradov, A. P. and Borovik-Romanova, T. F. (1945), Geochemistry of strontium. C.R. .Acad. Sci., U.S.S.R., 46, 193-196. VoGEL, H. August von (1814), Researches analytiques sur le corail rouge. Ann. Chim., Pans. (I) 89, 113-134. Wattenberg, H. and Timmermann, E. (1936), Uber die siittigung des Seewasser an CaCOj, und die anorganogene Bildung von Kalksedimenten. Ann. d. Hydrogr. u. Mar. Meteor., pp. 23-31. Wattenberg, H. and Timmermann, E. (1938), Die Loslichkeit von Magnesiumkarbonate und Strontiumkarbonate in Seewasser. Kieler Meeresforchungen, 2, 81-94. Webb, D. A. (1937), Studies on the ultimate composition of biological material. II. Spectrographic analysis of marine invertebrates with special reference to the chemical composition of their environ- ment. Sci. Proc. Roy. Dublin Soc, 21, 505-539. Papers in Marine Biology and Oceanography, Suppl. to vol. 3 of Deep-Sea Research, pp. 40-44. Variation, en mer, de la teneur en oxygene dissous au proche voisinage des sediments Par Jean Brouardel et Louis Face Institut Oceanographique, Paris Sununary — The experiments reported in this paper show that the oxygen content of the water under- goes a rapid decrease in the immediate vicinity of the bottom. This decrease is related to the very rapid oxidation of the sediment and the very slow diffusion of the dissolved gas. A LA SUITE d'une serie de dosages executes au large de Monaco en 1952 (Brouardel et Fage, 1954), sur des echantillons d'eau preleves a I'aide de carottiers de divers modeles, par des fonds de 200 a 1000 m, nous avions conclu que la teneur en Og dissous decroit brusquement au proche voisinage des sediments et nous emettions alors I'hypothese que cette diminution de la teneur en O2 pourrait etre plus importante encore au ras meme du sediment. De nouvelles recherches poursuivies en 1953-54 a I'aide d'appareils construits en consequence ont eu pour but de preciser ce point. Trois appareils differents ont servi aux prelevements destines a ces recherches, tous trois ont le caractere commun d'etre entierement en matiere plastique. Le tube preleveur est constitue par un cylindre de plexiglas completement ouvert a ses extrem- ites; ainsi, d'une part, il ne peut se produire aucune oxydation, d'autre part, il n'y a pas de perte de charge et done de melange d'eau du a des etranglements ou des soupapes. Carottier. Simple tube en plexiglas (fig. 1) de 1 m de long sur 5 cm de diametre. A I'extremite superieure un clapet en matiere plastique est maintenu completement eclipse a la descente de I'appareil par un ergot auquel est fixe le cable du treuil. Lorsque le tube touche le fond, le cable en prenant du mou libere le clapet qui, sous Taction de deux ressorts en acier inoxydable, maintient hermetiquement fermee I'extremite superieure du tube. L'extremite inferieure etant bouchee par la carotte, lorsque le tube rencontre le sediment il se trouve simultanement ferme a ses deux extremites et emprisonne une veritable carotte d'eau. Le fonctionnement du carottier, facilement controle au cours de plongees grace a la parfaite transparence du plexiglas, montre que, convenablement leste, il decoupe le sediment exactement " a I'emporte piece ", sans mettre en suspension de particules de vase au-dessus de la carotte. Bouteille de prelevement . Cette bouteille (fig. 2) est destinee a faire des prelevements non plus au ras du sediment, mais a une hauteur determinee au-dessus de celui-ci (5 m dans nos experiences). Le corps de I'appareil est ici encore forme d'un simple tube en plexiglas a chacune des extremites duquel un clapet, en meme matiere, est maintenu a la descente totalement eclipse par un ergot. Le systeme de declenchement des clapets est base sur un principe analogue a celui d'une arbal^te dont la corde est constituee par le cable du treuil. Le carotier, fixe a I'extremite du cable, maintient bandee cette arbalete par la tension qu'il exerce sur celui-ci. Des que le carottier touche le fond, le cable detendu libere les ergots des clapets qui se ferment simultanement sous Faction de deux ressorts. Preleveur de la surface du sediment (fig. 3). Le principe de cet appareil est celui du "Jenkins surface mud sampler " qui est employe en Limnologie. Mais, ici encore, cet appareil est entierement realise en matiere plastique ce qui lui procure une remarquable souplesse de fonctionnement indispensable a la mer. La bouteille elle-meme est un tube de plexiglas de 60 cm de long et de 7 cm de diametre supportee par un bati. Deux clapets, sortes de paupieres, eclipses a la descente, sont liberes des que 40 Fig. I. Carotticr a la descente. clapct ouvert. Fig. 2. Boutcillc dc preIo\cmcnt. Fig. 3. Preleveur de sLirface, clapets ouverts. Variation, en mer, de la teneur en oxygene au proche voisinage des sediments 41 I'appareil louche le fond, puis soumis a Taction de deux jcux dc quatre ressorts qui les entraincnt avec puissance, mais aussi avec ienteur grace a Taction d"un frcin hydraulique. Le clapet situc a la partie inferieure decoupe la pellicule superficiellc du sediment (2 a 3 cm d'epaisseur) ainsi retenue dans le tube. A le remontee les deux clapets ferment hermetiquement les extremites de la bouteille. Contrairement au carottier qui est descendu tres rapidement afin de faire une carotte sufTisamment importante pour boucher Textremite inferieure du tube, eel appareil est descendu exircmcment lentemenl des Tapproche du fond el verilablemenl " pose " sur celui-ci. L'expose suivant a trait aux resultats obtcnus avec ces appareils simultanement employes au cours des sorties effectuees en Juin-Juillet 1954 avec VEider, bateau du Musee Oceanographique. Distinguant parfaitement dans le carottier, grace a la transparence du plexiglas, la limite de la carotte et de I'eau limpide emprisonnee au-dessus d'elle, il est possible de faire des prelevements par siphonage, grace a un fin tube de verre. Les niveaux choisis pour ces prelevements ont ete les suivants: au ras du sediment, toutefois sans atteindre la carotte, puis a 5 — 10 — ^25 — 40 — 55 cm de celui-ci. En outre, comme lors des experiences anterieures (Brouardel et Page, 1953), la bouteille-arbalete, suspendue au meme cable, etait disposee a 5 m au-dessus du carot- tier faisant done un prelevement d'eau a environ 6 m au-dessus du sediment. Les teneurs en O2 de I'eau des divers echaniillons obtenus dans ces conditions figurent au tableau L Ces teneurs y sont exprimees en milligrammes par litre. Tableau I Teneur en O2 de Veau prelevee a Vaide des appareils {Figs. sediment et 2) au-dessus du Profondeur: 100 m 140 m 210 m 220 //; 240 m 245 m 290 m 305 m Moyennes 5 m 7,89 7,33 1,Q1 6,95 6,94 6,37 6,41 6,34 6.92 mc 1 55 cm 7,88 7,23 6,84 7,20 6,86 6,81 6,27 6,21 6.91 40 7,76 7,23 6,93 6,80 6,58 6,79 6,50 6,48 6,88 25 7,83 7,23 7,13 6,78 6,60 6,40 6,42 6,36 6,84 10 7,71 7,56 7,11 6,76 6,39 6,46 6,47 6,22 6,83 5 7,83 7,13 6,67 6,58 6,75 6,47 6,37 6,42 6,78 au ras du sediment 7,66 6,92 6,59 6,65 6,67 5,61 6,31 6,35 6.59 Sur la Figure 4, nous avons reporte en abscisses les teneurs moyennes en O2 des prises aux differentes hauteurs (ordonnees) au-dessus du sediment. Cette courbe fait apparaitre, avec plus de nettete que celles obtenues lors des experiences anterieures, la diminution rapide de taux d'C, au proche voisinage du sediment. L'amelioraiion due au remplacement de I'afcodur du carottier par le plexiglas transparent permet, en effet, de faire les prelevements plus pres de la carotte, a des hauteurs mieux detcr- minees, et d'avoir ainsi Failure de la courbe dans les premiers centimetres. On voit qu'entre la teneur en O2 de I'echantiilon d'eau preleve par la bouteille a 5 m au-dessus du sediment et celle de I'eau prelevee a la partie superieure du carottier on n'observe pratiquement aucune difference. Par contre, la teneur en O. de I'eau renfermee dans le carottier diminue rapidement, et cela d'autant plus que Ton se rapproche de la carotte. Dans les experiences anterieures (carottier a clapet, puis carottier en afcodur) les differences entre les moyennes des taux d'Oj observes entre 42 Jean Brouardel et Louis Face les deux etages extremes etaient de 3,7 et 3,5 %. Ici, la difference moyenne entre les prelevements extremes faits dans le carottier, a 55 cm I'un de I'autre, est de 0,32 mg soil 5 %. Si le carottier employe ici presentait une amelioration sur les precedents il restait, dans son usage, un point sur lequel aucun calcul ne pouvait donner d'appreciation mais qui pouvait, pensions-nous, presenter de I'importance. En effet, pour que la colonne d'eau reste emprisonnee dans le tube il fallait a son extremite inferieure une carotte relativement importante. Pour cela une assez grande Vitesse de descente etait necessaire et nous ne savions dans quelle mesure cette vitesse ne risquait pas de perturber la position des couches d'eau lors de la rencontre du 6 - 4 - UJ 050 O 25 - ^- '///sgdiment////^ 1 1 1 1 66 67 68 69 Fig. 4. Teneurs en O2 de I'eau dans une couche de 6 m d'epaisseur au dessus du sediment (Carottier et Bouteille de prelevement). carottier avec le sediment. D'autre part, a la remontee la carotte risque toujours de glisser legerement le long du tube et de modifier peut-etre, la position de I'eau. C'est pour eliminer ces causes possibles de perturbation que nous nous sommes servis d'un appareil de principe different, c'est-a-dire du preleveur de surface dont nous avons donne plus haut une sommaire description. Cette bouteille " carottier d'eau " et le carottier en plexiglas ont ete utilises simul- tanement au cours des sorties de X Eider en Juin-Juillet 1954, puis elle a ete utilisee avec la bouteille-arbalete, par de plus grandes profondeurs, en Octobre 1954 a bord de la Calypso. Les prelevements, comme dans le cas du carottier, se faisaient par siphonage dont la technique etait d'ailleurs amelioree ici par I'emploi d'une " pige " qui guidait le fin tube de verre dans le carottier bouteille. Lors des sorties de Juin-Juillet, les prelevements faits a des hauteurs de 1, 5, 10, 25 et 40 centimetres, au-dessus de la pellicule decoupee par le clapet inferieur de la bouteille, ont donne les resultats qui figurent au tableau II et qui permettent de construire la courbe figure 5. Variation, en mer, de la teneur en oxygene au proche voisinage des sediments 43 6 - X m I - O 50 - 25 k SEDIMENT 6-5 66 X 6 7 6-8 6 9 O, mq/ 1 71 7-2 Fig. 5. Teneurs en Og de Teau dans une couche de 40 cm d'epaisseur au-dessus du sediment (Preleveur de surface). Tableau II Teneur en O2 de Veau prelevee a Taide de Vappareil (Fig. 3), au-dessus du sediment Profondeur: 85 ni 95 m 130 m 150 m 150 m 180 m 220 m 230 m 235 m 245 m 250 m 255 m 275 m 285 m 430 m Moyennei 40 cm 7,89 7,95 7,53 7,83 7,44 7,36 7,08 7,44 7,07 6,90 6,65 7,08 6,73 7 6,17 7,21 mg 1 25 cm 7,89 7,89 7,53 7,64 7,34 7,24 6,94 6,93 7,09 6,74 6,27 6,96 6,63 6.77 6,91 7.05 10 cm 7,68 7,51 7.45 7,42 7,31 7,07 6,90 6,60 6,89 5,92 6,14 7,07 6,54 6,46 5,88 6,86 5 cm 7,57 7,59 7,25 7,38 6,68 6,54 6,79 6,16 6,64 6,68 6,24 6,82 6,39 6,37 5.82 6.73 au ras du sediment 7,36 7,27 7,04 7,05 6,50 6,39 6,47 6,21 6,57 5,54 6,05 6,35 6,36 6,48 5.74 6,49 La courbe figure 6, deduite du tableau III a trait aux resultats obtenus en Octobre. Elle confirme, a une autre epoque et pour d'autres profondeurs, Failure de la courbe precedente. Si Ton compare maintenant cette diminution du taux d'Oj avec celle due normale- ment a I'augmentation de la profondeur, mais en operant alors largement au-dessus du sediment, on constate que celle-ci est seulement de I'ordre de 0,15 mg 1 par 100 m ce qui represente sur 10 cm une variation de — 0,00015 mg 1, et cela dans la region ou nous operions, a la meme epoque et a la meme profondeur moyenne. Or dans les 10 cm d'eau au-dessus du sediment cette variation est (tableau II) de — 0,37 mg 1, soil Tableau III Teneur en O^ de Peau prelevee a Taide des appareils (Figs. 2 et 3) au-dessus du sediment Profondeur: WO m \60 m 870 m 950 m 1 740 /// A/('it7;/;t-.v 5 m 7,36 7,01 6,97 7,36 6,45 7.03 mgl 40 cm 7,21 7,02 6,86 7,46 6,33 6,96 25 cm 7,18 6.98 6,85 7,04 6.24 6.86 10 cm 7,10 6,53 6,59 7,21 6.17 6.72 5 7,14 6,60 6,45 6,75 6,04 6.6 au ras du sediment 7,06 6.30 5.87 6,69 5.83 6.35 44 Jean Brouardel et Louis Fage 4 X 10* fois plus. On peut done veritablement parler d'une chute du taux d'oxygene au ras du sediment. Ces dernieres experiences ont done permis de preciser I'allure de la variation du taux d'Oa au tres proche voisinage du sediment et ont montre qu'au fur et a mesuxe que les techniques de prelevements s'ameliorent cette variation apparait encore plus nettement sur les courbes. Le sediment constitue done une couche puissamment reductrice, cause du phenomene que nous analysons. Nous avons alors essaye de determiner a quelle vitesse se fait cette oxydation. Avec les precautions necessaires, nous avons etale sur 5 cm d'epaisseur, au fond d'un bac, d'une contenance de 380 litres, 30 litres de sediment preleve au large et dont la couche superieure avait ete dessechee a I'etuve a 110° C. Le bac, haut de 55 cm, I ^.3 ^■•- I - 0-50 O 25 - SEDIMENT /^■^[^ _L 6-3 64 b 5 6-6 6-7 OjmyL 6-6 6-9 71 7-2 Fig. 6. Teneurs en O2 de I'eau dans une couche de 6 m d'epaisseur au-dessus du sediment (Preleveur de surface et Bouteille de prelevement). etant rempli d'eau de mer de teneur en O2 connue, nous avons, chaque jour, dose les variations de cette teneur a diflferents niveaux. II a ete constate que, dans la couche d'eau de 1 cm d'epaisseur au contact du sediment, la teneur en O2 s'abaisse si rapide- ment qu'au bout de 30 heures elle a diminue de moitie; au bout de trois jours, elle n'est plus que de I'ordre du milHgramme et, au bout de neuf jours, elle est si faible qu'elle ne peut plus etre mise en evidence par des methodes sensibles au 2/lOOe de milligramme : cette couche d'eau ne contient pratiquement plus d'oxygene. Pendant le meme temps, la couche situee seulement a 10 cm au-dessus n'a perdu que 0,5 mg/1, Ainsi sont mises en evidence, d'une part, la tres grande rapidite d'oxydation du sediment et, d' autre part, I'extreme lenteur de diffusion du gaz dissous. Ce qui est de nature a expliquer que la chute rapide de la teneur en O2 observee in situ, ne se produit qu'au proche voisinage du sediment. REFERENCES Brouardel, J. et Fage, L. (1953), Variation, en mer, de la teneur en oxygfene dissous au proche voisinage des sediments. C.R. Acad. Sci., Paris, 237, 1605-1606. Brouardel, J. et Fage, L. (1954), Variation de la teneur en oxygdne de I'eau au proche voisinage des sediments. Deep-Sea Res., 1 (2), 86-94. Papers in Marine Biology and Oceanography, Suppl. to vol. 3 of Deep-Sca Research, pp. 45-57. Foraminiferal faunas in cores offshore from the Mississippi Delta* By Fred B Phleger Scripps Institution of Oceanography, La Jolla Summary — Study of Foraminifera from fifteen cores shows presence of cold-water faunas interpreted as representing glacial stages and/or substages, and of warm-water faunas interpreted as post-glacial and interglacial stages and/or substages. These sequences are similar to those previously reported from the northwestern Gulf of Mexico. The amount of post-glacial deposition is greater on the lower continental shelf and upper continental slope than on the lower slope and basin. Variations in amount of post-glacial sedimentation within these topographic provinces are demonstrated. Two cores located in the bottom of Mississippi Canyon contain faunas and sediments which have been displaced downslope, presumably by turbidity currents. It is suggested that the turbidity current was confined to Mississippi Canyon, and that submarine canyons generally tend to localize many turbidity currents. INTRODUCTION Members of the Woods Hole Oceanographic Institution have pioneered in studies of the offshore sediments in the northern Gulf of Mexico. Extensive collections of surface sediments and longer cores were taken along 2,500 miles of traverses in the north- western Gulf of Mexico in 1947, using the research vessel Atlantis. The physical parameters of these sediments were reported and interpreted by Stetson (1953), chemical studies of the materials are discussed by Trask (1953), and the foraminiferal faunas are described and interpreted by Phleger (1951) and Phleger and Parker (1951). In 1951 Stetson made extensive collections aboard the Atlantis in the north- eastern area, from the Mississippi Delta to Florida, collecting surface sediment samples and long cores along several hundred miles of traverses. The foraminiferal facies in the surface sediments along these traverses have been interpreted by Parker (1954), and study of the sediments is being undertaken by Stetson. The present paper is a study of the vertical sequences of foraminiferal faunas in fifteen of these cores in a traverse extending southward from the Mississippi Delta. The purposes of this study are : (1) To discover whether there is a vertical sequence of cold- and warm-water faunas as reported from the western Gulf of Mexico and elsewhere; (2) To attempt to discover relative rates of deposition off a large delta; (3) To evaluate the role of turbidity currents in deposition in the area covered by the cores. The cores were studied at the suggestion of Henry C. Stetson of the Woods Hole Oceanographic Institution, who furnished them to the writer. The assistance of Jean F. Peirson in this study is gratefully acknowledged. Dr. Rufus J. LeBlanc, of the Shell Development Company, kindly arranged to have several of the cores sampled. The laboratory work was supported by the Office of Naval Research (Project NR 081-050, Contract Nonr-233, Task 1). * Contribution No. 21, Marine Foraminifera Laboratory; Contribution from the Scripps Institu- tion of Oceanography, New Series, No. 804. 45 46 Fred B Phleger LOCATIONS OF STATIONS AND DESCRIPTION OF THE AREA The cores were collected along a traverse (Table I, Fig, 1) extending from the deep Gulf of Mexico basin at 3,017 m to the lower continental shelf at 88 m. The near-shore, shallow end of the traverse is only a short distance off Southwest Pass, one of the main distributaries of the Mississippi Delta. .i? o ^ «5 ■S '-^ ■?: 5 o t: o o ^ c/: OS Foraminiferal faunas in cores offshore from the Mississippi Delta 47 Table I— Locations and depths of cores Core Depth in m .V. Lat. W. Long. 3 3017 26^01' 88=03' 4 2972 26= 07' 89= 09' 5 2788 26' 31' 89=09-5' 6 2468 26 ' 58-5' 89 12' 7 1875 27 26' 89 14' 8 1417 27=37-5' 89= 14-5' 9 1372 27=51' 89= 15' 10 1298 28=01-5' 89= 19' II 914 28= 12' 89= 20' 12 732 28= 18' 89 20' 13 631 28=23-5' 89 20' 14 471 28= 29' 89 22' 15 298 28=33-5' 89= 22' 21 142 28= 39' 89=25' 18 88 28M5-5' 89= 27' The topographic charts of the northwestern Gulf of Mexico, constructed by Gealy (1955), end at the approximate position of the Mississippi Delta, and all of the present core stations are off her chart except cores 7-12 (Fig. 2). Core 18 was taken at a depth of 88 m on the lower part of the narrow continental shelf Most of the other cores came from the continental slope, except those at the outer end of the traverse which are in or on the edge of the Gulf of Mexico basin. The continental slope in this area appears to be quite rugged, and is cut by the Mississippi Canyon. The Sigsbee Deep Scarp described by Gealy may not be present in the area traversed by the cores. This is an area of high runoff from the Mississippi River and is presumed to have a rather high rate of sedimentation. It has been shown by Scruton (MS.) that the highest sedimentation rate is close to the delta distributaries and decreases rapidly offshore. There appears to be rather rapid sedimentation for approximately 60 miles offshore, according to analyses by Phleger (MS.). It seems likely that sedimentation is more rapid farther offshore in this area than in any other part of the northern Gulf of Mexico. The physical oceanography of the area is not well-known. Offshore surface temperatures vary from a mean minimum of 20° C in February to a mean maximum of 29° C in August, according to Fuglister (1947). It thus has the surface-water temperatures of North Atlantic mid-latitudes in winter and of low latitudes in summer. A considerable amount of low-latitude water enters the Strait of Yucatan, and while much or most of this flows out the Florida Strait, its effect may be pronounced in the area of the outer part of the present traverse. Offshore salinities in the Gulf of Mexico are approximately 36"/^ 3. A near-shore wedge of lower-salinity water is expected in this high runoff area. Parr (1935) shows salinities of approximately 24°/^^ in the upper 50 m a few miles to the east of the present traverse. METHOD OF STUDY The cores were collected with a coring tube described by Hvorslev and Stetson (1946). The samples used in the present study were one-fourth of each core cut into sections approximately 5 cm in length, so that the entire core was sampled. Each sample was trimmed oi approximately I 8 inch of sediment to prevent contamination between samples and was washed free of tine sediment over a brass sieve having an average opening of 074 mm. 48 Fred B Phleger 84«»00' Fig. 2. Locations of cores 7-12 in relation to topography. Topography modified after Gealy (1955). Depths in fathoms The foraminiferal faunas were analyzed quantitatively. Only a fraction of the population was counted in samples having very large populations; quartering was done by a method described previously (Phleger, 1951,7). Occurrences of species are listed in percent of the total ; benthonic and Foraminiferal faunas in cores offshore from the Mississippi Dcha 49 Table II— Occurrences of Foraminifera in cores 3-6, in percent of total population. Plank tonic and hent/ionic populations computed separately «. _ ___ CORE 6 5 4 J, DEPTH IN METERS 2468 278e 2972 ,,-,; ,. DEPTH IN CORE IN CM. FROM TOP -i to o • -D OC (J ^ (J C It (J * 3^ CD u; o w g C 5 7 ? ? f w ut -^ a m w ut t> - O O (0 ro ~j O U> U> ~ — ro ro T 10 O - Ul .->) .UJ oi ui _ (r x» ^ c Ui ^ - .z - i^ ^ - A u) J 01 ^ ID r\> O ui ui — rvj w ui In V Ol UI Ol m bi a; ry M 01 09 01 01 9 UI UI -J - « ^5 "■ Ol 01 i> 01 - IB I* -. l^ UI .>* C Ui A 01 a [B UI fy - tn & u i _ e rj p ov 01 I* 1^ -^ ID — - . Ui ^ .--* l^ ry TOTAL PLANKTONIC POPULATION G'obigerina bulloides G, digitoto G eggeri G inflota G pochydermo Ui W fV) OI CO o o o o 4 .6 712 - - - b c c H ~ -.J o o 2 ~ 4 ~ C c c ic "en o o "4 2 o o 2 8 4 2 4 c - C ru c 8 5 8 ; p y ~ > b b w V o o o c o o o c 2 5 1 .s 7 9 12 1 b ^ O OOO OOO 4.9 8 1 11 11 10 <£> O O 7 3 CD O O o_ 1" Ol b o o Is rr Ui "oi o O ~ "7 l£ C "4 fi. JT X "x 7 b « -J c . 8 6 o> 2 7 01 Ui b Q 00 00 2 5 7 8 10 1 .2 - t\i ■* - IS 01 000 10 5 9 R 11 910 3 3 Globtgennella oeguiloterolis G lobigennito glutinota Globigennoides congloboto G- f u b f G sacculifera Gioborotalio hirsuta G menofdii 5 4 3 1 5 .3 5 23 ^31 _ _ - - 4 ; 75 23 «C "3 ~ 1 2 7 9 ±_ 5 ~ 2 "6 .6 5 .6 3 .8 43 27 4 1 8 6 5 49 4 26 12 42 4< 25 7 8 ^ 2 4 3 2 17 15 II e .4.3.3.3 55 27 34 57 91622 a: 4 4 5 ?4I7 9 1 52 44 56 9 7 9 3 3 57 6 7 ~ 7 11 8_ .8 10 3 8 2 5« 8 "4 3 36 5( i i .e 4 IC X X X X X x_ X X 3 4 13 .2 3< « i S 6 5 2 6 5 4 12 8 .9 .4 .9 42 90 3E 44 15 >e 16 7 .2 2.2 6 3 1 7 13 8 A 1 7 2 M 11 43 A 9 17 6 A 1 .4.3 G puncTulato 1 4 1 T~ ~7 ~3" 3 2 8 ,5 .7 1 2 5.2 6 "2 7 is "7 7 X IT x" 8 .1 .3 fi-. 8.7 4 G tr uncatutinoides G tumida 3 4 1 ^ _ _ — T I 3~ 3' 8 2 2 4 3 2 2 2 2 2 7 2 2 [6 1 J "5 X X - - 3 8 2 .6 2 2 1 8 2 1 2 .3 1 7 R Or bulino universa 3 "3 — — T — 6 ~ 2 is' 3 t 7 3 3 9 3 3 5 4 5 4 "4 "8 1 X X - - 1 6 3 3.2 3 4 6 .6 R 3 4 4 PuHeniotina o bliquiloculoto Sphoeroidinello dehiscens 3 b - - i 3 -- 4 6 2. 3 8 3 2 2 4 4 3 7 6 3 3 4 3 2 5 4 2 2 6 .4 X , 1 2 3 2 1 5 3 .9 TOTAL BENTHONIC POPULATION Ji o D O '--J O o o o In O o o o O o ^ O c ro no Ji o -J u> O - b w — o o x» o o o o o u' w t» O (7) — OOO -J O m D o -) o o o 01 ro (vj OJ b < < Ji Ui 01 rs> :> ~ i^ - D ^ ^ ru w Amphistegino (juvenile) "~ ^ .5 .6 .6 Angulogenno be Ma 7 Bohvjno olota .6.9 .7.8 1 4 .3 .8 .3 .3 7 6 olbotrossi .5 .4 2 .3 .3 B low mam .8 .5 r 2 2 .2 2 3 5 6 2 2.8 .4 1 1 1 .2 4 2 2 3 1 .5 1 B poula .8 2 2 1 1 .4 1 . 8.3 1 1 . 7 1 .6 B pulchella pnmiTivo 4 .6 6 .5 .4 4 2 .5 1 B striatulo spmoto 2 .4.4 B subaenonensis mexicono 4 u4 .4 .8 .6 4 1 .5. 7 1 B subspinescens 4 .3 .8 1 1 1 1 1 2.3 1 8 1 .5 . 7 Buhmino spicato .2 .5 4 6 B sfrioto meKicana 1 .8 1 .9 .6 2 .6 .3 .5 .3 Bulimineiio cf bossendorfensis 5 1 .4 .4 1 .3.5 .3 5 .7 .3 Concns oblonga 7 Cassidulina subglobosa 5 5 4 5 2 2 1 .5 4 4 .4.8 2 2 2 3 2 3 .7 .5 .7 .7 .5 2 9 2 2 5 5 2 1 .6.6 l£ Cossidu hnoides tenuis 4 1 .4 5 7 .4.8 _^ .5 1 » 53 .i .5 .3 Cibicides hultenberqp 1 1 1 3 2 2 2 1 1 .3 .4 .7 . B .2 1 C fobef tsonionus 4 8 7 3 3 4 2 2 6 7 .5 3 4 3.4 1 3.5 6 3 2 1 1 .7 1 7 7 .6 .9 .5 £ C wueiiersroffi i 8 12 18 13 10 12 13 1 1 6 2 4 13 7 14 20 3 5 12 2 8 10 7 20 24 1 3 24 21 21 5 10 20 4 13 2C 1 6 23 2121 iZ*X Eggerello brodyi 1 .4 ' .5 2 1 2 6 2 "2.4 1 .3 1 .7 . 3 .3 r2 3 J .5 1 1 .3 1 2 .6 Elphidium spp .7 ,3. 4 .6.6 1 Epistominello decorate i 4 2 2 1 4 3 4 1 1 19 .5 ac 9 8 2 1411 8 8 4 10 2 6 .7 .5 2 21 30 3 4 14 3 L 5 2 5 E eitguo 4 .4 .2 .8 2 1 .8 73 1 .3 1 1 E ponides pol'us 3 1 2 .5 4 1 ,5 8 6 2.4 .8 4 9 i 4 .2 6 4 .6 1 ^.6 E tumdulus 1 .8 1 19 .9 3 3 5 2.8 2.8 6 5 5 3 3 1 4 2 3 1 21 5 7 9 2 .6 5 9 E turgidus 4 2 3 1 2 .9 2 4 13 f 27 12 8 7 918 9 14 4 9 3 18 36 3e 2e 7 7 4 6 7 : 4 1331 3e le 9 Globobulimina offims 8 vors 5 12 8 1 1 14 5 5 9 6 18 4 344 7 1 4 45C 5 95« 10 5 .7 12 5 5C 55 36 9 2 ^411 i2€ S Gyrotdino neosoldami 2 .9 .6 .9 1 4 .4 .3 .3 . 4 .i 1 .7 .5 .5 3 .9 1 1 G orbicularis 3 5 .5 I 1 1 .5 I 2 .3 .i 1 .5 Haplophrogmoides brodyi 5 2 2 2 2 .5 . 7 Hoglundino elegons 4 14 14 VI ?R ?^ 4 ?8 1 1 10 1 .6 4 13 631 4 6 2 4 5 IC 2 1 3 8 5 9 5 3 2 Korrer lella brodyi Logenidoe ( 5 8 5 6 7 7 4 4 6 2 3 1 10 11 12 915 2 4 2 7 7 6 2 4 2 3 5 7 7 114 5 4 ■ Miholidoe ? 2 2 2 .6 2 .8 1 .4 2 .5 1 .8 1 .3 .6 1 1 2 t Nonion pompiliordes i 1 2 3 3 2 2 2 2 S.5 .6 2 5.8 3 2 3 2 i ' ) 4 2 1 i 5 1 2 7 7 6 1 2 4 Nonionella o t loniico .4 .7 .3 N opima 2 Plec lino opicu Ions ? ? 3 Pseudoeponides urn bona tus *> 3 3 3 3 3 3 2 1 1 1 f J 2 5C 1 2.4,4 2 3 .4 2 .6 2 5 9 3 1 2 . r.6 2 7 2 Pu llenio spp <■ * 3 f> ? ? f. 7 8 3 1 4 7 2 11 3 2 2 2 : 3 2 3 .7 .5 .6 4 5 4 2 2 5 3 Pyrgo murrhino .4 .5 9 6 3 8 3 i 1 1 .3 J 5 .7 2 .3 1 2 .9 2 6 .6 Qu inqueloculino sp R .5 1 9 8 2 .9 .6 .8 1 .5 3 5 £ .9.5 Robutus spp ? 4 B 2 .* > ■« ' RoTolio beccor n vors ? 4 5 "R" tronslucens ? — — ? .5 3 .8 .3 1 .5 Sig moilina distorto "" ■6 S schlumbergen » ? 1 ? 3 1 2 2 .5 29 7 .8 3,3 .7 ; 2 1 7 4 1 .6 .3 1 Siphonino brodyono .4 S pulchro "" ~ _ _ _ _ J Siphoteitulono rolshouseni .4 .4 1 .5 1 5 7 _ _ 1 .5 .S Tr iloculino tncorinoto 6 ? 1 4 1 2 .4 ■* J .6 .6 fi 3 ? 3 7 9 fi 1 .5 20 1 3 1 2 > I 7 ? ? 4 3 e 2 29 57 4C 2 1 2.4 4 1 .4 .6 2 5 3 3 _ _ 4 .7 .6 « 4 U Virquhno odvena .4 _9 __ ._ _ .6 2 5 .4 .7 ^ ± J - - - 5 .8 1 .6 5 1 2 V complanota / V mexicono - _i - - .6 — - - .7 1 .3 V pontoni Loticonnino pouperoto OTher species i i 1 5 1 8 "5 _5 3 it T 2 ? I 2 6| 3. 6 .6 15 4.4 7 4 6 5 2 26 .4 4 f J3 i 2 ^ 1 _ - _ .5 16 .5 7 f .3 5 7 3 1 • 2 50 Fred B Phleger Table III — Occurrences of Foraminifera in cores 7-11, in percent of total population. Planktonic and benthonic populations computed separately CORE 1 1 10 9 8 7 DEPTH IN METERS 914 1298 1372 1417 1875 DEPTH IN CORE IN CM. FROM TOP H O o (J) (Ti w o o io ro O o ro IN) -.J H O ■0 r\) At OD 01 U1 O o o o* OD ? ID (JI .^ ui ui (D Ul f OD C/i to ro (B 01 OI H O ■D (0 OJ o (j> o a> OD O OJ Ul o OD Ui ui CJl CJl ui Oi OJ OD OI o 9 Oi _ bi OI OJ 9 OI Oi yi OI (D Ul o o ro 9 ro TOTAL PLANKTONIC POPULATION o o 8 "b o o to 8 O o o o o o ro OJ 8 8 o o m o o O o o o 01 -J o o OJ ro o o >£l OI b o O OD o ro O ro O ro 01 O O O o b o o yi b o o rj b o o OJ O o 01 OI ID "ro o o Globigermo buHoides 4 4 8 4 6 6 2 ,4 .9 2 9 1 .5 2 1 13 6 2 .8 2 3 1 4 3 3 5 G eggeri 14 8 19 16 19 1 1 14 13 14 8 16 10 12 14 II 4 6 18 10 10 8 14 BC 8 10 II 9 15 9 G pochyderma 1 Globigerineilo aequHoteralis 1 2 .7 4 .3 ■8 1 6 2 .7 3 3 2 3 .8 2 1 7 2 4 5 6 1 Qlobigerinita glutmata 3 6 7 5 2 7 3 4 4 6 3 5 3 4 5 10 10 1 .6 3 6 2 2 2 6 6 8 8 Globigermoides congloboto A .3 .5 .6 2 .3 2 .4 3 2 .2 3 .6 .5 .3 .3 G rubro 33 37 35 45 4S 41 25 25 27 39 31 32 27 33 35 51 33 2! 29 22 43 35 33 32 41 3d 38 24 36 G socculifero 17 12 13 8 8 8 6 10 14 13 19 5 13 12 12 8 10 6 36 35 25 6 14 15 17 II 2C 24 17 Globorotolio menordii II 8 .6 X X X 9 13 13 8 8 3 13 9 II 7 6 9 7 9 3 7 .3 II 8 9 4 2 .3 G punctulota 2 5 4 6 II 10 1 2 .4 .6 2 2 1 3 .7 13 .4 1 1 .3 14 G scitulo .3 .6 2 2 .2 .5 .8 G truncatulinoides 6 8 4 2 4 5 13 II 12 9 7 7 12 9 10 6 3 12 5 5 4 4 3 5 5 7 2 6 3 G tumido 2 1 2 3 .5 4 3 3 3 5 2 2 4 2 1 1 4 .6 1 3 3 2 2 2 .9 5 2 Orbulino universo .9 ■9 3 1 1 2 3 4 .8 3 2 8 2 1 3 7 2 7 5 5 10 3 8 4 5 6 2 2 PuMeniatino obliquiloculQto 10 8 5 5 2 4 9 12 9 7 8 IC 8 II 7 4 13 12 2 5 7 II 5 4 7 4 5 2 2 Sphoeroidinello dehiscens .4 .3 .8 2 .2 .3 3 TOTAL BENTHONIC POPULATION 01 o O o o o O O o o o o tJI O ro O o a> ot O OD O ro o a> o O OI o -.J o 00 o> o OI O o OJ O 01 o OJ O OD ro O rj O O o 01 o rj oi O o o ro to O Ol o rj O O ro ro (9 O Angulogerino bello .6 Anomolinoides mexicono .3 .1 .3 .1 .2 .7 .3 .5 .1 .2 1 .6 .4 .2 Bolivtno alota . 1 4 .4 .9 .7 2 .1 3 17 3 2 50 S3 12 17 42 3C 33 45 32 46 B albotrossi 3 12 10 .7 1 1 8 II 10 7 ,9 .8 4 8 2 5 5 6 3 1 3 2 1 1 2 7 .3 4 1 2 2 e borbota 3 4 B lowmani 1 2 2 .1 .1 2 2 3 .7 7 1 1 2 2 1 .9 1 2 .2 1 1 .4 .7 B ordinorio 2 2 7 6 12 5 .5 .7 .2 .2 1 1 .5 .6 .4 B pulchello primitivQ .9 .3 B stnatulo spinoto 5 .4 .3 .5 4 .2 B suboenoriensis mexicana .2 2 /» 2 4 .2 6 .6 1 .6 .2 .7 2 2 Bulimino oculeato 4 3 1 6 21 16 8 II 15 18 10 13 21 15 10 21 23 16 14 1 2 1 1 1 7 9 B. olozonensis 6 5 1 2 3 5 4 3 3 1 1 1 2 2 3 2 2 1 2 3 .5 B morginoto 5 2 B spicoto 2 2 .9 4 .8 2 3 1 .7 .3 .4 .9 .3 2 3 .6 2 .5 2 B strjota mexicono .2 .5 2 2 3 .6 2 .4 .7 .8 .9 1 .9 1 4 4 .8 2 .3 .6 1 .7 Buliminello cf. bossendorfensis 1 2 13 .2 .3 1 7 5 1 .5 ,4 Cossidulino corincto .6 .3 1 4 2C 73 .9 .2 .3 ,2 .5 10 2C 2 .5 1 1 1 .7 2 1 C subgloboSQ 9 4 10 5 5 2 9 II 13 4 5 4 6 5 6 4 3 4 1 .6 1 .6 2 3 7 1 II 3 .4 ChMostomello oolino .2 .r .1 .9 .3 .4 1 7 1 22 .5 4 1 .5 16 .4 Cibicides off floridonus .6 2 5 4 3 .8 .2 4 5 5 5 3 3 2 2 3 3 1 3 4 .4 C fobertsomonus 2 .1 .1 2 .5 1 2 .8 1 2 2 3 .6 1 .2 .5 .6 2 3 5 3 C- wuellerstorf 1 .2 2 . 1 .3 .9 1 2 1 .8 5 4 2 2 4 3 2 1 1 3 2 2 .7 3 7 Cibicidino strottont 2 Elphidi um spp 4 1 .3 Epistominello decorofo .3 3 1 2 3 1 .6 2 4 1 4 2 8 5 3 18 1 10 E exiguo 4 5 12 II 3 .2 4 2 4 4 2 1 1 3 2 3 2 2 7 .5 2 .6 3 1 6 .8 .2 2 3 .5 1 E rugoso 2 .8 .9 1 .5 .4 .5 .1 .6 .5 E vitrec 27 19 Eponides polius .4 .2 .7 .8 1 1 .9 .3 2 .5 1 .3 1 .6 .8 .2 E tumidulus .8 .3 .9 .3 2 .2 .2 .6 2 1 E furgrdus 8 5 4 2 5 .6 3 5 3 6 7 8 10 21 12 3 .9 1 2 .6 3 1 .7 .6 3 10 5 Globobulimino offlnis 8 vors .6 .6 2 .4 .9 .5 3 2 .5 1 2 4 .8 .1 1 2 2 23 2 3C 35 II 10 8 2 2 15 34 14 G mississippiensis 2 Gyfoidino orbicularis 3 .8 .6 1 1 2 1 2 4 2 3 4 5 4 2 3 4 2 .6 1 1 ,7 5 3 Hoplophrogmoides brodyi 7 1 .3 .5 2 2 .9 3 .5 Hoglundino elegons .7 .2 .3 .2 .1 .8 .7 .4 3 4 2 1 2 .7 4 2 6 II 5 Logenidoe 2 2 4 3 .3 1 .3 5 4 9 3 4 4 6 4 5 6 7 .7 2 4 1 4 3 2 3 3 4 3 Loticorinino pouperoto 2 .6 1 1 .3 .9 2 2 2 1 2 2 2 1 2 2 .9 2 3 1 2 2 Mi liolidoe .8 2 .6 .5 .2 2 .9 2 .4 5 1 1 1 1 .4 1 1 3 1 .9 .6 1 2 4 3 1 .4 Nonionello otlontico .6 N opimo 6 .2 1 .7 Osongulorio cultur .6 7 13 .7 2 3 2 6 S 3 1 2 3 2 3 3 I 1 5 4 3 1 2 .3 2 3 2 .7 Pseudoeponides umbonotus .5 .1 •3 .1 .3 .5 .2 .4 .5 1 1 .3 .5 .3 .5 3 4 1 .4 .7 £ 1 .4 Pullenio spp. .6 2 1 1 1 .2 4 2 3 2 I 3 5 3 4 5 2 .9 2 3 .6 2 1 .3 1 4 .4 Rectoboltvino odveno .6 Robulus spp .2 .3 .3 .6 2 .2 1 .5 .9 .2 2 .7 1 2 1 .4 .2 .7 .6 .5 "Rotolio" beccoril vors. 4 4 "R", tronslucens 1 3 3 13 12 .2 .3 5 1 5 .3 .9 2 .6 .2 2 3 3 .6 1 .6 Uvigerino porvulo 3 8 U peregrino 1 14 21 22 10 5 .2 6 7 8 5 6 8 8 4 5 4 6 15 2 17 8 5 5 6 3 5 15 ,2 6 Virgulino mexicano .1 .5 7 .2 3 2C 3 2 .5 1 .4 .7 6 ,4 V pontoni 2 2 V tesselioto .2 .6 .1 .3 3 .5 .4 .7 .2 .3 .2 ,3 .2 Other species 82 3 8 16, la I 9 2 2S 8 6 8 —1 2 4« 10 9 24 4 10 4 8 «7 3 5 7 4 5 12 10 7 12 10 7 5 Foraminiferal faunas in cores offshore from the Mississippi Delta 51 Table IV— Occurrences of Foraminifera in cores 12-15, 1o ro 3 'o 8 (71 8 3 b 8 ~J tn ro OS en 10 JJl 01 ro eji Si rj 'm _rei o« ■150 10 1298 >150 11 914 >210 12 732 >205 13 631 >180 14 471 155 15 298 >195 21 142 >195 18 88 >165 Some useful generalizations may be made from these data. The amount of post- glacial deposition is smallest away from the major source of supply, the Mississippi River, and near the centre of the basin ; there is a general increase in amount of deposition closer to the source, as expected. In all but one core shoaler than 1,400 m the coring tube did not penetrate through post-glacial sediment. There is considerable local variation in the amount of post-glacial deposition. For example, in core 5 in the basin there is two to four times as much post-glacial deposition as in nearby cores from similar depths. Core 14 on the upper part of the continental slope has consider- ably less post-glacial sedimentation than nearby cores at similar depths and position. The amount of post-glacial deposition in the cores from the basin and lower continental slope in the present cores is comparable to that shown in cores from the basin and lower slope reported previously from the northwestern area (see Phleger, 1951, Figs. 29-33). The amount of such deposition on the upper slope appears to be larger than in the western area, although there are exceptions. Cores 9 and 10 contain sediments and faunas in their lower sections which appear to have been displaced downslope from shoaler depths, presumably in the form of turbidity currents. The presence of abundant sand, wood fibres and abundant shallow- water benthonic Foraminifera at water depths of 1,298 m and 1,372 m appears to be conclusive evidence for such displacement. Most of the benthonic Foraminifera are species normal for the depth at which the core was taken ; this demonstrates that the displaced material was deposited at the present water depths at those stations. The age of the displacement is post-glacial (W 1), as shown on Fig. 3. Cores 9 and 10 occur in the bottom of Mississippi Canyon (Fig. 2). Other cores in this traverse do not have displaced sediment, at least in significant amounts, and these are located on topography other than the canyon floor. This suggests that the turbidity flow was localized in the canyon. Studies by Shepard (1951) and Ludwick (1950) in the San Diego area have shown that displacement of sediment from shallow to deep water appears to be funnelled down the canyons in that area, and it seems probable that much displaced sediment flows down the channels of submarine canyons in other areas of the world. The restriction of displaced sediment to the Mississippi Canyon floor in the present cores is evidence in this connection. The lower section of core 15, below approximately 165 cm, contains abundant sand grains and plant fibres; this sediment contains no benthonic Foraminifera which are Foraminiferal faunas in cores offshore from the Mississippi Delta 57 characteristic of water shoaler than that at which the core was collected (298 m). It is possible that this sediment was displaced from shallow water, but the absence of displaced Foraminifera suggests that a turbidity current was not the mechanism of deposition. It appears more likely that this material may be river sediment which was carried offshore (approximately 20 miles) during flood stages of the Mississippi River. Shallow-water Foraminifera would not be expected under these conditions. It is suggested that the occasional specimens of shallow-water Foraminifera found in cores 3-8 and 13-14 do not indicate the presence of appreciable amounts of dis- placed sediment in these cores. These specimens may have been deposited by one or more of the following mechanisms : (1) They may have been carried somewhat beyond the limit of turbidity flows because of slow settling velocity or low effective specific gravity. If the specimens contained protoplasm this mechanism would be aided. (2) They may have been put in suspension by wave action when they contained protoplasm, and were carried to their present positions by currents. A few specimens of living, shallow-water benthonic Foraminifera have been reported in offshore plankton tows in the northwestern Gulf of Mexico (Phleger, 1951, p. 36). (3) They may have been deposited by several very small-scale turbidity currents which transported small amounts of sedimentary materials. The presence of specimens of Gumbelina in core 10 is diflicult to explain. These may have been carried out by the river and deposited at their present position, or they may possibly be contaminated by a salt plug bringing early Tertiary or Cretaceous sediments to the surface. REFERENCES FuGLiSTER, F. C. (1947), Average monthly sea surface temperatures of the western North Atlantic Ocean. Papers in Phys. Oceanogr. and MeteoroL, Mass. Inst. Tech. and Woods Hole Oceanogr. Inst., 10 (2), 5-25. Gealy, B. L. (1955), Topography of the continental slope in northwest Gulf of Mexico. Bull. Geol. Sac. Amer., 66, 203-228. HvoRSLEV, M. J. and Stetson, H. C. (1946), Free-fall coring tube: a new type of gravity bottom sampler. Bidl. Geol. Soc. Amer., 57, 935-950. LuDWiCK, J. C. (1950), Deep water sand layers off San Diego. Univ. of Calif, at Los Angeles, Doctor's Thesis. Parker, F. L. (1954), Distribution of the Foraminifera in the northeastern Gulf of Mexico. Bull. Mus. Comp. Zool., Ill (10), 453-588. Parr, A. E. (1935), Report on the hydrographic observations in the Gulf of Mexico and the adjacent straits made during the Yale Oceanographic Expedition on the Mabel Taylor in 1932. Bull. Bingham Oceanogr. Coll., 5, 1-88. Phleger, F. B (1951), Ecology of Foraminifera, northwest Gulf of Mexico. Part I, Forammitcra distribution. Mem., Geol. Soc. Amer., 46, 1-88. Phleger, F. B (1954), Foraminifera and deep-sea research. Deep-Sea Research, 2, 1-23. Phleger, F. B and Parker, F. L. (1951), Ecology of Foraminifera, northwest Gulf ol Mexico. Part II, Foraminifera species. Mem., Geol. Soc. .Amer., 46, 1-64. Phleger, F. B, Parker, F. L. and Peirson, J. F. (1953), Sediment cores from the Nort^h Atlantic Ocean. No. 1, North Atlantic Foraminifera. Repts. Swedish Deep-Sea E.xped., 7, 3-122. Shepard, F. p. (1951), Sand and gravel in deep-water deposits. World Oil, 61-68. Stetson, H. C. (1953), The sediments of the western Gulf of Mexico. Part I. The contmental terrace of the western Gulf of Mexico: its surface sediments, origin and development. Papers m Phys. Oceanogr. and MeteoroL, 12 {4), 1^5. ^. • . j- r Trask, p. D. (1953), The sediments of the western Gulf of Mexico. Part II, Chemical studies ot sediments of the western Gulf of Mexico. Papers in Phys. Oceanog. and Metrorol., 12 (4), 49-120. Papers in Marine Biology and Oceanography, Suppl. to vol. 3 of Deep-Sea Research, pp. 58-67. Seasonal changes in the phytoplankton as indicated by spectrophotometric chlorophyll estimations 1952-53 By Pamela G. Jenkins (Introduction by W. R. G. Atkins) Summary — Estimations of the chlorophyll content of the phytoplankton in the English Channel at station El were continued from September 1952 until August 1953 at ten depths from to 70 m. As before, the species of the phytoplankton were identified by a culture method. Minima of about 2 mg/m^ occur in winter and in June. Maxima at particular depths can occur in March, April or May, thus in 1952 the maximum was in a March surface sample, 34-2 mg/m^, whereas in 1953 sinking of the cells gave, in May, 78-8 mg/m*. The quantity found can be much influenced by the date of sampling. An autumn maximum late in September 1952 gave 21-1 mg/m^ at the surface. The collodion filter disks varied in colour from dark grey or chocolate to a light sandy colour and examination with a low-power microscope shows phytoplankton, stray fibres and sometimes copepods and other animals. Copepods were counted in spring and summer, a maximum of 24 on one disk being found at 25 m on April 27, got from two litres of water. The figures for the column indicate about 300,000 per square metre down to 70 m. The botanical composition of the phytoplankton was studied by the repeated examination, from first signs of growth onwards, of the chemically enriched samples placed in diffuse daylight. Fifty- four species of Bacillariphyceae were recorded. As before Skeletonema costatum, a Navicula sp. and Nitzschia closterium were the most common. Many species of Chaetoceros were identified in the autumn of 1952. Six species of the Chlorophyceae, five of the Chrysophyceae, one of the Cyanophyceae and three of the Cryptophyceae were recorded. The most common species of the first class was a Chlorella, and of the second a species of Coccolithophora grew in each sample. Phaeocystis globosa grew from January to May. The member of the Cyanophyceae was an Oscillatoria. Neither this nor Phaeocystis was recorded for El in the previous year. Hemiselmis rufescens appeared once more. INTRODUCTION In this " Festschrift " number it may well be pointed out that the roots of the science of the sea are sunk deep in time. The adequate study of the sea involves all the exact sciences, including even astronomy, and all the biological sciences. It is not, as often considered, a preserve for zoologists. The early devotees of the study of marine life were just biologists, mainly systematists, for of necessity one must follow Adam and name things. The beautifully illustrated papers of the early workers are highly educational, and remind one that it is not for us to misquote and say, " Surely we are the people and wisdom shall die with us ". I recall with sadness a morning in April 1941 when, in smoking Plymouth, I picked up one page of an old biological work — all that remained of our Athenaeum Library, which had housed so much of the older literature. Perhaps Dr. Bigelow may be considered as having begun at about the end of the old era of amateur biologists. He was studying the phytoplankton of the Gulf of Maine in 1913. He found the entire basin occupied by a peridinian plankton, but never found diatoms in abundance in July or August except close along the coast and on Georges Bank. 58 Changes in phytoplankton as indicated by spectrophotometric chlorophyll estimations 1952-53 59 So when, largely by his efforts, the Woods Hole Oceanographic Institution was founded, they could begin with a basic knowledge of the phytoplankton which is still lacking in Plymouth. But quantitative work on the production of phytoplankton in the English Channel has been carried out since 1921, first by calculation from the changes in hydrogen ion concentration brought about by photosynthesis (1922) and since 1922 from the consumption of phosphate (1923). The arrival of a modern spectrophotometer however made it possible to estimate the phytoplankton crop by extracting the chlorophyll from collodion filter membranes on which even the smallest green flagellates had been retained. This was done in the year 1951-52, and by a culture method, similar to that of bacteriology, the organisms were grown and multiplied so that even those originally very sparsely distributed were not missed (Atkins and Jenkins, 1953). The chlorophyll method of course gives the phytoplankton content of the water when sampled, whereas the phosphate calculations give the amount produced over a period. The two may be very different. A beginning was thus made in obtaining a better knowledge of what plants were present — and of when they flourished — also such work provides a basis for the study of the movement of water masses tagged by a known algal flora. I therefore asked my collaborator Miss P. G. Jenkins to continue this research and to give her results, which she has done as follows. W. R. G. A. ORIGIN OF SAMPLES AND THEIR EXAMINATION Water was collected with a Nansen-Pettersson water bottle at the international hydrographic station England No. 1 (El), twenty miles S.W. from Plymouth, at a series of depths from m to 70 m bottom. Two htres of each sample were filtered through a collodion (Gradocol) membrane of average pore diameter one micron. The phytoplankton cells and the suspended matter which remained on the disks were examined under the low-power microscope. Then 10 ml of an 80 per cent aqueous acetone solution was used to extract the plant pigments from each membrane. A " Unicam " spectrophotometer with 4 cm cuvettes served to measure the minimum percentage transmission in the red between 640 and 670 my.. These values were converted into concentrations of chlorophyll in mg 1 read off from a graph of the transmissions of 80% aqueous acetone solutions of a dry commercial chlorophyll plotted against their concentrations. This graph and the absorption spectrum of the chlorophyll may be seen in the 1953 paper. Using 10 ml of the aqueous acetone 10 extract the plankton from a litre of water, it is obvious that the chlorophyll has been concentrated one hundred times, so 1 mg/l as read oflFis equivalent to 001 mg, 1 or to 10 mg, m^. An allowance was of course made for the actual volume extracted. In winter, i.e. November to February, the colour of the extracts was slightly yellow and the green was almost imperceptible. The colour deepened with the spring growth to a deep olive green, and lightened in the summer samples. EXAMINATION OF THE PLANKTON ON THE DISKS The disks varied in their intensity from a dark grey or chocolate to a very light sandy colour. It was impossible to deduce the amount of chlorophyll in the extracts from these shades. That deduction could only be made when the disks were a uniform faint green. They were often covered with diatoms of various species, and at times had green spots due to the presence of some species of the Chloro- phyceae. The Dinoflagellate Ceratium tripos occurred at every depth on Aug. 10. 1953. Fibres were frequently seen. Many copepods were found on the disks and as they were so numerous in the spring of 1953 their numbers were counted and set out in Table I. The totals in the second line from the bottom'(S means) are based on the sum of the means for 2-5, 7-5, 12-5 67-5 m for 14 depths. This sum is then multiplied by 5 (for 5 m intervals) and the totals in the bottom line are expressed as 60 Pamela G. Jenkins the number of copepods in the column per square metre of surface, namely in 70 m^. (The amounts of chlorophyll in the water column (Fig. 5) to be considered later were conducted in the same way.) These totals were included because the phytoplankton and zooplankton crops are obviously related. In 1951-52 some medusae were found but none could be seen in the following year. Nudi- branch veligers were found from June to August, though not previously seen on the disks. Table I — Numbers of Copepods found on collodion membrances from April until August 1953, in 2 litres of water at station E\ Depth April May June July August (m) nth 11th nth 22nd 20th lOth 4 16 4 11 5 10 15 4 6 14 5 4 3 2 3 10 22 6 15 20 10 7 14 7 15 9 25 7 24 3 12 18 8 30 10 6 10 1 14 6 40 6 9 5 — 12 50 3 7 4 6 4 3 70 5 9 14 3 S means* 70 124 148 62 130 45 thousands per m^ in column 175 310 370 155 325 112 * See text, p. 59. Table II — Diatoms identified in enriched cultures exposed to light Signs : — Not found A present B more frequent C plentiful D very plentiful O not found in one of the two cruises Where two letters occur for one month, two cruises were undertaken Bacillariophyceae Years J M M O N D COSCINODISCACEAE Coscinodiscus sp. C. concinnus C. excentricus C. radiatus Melosira moniliformis Paralia sulcata Skeletonema costatum Thalassiosira sp. T. condensata T. decipiens T. gravida T. nordenskioldii T. subtilis Biddulphiaceae Biddulphia sp. B. regia B. sinensis Ditylum brightwellii Eucampia zoodiacus 1952 1953 1952 1952 1952 1952 1953 1952 1953 1952 1953 1952 1953 1953 1952 1953 1952 1953 1953 1952 1953 1952 1952 1952 1952 1953 1953 B — A B B B D CO AO AO AO B — DO BO BO — BO — — — — — A — — — — C — — — — — A — — B — A B — — OB B A C B B — C D — BO — — — — DO — — AO AO B — — — D D B D A D D D B B D DD D C D DD D B B D D D D D __________A — __________A — — — — — — — — — B D C A A — — — — — — — — OD— — A ___________ Changes in phytoplankton as indicated by spectrophotometric chlorophyll estimations 1952-53 61 {Table II cont.) Bacillariophyceae Years Chaetoceraceae Chaetoceros sp. 1952 1953 C. affine 1952 1953 C. ceratosporum 1952 1953 C. compressum 1952 C. convexicorne 1952 1953 C. convoliituin 1952 C. cur vise tuni 1953 C. danicum 1952 1953 C. debile 1952 1953 C. decipiens 1952 1953 C. densuin 1952 C. did V mum 1952 1953 C. gracile 1952 C. laciniosum 1953 C. lorenziamim 1953 C. pseudocrinitum 1952 1953 C. scolopendra 1952 C. septentrionale 1952 C. simplex 1952 1953 C. sociale 1952 1953 C. teres 1952 Leptocylindraceae Lauderia borealis 1952 1953 Leptocylindricus 1952 danicus 1953 Rhizosoleniaceae Rhizosolenia sp. 1953 R. alata var. indica 1952 1953 R. fragilissima 1952 R. hebetata 1952 1953 R. setigera 1953 R. shrub solei 1952 1953 R. styliformis 1953 Fragilariaceae Asterionella japonica 1952 1953 1952 Fragilaria sp. F. oceanica «[ 1953 1952 F. striatula 1953 Tabellariaceae Thalassionema 1952 nitzschioides 1953 F M M J S O N D B— — — — — — D C — A C A ABO — — OBB C DOD— — — — — — — — — — A A A — A __ — — __D — — — — — — — — — — — — A — — A — — _ — _OA— — — — — D — — — — — — — — C D C H A ______ — — — — — A A A — — AO— — OA— — — ODD — _-______ — A _ — — ______ — — — A D — B — — — — — — — — — — — __ — ___ — — B __ — — — — — — — A _ — — — — _____ — — — — A — A — — — — — — — A _ — — — ____ — — — — A A — — ____ — — — — A — — — ____ — _ — — A _ — — A _— DO— — A _ — — — — zE^zzz = = = = I = ___ — — — — — — — — B C _ DO BO — — — A — — — — ____ — — C B A A— — ______ _ C DDD— — B _ — — C— — — — — — — ___ — — -A C A A-- ___ — — OBD D — — — — ZZZZZZa — DDD— — ___--OA----D D ______ — — A A — — _ — OA — — OD D — - — - A — B R \i C D — — l)D B A — A — C D DA AO OD — A A — — B B — — AO BO D A D C D _ D OD D D 62 Pamela G. Jenkins {Table II cont.) Bacillariophyceae Years J F M A M J J A 5 O N D Naviculaceae Navicula sp. 1952 — B C D D D C B C D D B 1953 A A AA AB C DC — A C DD N. membranacea 1952 — — — — — — — — — A N. vanhoffeni 1952 — — — — — — — — — B — 1953 A — DD AO — OA c — — — D Pleurosigma sp. 1952 1953 B A AO — — — — — A C D D Bacillariaceae Bacillaria paxillifer 1952 — — — — — — — — B — — — Nitzschia closterium 1952 D C C D A C D C B D D D 1953 D D DB DD D C D D DD D D N. delicatissima 1952 — — — — — — D C A D B 1953 D — — DB B AO D D — — — — N. seriata 1952 — — — — — — — B D D — A 1953 — — — AO — — — — D DD D D SEASONAL VARIATIONS IN CHLOROPHYLL AND THEIR CONVERSION INTO WET WEIGHT OF PHYTOPLANKTON Water samples were taken at intermediate depths from the surface down to 50 m from August to October 1952, then down to 70 m from November 1952 to August 1953. It was decided to extend sampling down to the lowest depth possible, 70 m to give a more accurate survey at El, so that the chlorophyll content for 70 m was known and mq/m' Fig. I. Variation in chlorophyll from August-December 1952, at Station El. Ordinates show depth of samphng in metres. Abscissae show concentration of chlorophyll in milligrams per cubic metre. Changes in phytoplankton as indicated by spectrophotometric chlorophyll estimations 1952-53 63 need not be assumed to be the same as at 50 m, which was done in the 1953 paper. Fig. 1 illustrates the autumn growth from August 1952 (linking up with Atkins and Jenkins, 1953, Fig. 7) to October, with a surface outburst in September. Growth fell to a value almost uniform with depth during November and December in an almost isothermal water column. Table III — Algae identified in enriched cultures exposed to light Signs : — not found A present B more frequent C plentiful D very plentiful O not found in one of the two cruises Where two letters occur for one month, two cruises were under- taken Chlorophyceae Years F M A M J O N D Chlamydo- monadaceae Chlamydomonas sp. POLYBLEPHARIDACEAE Pyramimonas sp. Chlorellaceae Chlorella sp. Ulotrichaceae Stichococciis sp. Ulothrix subflaccida Chaetophoraceae Ectochaete sp. Chrysophyceae Chrysomonad sp. Coccolithophora sp. ISOCHRYSIDACEAE Dicrateria sp. OCHROMONADACEAE Ochromonas sp. Chrysoca psaceae Phaeocystis globosa Cyanophyceae Oscillatoriaceae Oscillatoria sp. Cryptophyceae Cryptomonad sp. Cryptomonas sp. Hemiselmis rufescens 1952 1953 1952 1953 1952 1953 1952 1953 1952 1953 1952 1953 1952 1953 1952 1953 — — B — C C BB OA D B D 1953 1953 1953 i A 1953 1952 1953 1952 1953 I B 1952 ! D 1953 — DA C B D D — A A C — B— — — — — — B A — OC— — A _____ _DDDCBAADCD BBB— ADB— — — — — — — C DC C B— — — — — — OB OA — DO — — — — D — A _____ — — — — — AAOAO— — — — — — — — — C D A C — — B A A A — AO— — OC— — — — — — _____B ___ — — C---------B D D D D D D D CD DC D DD B D D DD D D __OC — BC — — — — — — A — — — — — — — — — — AOCDBD — — — — — — — _— OA— — — — — — — — ___--B----B _ _ BA - - -• - - 1^^ ^ A £3ZZZZZ — — D D _ OC - _ — B OD — 64 Pamela G. Jenkins nt 30 Fig. 2. Variation in chlorophyll, January-April 1953 at El, in mg/m^. The two winter months showed no appreciable change (Fig. 2), but from March onwards the samples gave a high value of chlorophyll at the surface and even at 70 m, giving a bottom maximum at 70 m of 78 -8 mg/ra^ on March 25th. These high readings for the lower depths showed that the cells must have sunk. Riley (1941 b) observed this high concentration of chlorophyll from 40-70 m at Georges Bank during March and April. Marshall and Orr long ago (1928) reported that " during the spring m 30 "O " lO "^ ^O 30 3 40 SO 60 7C mq/m Fig. 3. Variation in chlorophyll, May to August 1953 at El, in mg/m^. Changes in phytoplankton as indicated by spectrophotometric chlorophyll estimations 1952-53 65 months it was not uncommon in shallow water to find more plankton near the bottom than anywhere else ". They also found that during the spring outburst the phyto- plankton caused such a great reduction in submarine illumination that the compensa- tion point (or depth) was at times raised to 5 m or less. The sudden drop from spring growth, May 1 1th to the summer minimum, June 8th, is seen in Fig. 3. The May readings were very high due to the abundance o^ Phaeocystis globosa. Two filtrations were necessary because the disks became clogged, so ordinary filter paper was first used and then the special membrane. Each of these was extracted twice. The later cruises show the gradual growth near the thermoclinc, building up to the autumn maximum, with a peculiar outburst at 25 mon August 10th of 31 -5 mg m\ This outburst must have occurred in a region where light intensity allows photosyn- thesis to take place and its occurrence at only this depth is probably due to the absence of sufficient nutrient salts in the upper 15 m. The results for surface and bottom chlorophyll throughout the year are seen in Fig. 4 with a surface maximum in September and April and a bottom maximum in SEPT 1952 APL I MAY 1953 rnmg^.rf ' The\?a «mp™tuT«\ surface, above, and for .he bo..on, ,70 n„ ,us, below ,hc ° ■ top line or the trame. D 66 Pamela G. Jenkins March, due to sinking. The thermoclines for the year were found on September 22nd, 1952, 15 m, 14-6° C; April 27th, 1953, 15 m, 12-5" C; June 22nd, 20 m, 13-0° C; August 10th, 15 m, 17-2° C. In both 1951 and 1952 the autumn maximum (Fig. 5) was in September, the amounts being respectively 0-59 and 1 -02 g/m^ The spring minimum was in Novem- ber 1951, but was three months later the following winter when it was reached in February 1953. But the values were identical, 0-25 g/m^. The spring maximum in April 1952 was 1-33 g/m^ but this value was obtained on the assumption that the amount of chlorophyll was uniform beyond the last depth examined, 50 m to 70 m, bottom. Later work showed that, on account of sinking of the cells, this was probably an underestimate. The maximum was in May in 1953, 3-68 g/m^ (very much higher) with much Phaeocystis present. Both the summer minima fell in June, 0-15 g/m^ for 1952 and 0-35 g/m^ for 1953. 40 3 O 2 O g/r lO - A - - A \\ - - f - - 19 i2-5: 1 — ( ^ \ 951-5 2 - \ \ / \ 1 >A, — - o \ / i \ . / > V - ^ *J ^;r=t9=- »r r x-, o // ,. e o BiGELOw, Henry B. (1926), Plankton of the Offshore Waters of the Gulf of Maine. Bull. L.6. Bur. Fish., 40 (2), Document No. 968, 1-509. ^. , ,^ „. , ^ Hendey, N. Ingram (1954), A preliminary check-list of British marine diatoms. J. Mar. Biol. Assoc, 33,537-559. ^ ^. , . , , ., Marshall, S. M. and Orr, A. P. (1928), The photosynthesis of diatom cultures in the sea. J. Mar Biol. Assoc... \5,7>2\-60. . d- ; « it Parke, Mary W. (1953), A preliminary check list of British marine algae. J. Mar. Bu>t. .Assoc. JZ, 497-520 Riley, G. A. (1941 a). Plankton studies 3. Long Island Sound. Bull. Bingham Oceanogr. Coll., 7 (3), 1-93 Riley, G. A. (1941 b), Plankton studies 4. Georges Bank. Bull. Bingham Oceanogr. Coll., 7 (4), 1-73. Papers in Marine Biology and Oceanography, Suppl. to vol. 3 of Deep-Sea Research, pp. 68-73. Water replacements and their significance to a fishery By H. B. Hachey Chief Oceanographer, Canadian Joint Committee on Oceanography Summary — Attention has been directed to some of the major and more apparent effects that water replacements may have on various fisheries. It has been indicated that such interchanges may be responsible for the destruction of a fish population or the extension of others. The loss of larvae to scallop and haddock areas has been considered as the result of movements of a water mass, and it is suggested that the availability and catchability of certain species is effected by certain water replace- ment phenomena. When consideration is given to the cycle of life in the sea, it will be quite evident that many indirect effects may follow from these water replacements. While the processes of long- term replacements are not too well understood, wind action is one of the more apparent major casual factors in short-term replacements. INTRODUCTION While the changing characteristics of a water mass, such as temperature and salinity, are followed in detail in the study of a fishery, the replacement of one body of water by another is a process which, generally, only attracts attention when the characteristics of the water bodies concerned are in considerable contrast. Whether it be of the estuary, a coastal area, or the open ocean, the particles of a water mass are never at rest, being subjected to the internal and external forces of gravity, pressure, wind, tide and the Coriolis force. The resultant movements of the water particles under these forces bring about the flushing of estuaries, the replacement of water masses in a bay or coastal area, the removal of water masses from a fishing bank, and the transportation of large masses of ocean water from one location to another. The flushing of estuaries has been given considerable attention during recent years, due to the need of considering the many problems of pollution. The principles derived from such studies furnish an insight into the mechanism involved in the replacement of waters in a comparatively small area, almost completely land-bound. In contrast, and on an ocean-wide scale. Cooper (1954, 127), has called attention to the large variations in the phosphate content and biological productivity of the English Channel in the last thirty years. He seeks an explanation in the replacement processes involving the replenishing of deep Atlantic water by potentially rich northern waters, these deep Atlantic waters eventually upwelling to determine the nutrient supply of the coastal waters of north-eastern Europe. Attention is directed herein to many of the direct consequences to a fishery of the various processes of replacement. THE DESTRUCTION OF A FISHERY BiGELOw and Welsh (1925) describe the disaster to the tilefish, which first came to light in March, 1882, when multitudes of dead fish were observed floating on the surface between the latitudes of Nantucket and Delaware Bay on the Atlantic coast. The area of destruction was at least 170 miles long by 25 miles broad, and covered the entire zone inhabited by the tilefish north of Delaware Bay. It is estimated that 68 Water replacements and their significance to a fishery 69 at least a billion and a half dead tilefish were sighted. There is evidence to indicate that the destruction of this fishery was caused by a sudden temporary flooding of the bottom by abnormally cold water. It has been shown that the tilefish of the Atlantic coast occupies a very definite environment, for it lives only along the upper part of the continental slope where the water temperatures are approximately 10" C, and never ventures into the lower temperatures on the shoaling bottom nearer land. We have in BiGELOW and Welsh's account of the disaster to the tilefish, a well-documented record of the significance of a temporary incursion of waters of contrasting tempera- ture, which in this case brought disaster to a fishery. On the basis of present knowledge, the source of the abnormally cold water responsible for such a flooding is to be found to the eastward. A study of the slope water off the Scotian Shelf (McLellan, et al, 1953), has shown that, between the northern boundary of the slope water and the continental slope, there is found in varying quantities a body of cold water, less than 0-0° C off" the Grand Banks, and less than 4-0'' C east of Sable Island. The quantity of such water, acting as a cushion between the slope water and the continental slope, decreases with westward progression until the slope water generally makes contact with the continental slope to the west- ward of Emerald Bank. Northerly and southerly migrations of the northern edge of the slope water regime, and westerly progressions and easterly withdrawals of the colder waters along the continental slope, provide the mechanism for producing sharp and sudden changes in the water temperatures on the continental slope and over the outer areas of the continental shelf. While the disaster to the tilefish was of major import, and the westerly progression of cold water must have been greater than normal, similar phenomena on a less spectacular scale have been noted in recent years. THE EXPANSION OF A FISHERY In contrast, both as to time and extent, to the temporary incursion of colder waters, bringing about the destruction of a fishery over a limited area, is the historical record of the expansion of the Greenland fishery in recent years with the strengthening of the Atlantic influence in northern latitudes. Passing over earlier history, about 1820, cod were known to be present in enormous quantities in West Greenland waters, as far north as Disko Bay (Jensen and Hansen, 1931). Thereafter, they were absent for a long period of years. Between 1845 and 1849, cod were again plentiful in the Greenland area and then entirely disappeared. From 1917 there was a marked upward tendency in the fishery. The catch increased from approximately 1,000 tons in 1925 to greater than 12,000 tons by 1945, and this fishery has persisted to the present. In dealing with the state of the West Greenland Current up to 1944, Dunbar (1946) points out that, by 1928, the waters for the west coast of Greenland were considerably warmer than in the preceding period, and that the peak warm year was reached within the period 1930-40. The increasing Atlantic influence however was clearly evident in Latitude 72 N in 1942. A comparison of oceano- graphic conditions in Hudson Bay and Hudson Strait, as observed in 1930, with those of 1948 (Bailey and Hachey, 1951) has shown that the observed higher temperatures and salinities of 1948 are indicative of the increasing Atlantic influence in northern waters generally. A meteorological study of Sherbog (Dunbar, 1946), has suggested that the warming of the Arctic and sub-Arctic regions from Greenland to Siberia, which has taken place in recent years, is one manifestation of a large scale climatic 70 H. B. Hachey cycle of a period of about 225 years, which has presumably passed its maximum. It must be stressed here that the expansion of the Greenland cod fishery is associated with the extension of warmer Atlantic waters into more northerly latitudes, although such an extension is probably associated with the climatic cycle. THE LOSS OF LARVAE Studies on the replacement of Bay of Fundy waters were initiated some years ago (Hachey, 1934), and it became evident that the main factors involved in the replace- ment of these waters were land drainage, wind, and tide. Other things being equal, an excess of south-westerly winds favoured the retention of surface waters within the Bay and thus nullified the normal dynamic tendency for surface outflow and renewal of the waters at greater depths. Under such conditions a type of " closed circulation " is set up within the Bay, as opposed to an " open circulation " when surface waters are carried out of the Bay to be replaced by inflowing deeper waters. During the summer months the " closed circulation " favours higher surface water temperatures, and the surface temperatures, as well as the temperature gradient in the upper layer, can be used as an indication of the type of circulation prevailing. Dickie (1955) has made use of the temperature data for the Bay of Fundy to show that successful year classes of scallop are produced in those years when the " closed circulation " prevails. Not only do the warmer surface waters hasten development of the scallop larvae to the setting stage, but the " closed circulation " favours the retention of the larvae within the Bay where they settle and grow on suitable scallop bottom. Under the " open circulation " system, as indicated by lowered surface temperatures, the larvae are carried out of the Bay and are lost to the Bay of Fundy scallop fishery. The replacement of Bay of Fundy waters is not necessarily a regular progressive process, even when the " open circulation " system prevails. Ketchum and Keen (1953) have worked out various mean flushing times and exchange ratios for various parts of the Bay of Fundy, and they point out that although their conclusions were based upon the distribution of river water, the same exchanges may be expected for any material or organism transported by the water. Ketchum and Keen calculated an average flushing time of 74 days for that part of the Bay of Fundy between Cape Chignecto and a line south of Grand Manan. Bailey (1953) has shown that within the period October 6th, 1952, and November 2lst, 1952, the waters of the Bay of Fundy were almost completely replaced, a replacement that only became apparent when it was found that the salinity throughout a section had increased by more than 0'5%o within the period. It might be emphasized that within this period of forty- six days (or less), practically all free-moving larvae in the surface layers would have been carried out of the Bay of Fundy. So too would all free-swimming forms which were feeding at random. The higher salinities as found on November 21st indicate that the replacement involved the inward movement of waters originating from sub-surface depths, and these waters would necessarily carry into the Bay non-swimming forms, and free swimming forms which were feeding at random. According to Walford (1938) the survival of the eggs and larvae of the haddock on Georges Bank is dependent on the variations in the current system in this area, these variations being controlled to a large extent by winds (Bigelow, 1927, 857). He suggests that the loss of the eggs and larval population in 1932, which occurred some time after the first week of March, was due to the removal by currents. Similarly Water replacements and their significance to a fishery 71 BiGELOW (1926, 77) wrote: " at the time of our March and April visits (to the north- eastern part of Georges Bank) in 1920, the presence of newly spawned eggs in abund- ance right out to the 1,000 metre contour proved that a drift out to sea was then taking place from the southern point of the Bank." Walford (1938, 49) states: " there was a very important difference between the circulatory picture in the season of 1932 and that of the corresponding period in 1931. While in 1931 the water movements were such as to permit the bulk of the eggs to remain on the bank and hatch there, in 1932 there were currents carrying eggs off the northern and southern edges into deep water where they were probably lost to Georges Bank." In summary, Walford (1938, 55) points out that a change in the circulatory system on Georges Bank may be disastrous to the haddock brood, and consequently, may be an important cause of fluctuations in abundance. Carruthers, et a/. (1951) have directed attention to the variations in brood strength in the North Sea haddock in the light of relevant wind conditions, and the results of these studies have directed considerable effort to the analysis of wind systems as related to other North Sea Fisheries. AVAILABILITY OF THE FISH The displacement of surface waters on a grand scale by wind is amply demonstrated by the great ocean currents of the North Atlantic, and the fundamental principles have been thoroughly outlined in the classical work of Ekman (Sverdrup, et al., 1942, 492). Ekman has shown on a theoretical basis that a surface current set up by wind is directed 45° to the right of the wind direction in the northern hemisphere, and this has been shown by observation to be a satisfactory approximation in deep water. In shallow water the deflection of the surface wind current is smaller. A wind blowing parallel to a coast is thus effective in transporting surface waters towards the coast, when the coast is to the right of the wind (cum sole from the wind direction) in the northern hemisphere, and away from the coast when the coast is to the left of the wind. The capture of young herring along the open coasts of the Bay of Fundy and the Gulf of Maine is chiefly dependent upon fixed shore weirs. Huntsman (1934) states: " that the herring are to a considerable extent quite near the surface during the twenty-four hours of the day is not only a matter of direct observation, but a requisite for successful operation of the weirs." Huntsman also states (1934, 96) that '* the herring may be treated as a planktonic form ". It then becomes evident that herring in the surface waters, and considered as a planktonic form, will be moved on or ofl the shore with the varying direction of the wind. This is particularly pertinent on an open coast, such as that of Maine, and the south coast of New Brunswick, where the availability of the herring to the shore-fixed weirs must be determined in part by the varying strength of the prevailing south-west winds during the summer months. TEMPORARY ADJUSTMENTS IN A FISHERY Large scale water replacements of a temporary nature sometimes bring about sharp and sudden changes in water temperature, the changes in environment being sufficient to cause a body offish to either: (a) move away from a fishing ground, or (b) cease feeding and thus fail to be attracted to the bait. 72 H. B. Hachey The water replacements in the Halifax area associated with the formation and subsequent movement of a tropical cyclone have been described (Hachey, 1934), and it has been shown that bottom water temperatures in some of the inshore areas increased from 2-0" C to greater than 15-0° C in less than one week. Under these circumstances, in these areas, all fishing for cod and haddock ceases, and does not resume until temperatures return to more normal values (Hachey, 1934; McKenzie, 1934; Vladykov, 1933). Incursions of warmer slope water over the Scotian Shelf have been observed (Hachey, 1953), when bottom temperatures reached values as high as 12-0° C, about five degrees above normal seasonal temperatures. While no records of fishing effort are available for periods in which such incursions have been observed, it is well known that such bottom water temperatures are unfavourable to cod and haddock, the groundfish probably moving out of the areas subjected to the incursions of waters of such temperatures. BARREN SEA FLOOR In the Gulf of St. Lawrence comparatively extreme variations in temperature and salinity have been observed at depth during the summer months (Lauzier, 1952) brought about by oscillations of the various water layers of contrasting characteristics on a gradually shoaling sea floor. It has been pointed out by Lauzier that organisms which cannot tolerate these sudden changes in temperature and salinity will not form important populations along the margins of the Magdalen Shallows, and Hunttsman (1918) has shown that there are bands of the sea floor between Cape Breton and the Magdalen Shallows which are comparatively barren. The oscillation of these water layers of contrasting temperatures is probably responsible for the periodic complete destruction of scallop beds which become temporarily established and fishable under marginally satisfactory conditions in the Magdalen Shallows (Annual Report for 1953, Fisheries Research Board of Canada, 34). MARGINAL FISHERIES In Canadian Atlantic waters, which are contained in the area of confluence of three current systems with waters of contrasting characteristics, extreme contrasts in water characteristics, chiefly temperature, are to be observed. Very sharp boundaries, both vertically and horizontally, are thus encountered, and various marine organisms suitable to one environment (frigid) or another (temperate) exist on a marginal basis. A small vertical or horizontal change in the margin sometimes exerts a very pronounced effect on a fishery. With progressive warming of the waters during recent years we have witnessed the northerly expansion of successful oyster production in the Gulf of St. Lawrence, and the northerly extension of the green crab to the Bay of Fundy (Annual Report for 1953, Fisheries Research Board of Canada, 25). As mentioned earlier in this paper, the expansion of the west Greenland cod fishery is probably the most outstand- ing result of the increased Atlantic influence in northern waters. It is to be expected that a downward trend of temperatures would bring about a recession of these extensions. In some cases, man is responsible for changes in environmental conditions, and this is particularly true in estuaries where pollution problems arise with present-day Water replacements and their significance to a fishery 73 industrial developments. Of a different nature are the possible changes in ihc environment of Georges Bay, between Cape Breton and the Nova Scotia mainland, which may follow from the completion of the Canso Causeway. By cutting off the interchange between Gulf waters and those of the open Atlantic through Canso Strait, the strong possibility exists that the waters of Georges Bay will be warmer than heretofore. Slight upward changes in the water temperatures in this area may be sufficient to exert some influence on the lobster populations, and to bring about a southward extension of the oyster populations of neighbouring areas. REFERENCES Bailey, W. B. and Hachey, H. B (1951), An increasing Atlantic influence in Hudson Bay. Proc. Nova Scot. Inst. Set., 12 (4), 17-34. Bailey, W. B. (1953), Seasonal variations in the Bay of Fundy. Fish. Res. Bd., Canada, Manuscript Kept., No. 551. BiGELOW, H. B. (1926), Plankton of the offshore waters of the Gulf of Maine. Bull. U.S. Bur. Fish.. 40 (2), 1-509. BiGELOW, H. B. (1927), Physical oceanography of the Gulf of Maine. Bull. U.S. Bur. Fish., 40 (2), 511-1027. BiGELOW, H. B. and Welsh, W. W. (1925), Fishes of the Gulf of Maine. Bull. U.S. Bur. Fish.. 40 ( 1 ), 1-566. Canada, Fisheries Research Board (1953), Annual Report, Ottawa, 185 pp. Carruthers, J. N., Lawford, A. L., Veley, V. F. C. and Parrish, B. B. ( 1951), Variations in brood- strength in the North Sea haddock in the light of relevant wind conditions. Nature, 168, 317-319. Cooper, L. H. N. (1954), Deep sea oceanography and biological productivity of shallow seas. Repts. and Abstr. of Communications, Assoc. Ocean. Phvs., Union Geod. Geophys. Int., General Assembly, Rome, Sept. 1954, 148 pp. Dickie, L. M. (1955), Fluctuations in abundance of the giant scallop, Placopecten magellanicus (Gmelin) in the Digby area of the Bay of Fundy. J. Fish. Res. Bd., Canada (in press). Dunbar, M. J. (1946), The state of the West Greenland Current up to 1944. J. Fish. Res. Bd., Canada, 6(7), 460-471. Hachey, H. B. (1934), The replacement of Bay of Fundy waters. J. Biol. Bd., Canada, 1 (2), 121-131. Hachey, H. B. (1934), The weather man and coastal fisheries. Trans. Amer. Fish. Soc, 64, 382-389. Hachey, H. B. (1935), The effect of a storm on an inshore area of markedly stratified waters. J. Biol. Bd., Canada, 1 (4). Hachey, H. B. (1953), A winter incursion of slope water on the Scotian Shelf J. Fish. Res. Bd., Canada, 10 (3), 148-153. Huntsman, A. G. (1918), The vertical distribution of certain intertidal animals. Trans. Roy. Soc, Canada, Ser. Ill, 12 (4), 53-60. Huntsman, A. G. (1934), Herring and water movements. James Johnstone Memorial Volume, 81- 96, Univ. Press of Liverpool, 348 pp. Jensen, A. S. and Hansen, P. M. (1931), Investigations on the Greenland Cod (Gadus callarias L.) with an introduction on the history of the Greenland cod fisheries. Cons. Perm. Int. Expl. Mer. Rapp. Proc. Verb., 52, 3-41. Ketchum, B. K. and Keen, D. J. (1953), The exchange of fresh and salt waters m the Bay of Fundy and in Passamaquoddy Bay. J. Fish. Res. Bd., Canada, 10 (3), 97-124. Lauzier, L. (1952), Effect of storms on the water conditions m the Magdalen Shallows. J. Fish. Res. Bd., Canada, 8 (5), 332-339. , ^ . McKenzie, R. a. (1934), Cod and water temperatures. Fish. Res. Bd., Canada, Atlantic Stas., Prog. Repts., No. 12, 3-6. , , ,-r ■ ^ ■ ou .r McLellan, H. J., Lauzier, L., and Bailey, W. B. (1953), The slope water off the Scotian Shelf J. Fish. Res. Bd., Canada, 10 (4), 155-176. ^ ^, ^ „ ,,„, SvERDRUP, H. U., Johnson, M. W. and Fleming, R. H. (1942). The Oceans, Preniicc-Hall Inc., New York, 1087 pp. ^,. ^,„ „.^ y., • Vladykov, V. D. (1933), High temperature stops haddock fishmg. Fi.sh. Res. Bd., Canada, Atlantic Stas., Prog. Repts., No. 7, \0-\l. , . ^ • , ru ^i c Walford, L. a. (1938), The effect of currents on distribuUon and survival of the eggs and larvae of the haddock (Melanogrammiis aeglifinis) on Georges Bank. Bull. U.S. Bur. Fish., 49, 1-73. Papyers in Marine Biology and Oceanography, Suppl. to vol. 3 of Deep-Sea Research, pp. 74-91. Sir C. Wyville Thomson's correspondence on the "Challenger" fishes By Daniel and Mary Merriman Bingham Oceanographic Laboratory, Yale University HISTORICAL ACCOUNT Wyville Thomson's correspondence with Albert Gunther, Keeper of the Depart- ment of Zoology in the British Museum, about the disposition and study of the fishes taken on the H.M.S. Challenger expedition, 1872-1876, and about the publication of results, extended from 1877-1881. In a sense this correspondence centred around the year 1879 — the year in which Henry Bryant Bigelow was born on October 3rd in Boston, Massachusetts.* Now-a-days it is common practice to call oceanography a " new " science. The speed with which it has advanced and, accordingly, with which our knowledge of the oceans has increased, is generally recognized. None the less, it is not altogether easy to maintain perspective and to reahze that "... the famous ' Challenger ' reports, which may be said to form the solid base upon which the superstructure of the science of oceanography has since been built " (Russell and Yonge, 1928), began first to be published only three-quarters of a century ago — in short, during the life-time of the man whom this volume honours. For this reason, it is perhaps worth looking with some care at the year 1879, both in its broad and special aspects, before turning to the hitherto unpublished correspondence which is the subject of this paper. The second half of the 19th century in Europe was marked by many political and economic changes stemming from the Industrial Revolution. As a result of the Franco- Prussian war (1870-1871) there emerged a united Germany and a unification, too, of the Italian States. Russia had effectively pushed the Turks back into Asia, but the western powers resisted her attempts to obtain Constantinople and a command of the eastern Mediterranean. With the Congress of Berlin in 1878 the independence of the Balkan countries, Montenegro, Servia, and Roumania, was established. And the alliance of Germany and Austria-Hungary in 1879 proved most aggravating to Italy, who feared it would block her potential control of the Adriatic ; to France, who saw it as a blow to her ambition to regain Alsace-Lorraine; and to Russia, who still wanted a foothold in the Balkans and the Mediterranean. China posed problems in the Far East, as the western powers and Russia both tried to gain concessions to her great natural resources. In the United States, 1879 was a singularly happy and prosperous year. Crops flourished, manufacturing and trade were stimulated, and the railroads were expanding to meet these demands. We were at peace with the world, except for Indian skirmishes in the west where Sitting Bull had been forced to retreat across the Canadian border. * The October 3rd, 1879, edition of the London Times carried an admonition that there were 12 different Bostons in the U.S.A., and that the name of the state on the envelope address would facilitate accurate delivery. 74 Sir C. Wyville Thomson's correspondence on the "Challenger" fishes 75 England, it is said, was not so happily situated that year. There were wide crop failures and a continuing depression. Ireland, with the loss of her potatoes due to adverse weather, was agitating for Home Rule and land rent reforms; the situation was so acute that the American Irish were sending 510,000 a week towards relief in the homeland. England was fighting in Afghanistan to protect her affairs in India. She was also engaged in military operations in South Africa to keep this portion of her Empire. And she was even sending warships to the west coast of South America, where Chile and Peru were fighting, in order to guard her "... interests in guano for the fields of Yorkshire ". However, England's firm policy of colonization was to stand her in good stead. In addition to India and South Africa, she had acquired control of the Suez Canal, and had long since colonized Australia, New Zealand, Tasmania, and Canada — this, as other countries, notably Spain, had tended to lose their colonial domains. In a different context, Charles Darwin at the age of 70 in 1879,* just twenty years after the publication of The Origin, was doing the experi- mental work at Down which led to the publication in 1881 of The Formation of Vegetable Mould through the Action of Worms, to be received "... with what struck Darw^in as 'laughable enthusiasm'" (Moore, 1955). And the " episco- phagous " Huxley had turned to more peaceful pursuits and was preparing to publish The Crayfish (1880). It was not, after all, such a bad year for England. Perusal of the London Times in the year 1879 yields amusing coincidence with today. The New York Herald was sponsoring a naval vessel's exploration of the North Pole.f The Macon (Georgia) Telegraph reported, and got space in the London Times on January 1, 1879, that there was a remarkable phenomenon on the Florida coast where "... dark, reddish water " was killing fish and creating "... a pestilential stench "; the account went on to state, " We have no other explanation of the poisoning " (reported to have extended as far as 150 miles into the Gulf) " than that it comes from inland waters — the everglades predominantly. ..." Panama had a revolutionary outbreak. The stevedores on the piers of New York were striking. A large underground cave in Algeria was found to contain blind fish.ij: Even the book titles have a familiar ring : we read reviews of " The Sea, its Stirring Story of Adventure, Peril, and Heroism " by F. Whymper. " The Broad, Broad, Ocean ", " Notable Voyages ", and " Episodes of the Sea " are a few others of the spate. And another Mr. Whymper** was bringing out " The Ascent of Matterhorn ". We are also struck by many changes. In 1879 Woolworth opened his first store in Utica, New York (in 1955 the Woolworth Company reported its sales for the previous year at an all time high of $721,312,990). In 1879 Harvard College offered 112 scholarships varying from $40 to $350— today it offers over 1,000 scholarships at an average worth of nearly $700 each. In 1879 the Shah of Persia was planning to undertake a pilgrimage to Meshhed with 10,000 troops, while this year the Shah's * " It was on . . . February 12, 1809 that the other man who along with Charles Darwin was most profoundly to influence their time, and perhaps the future, was born— Abraham Lincoln "' (Moore, 1955). t See for comparison, Walter Sullivan's articles about the Atka expedition to Antarctica in preparation for International Geophysical Year, 1957, in the New York Times. January-March. 1955. ± Some 37 years after the discovery of the famous Kentucky Blind-fish. Amhyhpsis xpclaeus, whose origin the National Geographic Society states in 1955 " . . . is a mystery to naturalists ". ** Edward Whymper, 1840-191 1, English Alpinist and wood engraver, who found a route up the Matterhorn in 1865. 76 Daniel and Mary Merriman travels have carried him to Sun Valley and Florida. In 1955 the production of syn- thetic diamonds has been achieved, finally bearing out the statement in 1879 by Nevil Story Maskelyne of the Mineral Department of the British Museum, "... that (this problem) will be solved, no scientific mind can doubt ". In Boston on December 3, 1879, the Atlantic Monthly gave a party for the seventieth birthday of the " Autocrat of the Breakfast Table " ; today it would be hard to rival the distinction of the gathering which included beside O. W. Holmes these others : H. B. Stowe, J. G. Whittier, H. W. Longfellow, R. W. Emerson, Mark Twain, W. D. How^lls, and J. W. Howe. The February 11th issue of the 1879 London Times carried the statement that, " Arrangements are being settled with the Societe Generale d'Elec- tricite for an experimental lighting of the reading-room of the Museum with the electric light ". And the April 10th issue reported that communication by telephone was established between the Royal Institution and Burlington House, "... with Professor Tyndall at one extremity of the wire and Professor Huxley at the other "; there was apparently much amusement among those present, and, " The feasibility of telephonic communication was clearly demonstrated, the voice being distinctly audible over the whole of a large room ". The London Times for 1 879 also contains numerous articles on science and natural history and makes reference to many familiar names. Thus Frank Buckland* and Spencer Walpole published a report on the Sea Fisheries of England and Wales about which it was written:! On these very surfaces and in these very depths there rages, has ever raged, and ever will to the end of time, a warfare compared with which historical battles sink to the dimensions of street rows or family jars. It is an incessant and universal war carried on between greater numbers than can be told, more species than can be well distinguished one from another, and every order of existence, from the scarcely visible and scarcely animated fibre or molecule up to the lords of creation. . . . Fish engaged in a universal internecine war devour many times more than we can do. Francis Day:]: read a paper to the Linnean Society on the instincts and emotions of fish. Edwin Ray Lankester, founder of the Marine Biological Association five years later, was Public Examiner in Natural Science at Oxford. Edward Cope was appointed head of the department of organic material of the Permanent International Exhibition of Philadelphia. Alexander Agassiz was chief of the scientific staff on the cruise of the George S. Blake through the West Indies. William B. Carpenter resigned as Registrar of London University; this was the man who had helped to induce the Admiralty, through the Council of the Royal Society, to place at his and Wyville Thomson's disposal first H.M.S. Lightning (1868) and then H.M.S. Porcupine (1869 and 1870) for deep-sea exploration (Thomson, 1873), and who was a prime mover with the Government for the Challenger expedition "... to explore and make known the conditions of life in the great oceans " (Herdman, 1923). Huxley received an honorary LL.D. from Cambridge in company with Robert Browning. And at younger levels, William Herdman received his B.Sc. and the * Author of The Natural History of British Fishes, London, 1880. t London Times, December 6, 1879, p. 9. X Author of The Fishes of Great Britain and Ireland, two volumes, London, 1880-1884, and of British and Irish Salmonidae, London, 1887. Sir C. Wyville Thomson's correspondence on the "Challenger" fishes 77 Senior Bursary in anatomy and physiology from Edinburgh, while the Junior Macken- zie Bursary went to one D'Arcy Wentworth Thompson for "... the greatest industry and skill in the particular anatomy work during the winter session ". Francis Maitland Balfour, 1851-1882, had just published his monograph on the Develop- ment of Elasmobranch Fishes* German carp were introduced in southern United States waters, and the previous year the U.S. Fishery Commissioners had "... made a present of a million ova of the California salmon ... to the Government of New Zealand ", about 95% of which were reported to have produced " healthy fish ".f Over 100,000 individuals were employed directly or indirectly in the Scottish fisheries, and the Herring Board stated that: With the exception of the occasional and uncontrollable influences of the weather which cause temporary fluctuations in the catch, the sea fisheries of Scotland and the herring fisheries in particular, are beyond the reach of any power to impair their abundance. The British Association Meetings for 1879 opened in Sheffield on August 20th, and Professor G. J. Allman chose for the subject of his presidential address, " An Account of the Most Generalized Expression of Living Matter ". He made particular reference to the grey gelatinous material which appeared in preserved samples of the deep-sea dredgings made from the Porcupine at depths of 5,000-25,000 feet, and of the fact that it had appeared to be " . , . obviously endowed with life '". He recounted how it had been examined by Huxley, who declared it to consist of protoplasm and envisaged this living slime as extending over wide areas of the sea bottom as a sort of pabulum on which the animals living at these depths fed in the absence of plant life. Huxley had named the material Bathybius haeckelii, and Haeckel had fully supported Huxley's conclusions. Allman went on to state that the reality of Bathybius had not been universally accepted and that the Challenger did not find it. It remained for J. Y. Buchanan, Challenger chemist, to prove that the material was an inorganic precipitate owing to the action of the preserving fluid, alcohol. Huxley, in thanking Allman at the conclusion of his address, admitted that he had christened Bathybius. " He had hoped, indeed, that his young friend Bathybius would turn out a credit to him, but he was sorry to say as time had gone on Bathybius had not verified the promise of his youth " (London Times, August 21, 1879). This was the great early period of oceanography. Wyville Thomson's cruises on the Porcupine and the Lightning in the North Atlantic had destroyed Edward Forbes' conception of the azoic zone. In 1870 Thomson, now forty, and having spent seven- teen years teaching in Ireland, succeeded Allman in the Chair of Natural History at Edinburgh. The Challenger expedition followed, with Thomson as director of the civilian scientific staff on board. As Murray (1895) reports, " After circumnavigating * Macmillan and Co., London, 1878; reprint of papers in \hQ Journal of Anatomy ami Physiology, 1876-1878. The present-day classics on Elasmobranchs are by Bicelow and ScHROEDtR (1948 and 1953). t At the same time the question of introducing California salmon into British waters was the subject of much debate in England. One writer, Sir Rose Price, emphasized the "... extremely risks nature of the experiment ", and claimed these fish would not take a fly and had no flavour. He concluded by stating, "The mortality among salmon in California is simply incredible" (London Times, April 16, 1879). 78 Daniel and Mary Merriman i the globe, and carrying on deep-sea and other investigations in many regions of the ocean, the Challenger returned to England in May 1876, and the crew was paid off after the ship had been in commission for over three years and seven months ". Then came the question of how best to work up and publish the results. Quantities of data in all branches of oceanography had been recorded, and the collections, which had been sent back to England from the expedition's different ports of call or brought back on ship-board, were wonderful in their extent and variety. There was extensive correspondence, in which members of the Royal Society, the Admiralty, the British Museum, and the Treasury all had a hand. Murray {he. cit.) again writes, " It was further determined that the records of the various observations and marine collections should remain in the meantime in the hands of those who had taken part in the Expedition, and that a temporary Government department, with a small annual grant, should be created, the duty of which should be to direct the discussion of the physical and biological observations, the examination of the collections, and the publication of the scientific results, so far as these had a bearing on the science o Oceanography ". Her Majesty's Stationery Office was to publish the results, and the " typical collections " were eventually to be deposited in the British Museum. In 1877 Wyville Thomson was appointed Director of the Challenger Expedition Com- mission, with headquarters in Edinburgh and John Murray as first assistant. Wyville Thomson fell into ill health shortly thereafter, and he lived only five more years — to the age of 52. In that time he settled the style of the publications and sent " a consider- able part " of the collections to the specialists* who were to examine and describe them. In fact, by 1882, the year of his death, 22 of the Challenger Memoirs were in print, the first having appeared in 1880. It remained for Murray to see the job through; the last of the " Fifty large Royal Quarto Volumes " appeared in 1895. Thomson's letters to GuNXHERf resulted in three memoirs: on the shore fishes (1880), on the deep-sea fishes (1887), and on the pelagic fishes (1889). The corre- spondence, not all included here and in a most unhappy long-hand, provides fair evidence of the tremendous pains to which Sir Wyville went, of his attention to detail, of his ability to prod the contributors to the Challenger Reports when occasion demanded, and of his insatiable desire to see the whole series done to the highest degree of perfection. University — Edinburgh March 17, 1877 Dear Dr. Giinther: We have now gone over the greater part of the " Challenger " spirit collections and the Fishes are nearly ready to be handed over to you if you are inclined to take them up. There are two distinct sets — those from the shallow water and from the marshes. This col- lection is not large as such a collection might easily be made with more time but I have no doubt there are many undescribed forms from the more remote places. The other set is from deep water and many consist of a couple of hundred specimens (more or less) of forty or fifty species of which a large proportion are undescribed. They are mostly allied to the deep-water things which have come home from about Madeira. * The completed Challenger Reports contain contributions from 76 authors. t Published with the permission of the Trustees of the British Museum. We here express our best thanks to the several persons who helped us with the original letters of C. Wyville Thomson in the B.M.N. H., particularly Mr. Mugford of the Mineralogical Department Library. Sir C. Wyville Thomson's correspondence on the "Challenger" fishes 79 If you undertake the Fishes the arrangement approved by the Treasury and by the Trustees of the British Museum is the following. " That the fishes be sent to Dr. Giinther for determination and description and that Dr. Gunther be requested to select a complete set for the British Museum including all unique specimens and two specimens of all species of which there arc more than three; the remaining duplicates to be returned to me for distribution with the sanction of their Lordships." There is one point with regard to the deep-sea fishes especially which I must mention. Mr. Murray has had charge of these and has devoted special attention to the circumstances under which they have occurred making careful notes in each case. No description would be complete particularly in its bearings on physical geography without such information and 1 think it would be very desirable that Mr. Murray should be associated with you in the description of this section; this is a matter, however, which I must leave in your hands. I should wish all new species and all species which have not already been well figured in readily accessible publications to be fully illustrated with any necessary anatomical details. Of course I am prepared to defray the expense of illustration. I should like, if possible, to have some at all events of the plates done during the next financial year and I would be greatly obliged to you if you would, when you see the specimens, give me a rough estimate of the number of plates which will be required and of the approximate costs. In the meantime, will you kindly let me know your views generally on the matter and when you would wish the Fishes sent. I enclose a proof of a list of observing Stations and will send a corrected copy with the chart shortly. I send a rough proof of one of the plates to give the size. Yours faithfully, C. Wyville Thomson Bonsyde Linlithgow, N.B* July 10, 1877 Dear Sir: I do not think that there is the slightest objection to your publishing the Kerguelen Fishes in either way you prefer — Annals or Phil. Trans. The deep-sea series I should think of course to form a part — or volume if need be — of the official report. Would you be good enough to give me so far as you can a rough estimate of the amount which you expect to be able to undertake — plates and letter-press, during the current financial year, and the total expense for this year— the plates on the stone and the letter-press ready to go into the printer's hands. I am glad that you are finding so many new and interesting forms. I think you may depend upon the condition of the specimens being good, for certainly no care has been spared either in that or any other group of marine forms. 1 am yrs faithfully, C. WvviLLE Thomson University of Edinburgh 18 October, 1877 Dear Sir: I now enclose a plate as a sample of size and style that we may have the whole series as nearly uniform as possible. The tinting, the additional expense for which has been sanctioned by the Stationery Office, gives a great advantage in brightening of the figures by the use of white. The exact size of the tinted portion of the plates, well within which the figures must be kept, is 10 X 8 inches. It would be well to keep the lettering the same as on the sample plate, altering * Thomson's ancestral country home, where he was born and where he died The centre window of the aose of the parish church of St. Michael of Linlithgow was done m 1SS5 in memor>^of Sir WYvfLL? the ubject s a representation of the 104th Psalm. God's mamtcstation ot Hiniself mthe Works of Creation, and in the lower parts of the window are illustrated the great and wide sea . 80 Daniel and Mary Merriman the legend on the right hand upper comer as required, e.g., Foraminifera PI. Ill Ostracoda PI. IV. I should like that two proofs of each plate should be sent to me before the requisite number of impressions, which will probably be 525 are printed off. The paper will be supplied by the Stationery Office. I am, sir. Yours faithfully, C. Wyville Thomson* Edinburgh November n, 1877 Dear Dr. Giinther: I have written to Spence Bate, t Under all the circumstances it may perhaps be as well to allow the Brachyura to stand over for the present. Will you kindly let me know for the information of the Stationery Office how you propose to have your plates lithographed — through what firm, and at what cost — the plates to be of the same size as the pattern plate sent and the number of copies probably about 730. The Stationery Office supplies the paper, so it is merely the use of the stones and the printing. Thank you for your interesting account of our Japan species. What a number of new forms there seems to be. Believe me. Yrs. faithfully, C. Wyville Thomson Edinburgh May A, 1878 Dear Dr. Giinther: I should think that there cannot be the least doubt of the advantage of the course you propose. The abstract of the description in the Annals should be " published with the permission of the Lords Commissioners of the Treasury ". I have heard nothing about fishes from Agassiz. He has got a fine haul of invertebrates. Let me know what arrangement you make when you begin the plates so that we may be all square with the Stationery Office. Yours faithfully, C. Wyville Thomson University Edinburgh July 5, 1878 Dear Dr. Giinther: Thank you for the signed list. It keeps things all square. I am very glad to know that the illustrations of the Fishes are going on. I am glad also that the condition of the Fishes pleases you. Murray took a great deal of trouble. Yrs. faithfully, C. Wyville Thomson * This letter is not in Thomson's own hand, though the signature is clearly his. t See Challenger Report on the Crustacea Macrura, Vol. 24 (Zool.), Pt. Lll, 1 Sir C. Wyville Thomson's correspondence on the "Challenger" fishes 81 EdinhtiiKh ^ r^ ^. .u Octubcr 12, 1878 Dear Dr. Gunther: 1 am delighted to see the plates. They seem to be excellent. We have a contract with Minlcrn's so all you have to do is to order from him is to order {sic) 750 copies of each plate on the terms of his contract for the Challenger work. 1 think the best plan would be to leave all the plates at Mintern's except 20 copies of each which are allowed for editorial purposes, and which I should like to have sent to me. I do not care to have all the plates stored in one place. It concentrates the risk too much. I suppose Mintern's people have a Hre-proof store. The faster the plates come in now the better. I fear it will be impossible to complete the whole work within the time I gave* — but — we are doing our best. 1 send a copy of Lyman's synopsis of the first division of the Ophiuroids so you see what you have to expect in the way of new forms. 1 have not yet got the type specimens of the Echinids but most of the plates are done so you will get them soon. Believe me Yrs. faithfully, C. Wyville Thomson Edin. Decembers], IS78 Dear Dr. Gunther: Of the enclosed documents those marked I and II were returned to me from the Treasury on the grounds that Plates 3 and 6 had been charged twice. I had signed both accounts supposing that the double marks referred to the two series — shore and deep-water fishes. Mintern's people seem to be careless in these matters. If this is the second time a mistake has occurred, would you mind asking them to send their accounts to you and initialing them if you find them correct? Yrs. in this C. Wyville Thomson Edinburgh January 25, 1879 Dear Dr. Giinther: Will you very kindly try and fill up the accompanying sheet. 1 am anxious to make out as clearly as possible where I am and what I may expect. Yrs. faithfully, C. Wyville Thomson Bonsyde Linlithgow, N.B. Nov. 12, 1879 Dear Dr. Giinther: I am now going on with your report and I will send you some proofs in a few days. It is somewhat different in form from the others. I thought the plates had been printed oflT. I suppose Mintern had better send to me a demand for the paper they require and I will send it up to the Stationery Office. Will you kindly let me know to what the numbers attached to the species of shore-fishes refer. * Murray (1895) writes, " In the year 1889 Her Majesty's Treasury declined to ask Parliament to renew the annual grant for the continuation of the work relating to the scientific results of the Expedition the time estimated for the completion of the publications havmg expired. However, after some correspondence, in which I offered to finish the Report at my own expense, the Govern- ment agreed to set apart the sum of sixteen hundred pounds for the completion ot the ofiicial publica- tions in the same style as that in which they had hitherto appeared. This sum has been the only pay- ment from Government funds in connection with the Challenger Expedition during the past six years. . . " 32 Daniel and Mary Merriman As to the pelagic fishes if you think they would be better on woodcuts I have no objection. If you can employ Mr. Cooper 188 Strand to cut the blocks it would be convenient as we shall have an account with him at all events. I shall be very glad to see the deep-sea fishes. Of course in this report the interest centres on the distribution and the bathymetrical range of the forms, and I should like it to be as complete in this respect as possible so as to form a basis for future work. I would be glad if you would kindly give very full lists of the deep-sea fishes which have been described hitherto, and if you would cause to be figured anew any species taken by us, which have been only described and not figured, which have been badly figured, or which have been figured in not easily accessible books. You are no doubt aware that Agassiz's deep-sea fishes are in Steindachner's hands and now well advanced. As the abyssal fauna is very uniform many of our species are in that series. I have heard from several of your London acquaintances that you are in some way dissatisfied with the form in which you are receiving the British Museum Series of specimens. I do not know what the cause of complaint is, but if there be any it would be better perhaps to refer it to myself. I send a proof which happens to be lying before me to show you the form which the reports are taking. Yrs. faithfully, C. Wyville Thomson Bonsyde Linlithgow, N.B. Nov. 17, 1879 Dear Dr. Giinther: Thank you so much for your note. You have not however told me to what the nos. attached to the Shore-fishes in your list refer. Please let me have a post-card as soon as you can. I assure you if I had only heard of your supposed dissatisfaction casually or from one source I should have said nothing about it. Since, however, there seems to have been some mistake there is no use in taking further notice of the matter. I consider it perhaps the principal part of my duty in connection with the working up of the " Challenger " collection to place the type specimens and whatever else seems necessary for full illustration, of every species, in the British Museum. And this 1 will carry out to the best of my power. As I told you at the time the specimens which you got from Agassiz belong to known species, and to a few new species of which many samples were found — possibly some of them are unnamed, and I sent them to you just as they came as I am aware that each transfer however carefully managed does some little damage, and I did so mainly that some pretty things such as Coelopleurus, Salenia, Asthenosoma, etc., might be seen at once.* You will get the type speci- mens whenever they are figured and described. Thank you for your kind reference to my late illness. I fairly broke down with over-work, but I am now nearly as well as ever. Believe me Yrs. faithfully, C. Wyville Thomson Bonsyde Linlithgow, N.B. Feby \3th, 1880 Dear Dr. Giinther: I send you at last a proof of your report on the Shore-fishes. I thought you would have had it long ago but some other things occupied us very fully for a time. As it turns out however this delay has been of no consequence for it will be a month yet it seems before your plates are printed off. '"^ ^ What I send you, then, is a first-proof, corrected as far as we could manage it, but still needing * See Challenger Report by A. Agassiz, Vol. 3 (Zool.), Pt. IX, 1881. Sir C. Wyville Thomson's correspondence on the "Challenger" fishes 83 to be put into shape in certain respects. The Geographical list* I propose should come first— as an elaborate Table of contents. Then the descriptions and the systematic list at the end. There are several mistakes and among them a few which ought to be corrected. For example the fresh- water fishes from the Mary river were not presented to us but caught with some labour by a little party consisting of Murray, Lieut. Aldrich, and myself, who squatted on the bank of the river for a fortnight in the hopes of getting young Ceratoclus. We took two of the mature Cera- todus however and it should have been in your list. The specimens were not sent to you as ihey were not put up with the rest of the fishes. I know you had plates of it so I suppose you do not wish them. If you do you can have one. I should like to keep the other here as a memento of a pleasant holiday trip. To save you trouble it would be better perhaps that I should add when necessary a note indicating anything special about the fishes— such as the mode of preserving them at different places. The account is rather bald without a few such details, t Would it not be as well to substitute some other specific name for Sancii pauli.% The French are just describing the fauna of the other St. Pauls which is so much better known. In the introduction of your preliminary notes on the Deep-sea fishes you make some remarks as to the extent and condition of the collection of fishes. I suppose you have no objection to these remarks being repeated here. I send you two copies of the proof. If you want additional copies please send me a line- also if it is necessary to send the Mss. I retain a copy and will add such additional notes as I think are necessary, and then, when I get your corrected copy, 1 will have a clean proof drawn and send it to you before printing off, in case you have any further alterations. I will be glad of the deep-sea fishes whenever I can get them. The first spurt is about over and we shall need as much material as we can get to go on with. You will get the type collection of the Echini very shortly now. I hear from Agassiz that he is very nearly done. I hope to be able to be in London in March but I am not quite strong yet and am shirking the journey as long as possible. Yrs. faithfully, C. Wyville Thomson Bonsyde Linlithgow, N.B. February lA, 1880 Dear Dr. Giinther: I am afraid it was trespassing altogether too much upon your time to ask you to take charge of the Report on the Challenger Fishes, but I followed the principles I tried to work on as far as I could, and applied to the most distinguished Ichthyologist I knew. I must ask you however to allow me to bring your list as nearly into the form which has been adopted after much consideration for the report, as its nature will allow. I am taking the utmost care that the type collection of everything goes to the British Museum in its thorough completeness but I do not mean to make the Challenger Report a Museum Cata- logue in any sense. The data I mean to publish are those which have reference to the Expedition. No doubt all the letters and references to specimens in and out of the Museum will appear in your own catalogues when these Fishes are added to them. Such a list as you send me should be published by the Museum if it is required. I suppose that the Shore-Fishes are not of much importance but I do not wish to publish matter which from my point of view is wholly irrelevant, and I must ;idd a few notes about the * No geographical list as such appears in the printed version; the descriptions arc made under broad geographic headings, Atlantic, Temperate Zone of the South Pacific, etc. t To make the account of Ceratodus less " bald ", Sir Wvvilli , who evidently felt strongly about these fish, wrote what must surely be one of the longer footnotes in ichthyology— almost a thousand words (GiJNTHER, 1880). X GUNTHER apparently disregarded Thomson's suggestion, for Holoccnirum sancti pauli. n.sp. appears on page 4 of his Report on the Shore Fishes (1880). 84 Damel and Mary Merriman Ceratodus and means by which the fishes were procured — to do my duty fairly as Editor. I have not time for many, but what are added you will see before they are printed off. The report on the Shore-Fishes might be as good as that on the birds which will give all the information we have. I will send you tomorrow the Mss. — and you will see that we have taken no little trouble correcting spelling, adding authorities, etc., etc. I would have certainly left out that long Geographical list altogether. It seems little more than a repetition of the main list. I will leave it out yet if you have no objection for I do not think it improves the appearance of the paper. I have not the proof at hand — it is at the Office in Town but I will see it tomorrow and write again what I think had best be done. Yrs faithfully, C. Wyville Thomson Bonsyde Linlithgow, N.B. March 10, 1880 Dear Dr. Giinther: I daresay your plates are now nearly finished and the rest of the volume is ready. All you need to do is to correct any errors in spelling and so on in the text, and I would like to have it as soon as convenient. Yrs. faithfully, C. Wyville Thomson Bonsyde Linlithgow, N.B. March 22, 1880 Dear Dr. Giinther: I must apologize for having entirely forgotten to send Ceratodus. I suppose you have been waiting for it before sending the corrected proof. I will be glad of the proof whenever you are ready for we are anxious to set the type free. There is another little matter which I ought to mention. I see you have communicated a paper to the Linnaean Society on some of the deep-sea fishes. If any report or abstract of that paper is published I would be greatly obliged if you would add to the heading " Published by permission of the Lord Commissioners of the Treasury ". Of course in making this request I am only obeying my own instructions from the Treasury. Yrs faithfully, C. Wyville Thomson 29 March 1880 Dear Sir Wyville:* The specimen of Ceratodus arrived safely on Saturday last, and I return it today with the proof sheets. The proofs are corrected with the exception of 1 . The Geographical List which you propose to leave out. I see no objection to it and only regret that it was set up in type, time and expense being thereby saved. 2. Of the enumeration of the specimens of each species which I have left as you have had it set up. With regard to my discourse at the Linnean Society, it referred to results of examinations made by myself long ago or others more recently independently of the Challenger Collections. * This unsigned letter, much emended, appears to be the first draft of Gunther's answer to Thomson's of March 22, 1880. Sir C. Wyville Thomson's correspondence on the "Challenger" fishes 85 Bonsydc Linlithi^'OM, N.B. April 1. IS SO Dear Dr. Gunther: Many thanks for the corrected proofs. They will be put in hand in"rmcdiately and I uill send you a second proof before the paper is printed off. I see most of your corrections refer to matters oi form rather than Ichthyology. I do not suppose you attach much importance to the relative positions of n and sp! Our way of putting it was adopted after an amount of consideration sufficient for the subject, I think. It means either " new species " or " nova species " as you choose. On what ground it should be inverted, unless it came in as part of a Latin sentence, I am not aware. The single i terminating proper specific names ending in a consonant is in accord- ance with the " Strickland Code " and is I believe correct. The first Zoological volume will now be out at once. I have not yet got your plates from Mintern's however — but I suppose they are ready. I suppose you will have no ditficulty in arranging the account of the deep-sea fishes zoologically and putting the report into the same form as the others. It will save a deal of trouble. The sooner I can get the reports, the materials for which have been long in the hands of the authors, into print the better — for there will be a great accumulation towards the end. Yrs faithfully, C. Wyville Thomson Bonsyde Linlithgow. S.B. May A, 1880 Dear Dr. GUnther: I hope this will answer now. I have, as you suggested, taken out the reference to the B.M. Catalogue, and have made the Report more comparable in shape of the rest. I trust you will not find it to require much more correction— and that you will be able to let us have this Mss. proof at once, for the succeeding volume is almost finished and (we) are greatly pressed to put the first out of hand. r ..r n ^ Yrs faithfully, C. Wyville Thomson Bonsyde Linlithgow, N.B. May 6. 1 880 Dear Dr. Gunther: I think the addition of the paging to the systematic list would be a great improvement. I would have had it done here but our hands are very full. Delighted to hear that we shall soon see some deep-sea fishes. ^^ fai,h,-u||y C. Wyville Thomson Bonsyde Linlithgow. S.B. July 5, 1880 Dear Dr. Gunther: I was unlucky in missing you the last time I was in London. ^r .u. „, , .. You wrote me some time ago that I might expect w.thm a very short fmc some of the pLtes of the deep-sea fishes. Could you drop me a line when you thmk these will be ready. I wish to arrange for the next set of volumes and I want to know when your memoir will come m-as '"h^^^u u::^ci:^dr;ncorporaced or done anything with the set of Echinodern. . sent you^ I find that they were sent in a certain sense by mistake, and I could select your full set much 86 • Daniel and Mary Merriman more satisfactorily if I had them back again. I have had no official receipt for them. They were only meant in a provisional lot in case anyone wished to see them and if they are not exhibited or entered if you wish to send them back to me I will send you the type lot complete. Yours faithfully, C. Wyville Thomson *I am sorry that there has been any mistake about the preliminary set of Echinids; but, re- reading from letters, I must acquit myself of any share in the error. The specimens have not only been put into different bottles, but have been registered, incorporated and reported to the Trustees; and by them to the House of Commons. It is now, therefore, impossible for me to acquiesce in your request that they should be returned. It seems to me that the best thing to do will be to send me the type-series, when I will have them compared with what have been already sent, and, if I find that it is possible to return any of the freshly sent specimens, or finding that the characters and distribution of the species collected is already well enough represented by the previous series, I will certainly do so, in order that they may form a part of a good set for the museum, which has the second claim on this National collection. So far as the specimens already sent are concerned, I may add that the assistant in my Depart- ment who is especially charged with the care of this group has engaged himself to remain in London until the end of August, and I may safely promise for him that he will spare no trouble in giving all assistance in his power to the artist (2) or the describer (1), if they are desirous of having another examination of the specimens already in the British Museum. Bonsyde Linlithgow, N.B. July 10, 1880 Dear Dr. Giinther : I am greatly obliged to you for the proofs which are already in the printer's hands. Yrs faithfully, C. Wyville Thomson Bonsyde Linlithgow, N.B. July 15, 1880 Dear Dr. Giinther: Thank you so much for your note and for the first deep-sea fish plates. I suppose in these plates you will simply put the name of the Fish beneath as we have done in the other groups without locality or anything further. Either the generic name only or the generic and specific as you think proper. It is all right about the Echinoderms. I thought it very possible you had incorporated them. I daresay you could without much trouble send me a list of the species already sent. I think I kept a duplicate list but I have changed my Secretary since, and I cannot at this moment lay my hands on it. I want to be sure that you get every species the old as well as the (sic) those not previously described. Yrs faithfully C. Wyville Thomson P.S. Can you readily send me by return the date when you got the shore-fishes and the date when you returned me the Mss ? It has been suggested to me that these should be published in all cases and I did not think of it at the time. C.W.T. * This letter, unsigned and undated, but in the same hand as that of March 29, 1880, is apparently a first draft, Gunther to Thomson. Sir C. Wyville Thomson's correspondence on the "Challenger"' fishes ' 87 Bonsyde Linlithgow, N.B. October 1, 1880 Dear Sir: I have forwarded your note to Dr. Sclater* and see no reason why you should not enter the birds in your forthcoming volumes. The Pteropods are not even commenced. They are not so numerous as might be expected in the Collection and, as they are greatly scattered on slides and in bottles of tow-net matter, it will be some time before they are ready. We must get through with the hii^ger things first. You will get probably tomorrow or next day, a lot of fishes from the deep water off the Faroes ( ?) some of the spoils of the Knight-Errant.] 1 will be greatly obliged to you if you will simply add them to the Challenger things ...(?) That is look them over keep what you require for the Brit: Mus: and return us the remainder named. Only it would be a great favour if you would keep this lot separate, send us a specimen of all of those of which there are two in the same condition, and let me have a separate short report on them for a paper on the Faroe Channel which I am going to read at the R.S.E. You will get lots of things in other departments by degrees from the same cruise, but I am having them all named and worked up with the Challenger things in the mean time. Yrs faithfully, C. Wyville Thomson Bonsyde Linlithgow, \'.B. October 25, ISm Dear Dr. Giinther: Dr. Sclater promises to send you the birds as soon as possible after his return home. I am looking out most anxiously for the plates of the deep-sea fishes. They are now pressing me to finish my work on the Collections and leaving such a crush of printing for the end that I hardly see how I can manage it. I would prefer having, as in other Memoirs, the name of the Fish only on the plate. 1 would especially rather not have the depth— for although of course the depth of the sounding is given in the station for the Fish, we can seldom be absolutely sure that the fish actually came from that depth— particularly in the case of using the trawl. faithfull C. Wyville Thomson ±The bathybial Fish-fauna which surrounds the British Islands was hitherto almost unknoNsn. Beside the stray specimens which now and then were found thrown ashore or floatmg on the surface no further evidence of the existence of this fauna was obtained, except on two occasions, viz on a dredging-excursion of Dr. Gwyn-Jeffreys in 1867 from a depth of from SO to W fathoms;* and during the cruise of H.M.S. Porcupine in 1869 from a depth ot from 200 to 500 fathoms, t *See Ann. & Mag. Nat. Hist. 1867, xx, p. 287. t/W.. 1874. XML p. 138. * See Challenger Report by P. L. Sclater, Vol. 2 (Zool.), Pt. VIII. 1881. t The cruise which led to the deliniation of Wyville Thomson ridge: see Herdman. p. 5!^. 1923. J A^*^A or>H miirh emended is in the same hand as that of March 29 and .he^T.eiJjSw^col'-o-e t P. itZt aTpTremiy ano.her fi., draf. GO,.„h« .o THO^■so^. 88 Daniel and Mary Merriman Neither of these two contributions can compare as regards interest and number of specimens with the series obtained during the cruise of the " Knight-errant " ; and it would seem as if now only the rich spoil which I ventured to indicate in 1 867 as resulting from an exploration of the Deep Sea round the British Islands, were being gathered. Six out of the ten species obtained^ are new to the British Fauna ; and of course represent but a small fraction of the actual number of Brit, deep-sea fishes. Much, therefore, remains to be done. The laws which govern the bathymetrical distribution of Fishes, are still obscure ; and it is evident that a series of continued methodical observations, such as can be made in a limited oceanic district like that round the Brit. Islds., whose hydrographic conditions with its surface and coast fauna are so well known, is most likely to reveal a chain of facts which cannot be recognized in disjointed observations made at distant localities. Besides, there are not a few obscure points in the life-history of our food- fishes which may be well expected to be cleared up by the deep-sea-dredge, such as the unaccount- able disappearance from certain parts of the coast of fishes like the Haddock, the change of habitat of many fishes according to the season, a change which evidently much more frequently takes place in a vertical than horizontal direction, etc. It is therefore to be hoped that the present successful expedition will be followed by equally well conducted efforts. The collection submitted to my examination contains a much greater proportion of arctic forms, than of southern; and in this respect differs entirely from that made by Mr. Gwynn" Jeffreys at a less depth. The only southern form is Haloporphynus lepidion which we knew previously from the Mediterranean and Japan. Singularly, again, no trace of a Trachypterus or Regalecus was obtained; and we can account for their absence only by the supposition that it is difficult to enclose these long snake-like fishes in the dredge, and that young specimens from their extreme delicacy of structure are probably torn into fragments or lost long before the net reaches the surface. Some of the species have been previously obtained by the Scandinavian Expeditions in similar latitudes. As all the species will be fully referred to or described in my Report on the *' Challenger " Deep-sea-fishes, only a few notes on them are appended here. Bonsyde Linlithgow, N.B. Oct. 26, 1880 Dear Dr. Giinther: I had written you a note just the post before I got yrs. I am very much obliged to you for the report and greatly pleased that the fishes have interested you so much. I have been long looking forward to a careful overhaul of the Faroe Channel, and I have every hope that we may have another investigation this next summer under more favourable conditions. So far as this years work is concerned I am at liberty to ask you to make what use you choose of the duplicity which I have much pleasure in doing ( ?) — only send me back what you can spare Yrs faithfully, C. Wyville Thomson Nov. \st (?) — no address Dear Dr. Giinther: I have just received a list signed by you from Mr. Moseley. You will exercise your own discretion in selecting the Brit : Mus : set. Of course my prime object is to make that as complete as possible, but, that done, it would be a convenience for (me) to have as many species here for comparison as I can get. I have a note from Dr. Sclater that he has handed over or is about to do so, the birds. I suppose you will send me a list. I expect to send off the Pennatulida the end of the week. For some whale bones and seal bones you will have to wait till the part on the bottom deposit is finished. I am anxiously looking for the deep-sea fishes. This is a most laborious job ! Yours faithfully C. Wyville Thomson Sir C. Wyville Thomson's correspondence on the "Challenger" fishes 89 Bonsydi' Linlithgow ^ ^ ^.. ^ Nov. 2, 1880 Dear Dr. Gunther: From having heard nothing from you with regard to the Corals of the Challenger Expedition I suppose I am right in concluding that you have not received them yet from Mr. Moselcy. As some difficulties have arisen in this department I have asked Mr. Moseley, to avoid any further complications, to send the whole collection, the type specimens, the second selected set. and the duplicates to you. I will be very much obliged to you if you will kindly select the first set accord- ing to the instructions sanctioned by the Treasury— and return the rest to me. I am sorry to give you this trouble— but this is I think the only case. I will send you in the course of a few days the Ostracoda and the Pennatulida. I mean in all cases to send the specimens to the Museum as soon as possible after the Memoirs arc published. It would be scarcelv fair to do so much before. I have been reading your book on Fishes with much pleasure, and I think I know more about them then I did before. It is a resume which was much wanted. Yrs. faithfully C. Wyville Thomson Bonsycle Linlithgow. N.B. December 17. 1880 Dear Dr. GiJnther: I know pretty well all about the advantages of complete collections for reference. I am only very glad that so much attention is now being paid to these minute ( ?) groups in the B.M. My great object has been to make the collection from the " Challenger " in all branches in the National Collection as perfect as possible. Beyond a certain point I cannot force this but I will do the best I can. Believe me yrs faithfully C. Wyville Thomson P.S. I will send you today the whole of the remaining material returned by Dr. Brady as duplicates. I forwarded your letter and list to Dr. Brady.* Bonsyde Linlithgow, N.B. January 16. 1881 Dear Dr. Giinther: It has been suggested to me that perhaps I ought to have let you know that it was in my power to offer a moderate honorarium for literary work in connection with the Challenger Report. To tell the truth I had some delicacy in doing so remembering the strong representation which the Brit. Mus. officers made on that matter to government. I do not see howe\cr that preparing such a report, especially as I distinctly objected to its being in the form of a Brit. Mus. catalogue, could be considered a part of your regular work. If you desire it I will send in an account for the sum to which you are entitled under my instructions. Will you very kindly let me know how the Deep-sea Fishes stand. Yours sincerel> C. WVVILLK THOMSt)N * Either G Stewardson Brady on the Copepoda (Challenger Report Vol. « (Zool.»- P' \\' • 1884) or Henry BOWMAN Brady on the Foraminifera (Challen.^er Report. Vol 9 (Zool.). Pt. WII. 1884). 90 Daniel and Mary Merriman Bonsyde Linlithgow, N.B. February 11, 1881 Dear Dr. Gunther: I am extremely sorry to hear that you are still on the sick list. I hope you will shortly be in condition to resume your work with comfort again. As to future arrangements I will be glad to meet your wishes in every way, as far as I can. Yrs faithfully C. Wyville Thomson Bonsyde Linlithgow, N.B. March 1, 1881 Dear Dr. Gunther : Let me introduce to you my Secretary Dr. W. A. Herdman, F.Z.S. — from whom I think you have heard from time to time. I will be very much obliged to you if you will let him overhaul your Ascidians. He is doing ours and I know that he is thoroughly up to them. If you can help him in any way you will do me a great favour. Also on all " Challenger " matters talk to him as to myself. I sincerely hope that you are now all right again. I hope to be in Town in about a fortnight or so. Yrs faithfully C. Wyville Thomson Bonsyde Linlithgow December A, 1881 Dear Dr. Giinther: I am getting very anxious about your paper on the deep-sea Fishes and would be very glad to see some of the work. Both the Royal Society and the Treasury are expressing some impatience and I may be landed in difficulties if some of the promised memoirs are much longer delayed. I would be greatly obliged to you also if you would send me a receipt for the Echinoidea sent by Agassiz according to his Memoir. There are several other Memoirs which will be ready for delivery shortly. Believe me Yrs faithfully C. Wyville Thomson Murray (1895) speaks of the fact that the Challenger Reports cover "... about twenty-nine thousand five hundred pages, illustrated by over three thousand litho- graphic plates, copper plates, charts, maps, and diagrams, together with a very large number of wood-cuts ". He goes on to say, " From beginning to end the history of the Challenger Expedition is simply a record of continuous and diUgent work ". In a sense it is just that. But who, most of all, had the perspective and pertinacity to initiate this first real study in deep-sea research ? If Thomson were alive today, it is fair to speculate that he would be astounded at the developments in oceanography since 1879. His was an unusually broad, inquiring mind — to which his writings and editing testify abundantly; his was the imagination that resulted in the Challenger Expedition; and his was the guiding hand that led to the foundation of the modern science of oceanography. In mute testimony, his name appears on the title page of Sir C. Wyville Thomson's correspondence on the "Challenger" fishes 91 all Challenger volumes— whether produced before or after his death -" Prepared under the Superintendence of (the late) Sir C. Wyville Thomson, Knt., F.R.S., &c. Regius Professor of Natural History in the University of Edinburgh, Director of the Civilian Scientific Staff on Board". REFERENCES BiGELOW, H. B. and Schroeder, W. C. (1948), Fishes of the Western North Atlantic. Pt. I. Sharks. Sears Foundation for Mar. Res., Mem. I, 59-576. BiGELOW, H. B. and Schroeder, W. C. (1953), Fishes of the Western North Atlantic. Pt. 2. Saw- fishes, Guitarfishes, Skates and Rays. Chimaeroids. Sears Foundation for .Mar. Res., Mem. I, 1-588. GiJNTHER, Albert (1880), Report on the Shore Fishes procured during the Voyage of H.M.S. Challenger in the Years 1873-1876. Report on the Scientific Results of the Vovage of H.M.S. Challenger, etc., Zool., I, 1-82. GiJNTHER, Albert (1887), Report on the Deep-Sea Fishes collected by H.M.S. Challenger during the Years 1873-1876. Report on the Scientific Results of the Vovage of H.M.S. Challem^er. etc.. Zool., XXII, I-LXV and 1-335. GiJNTHER, Albert (1889), Report on the Pelagic Fishes collected by H.M.S. Challenger during the Years 1873-1876. Report on the Scientific Results of the Vovage of H.M.S. Challenger, etc., Zool., XXXI, 1-^7. Herdman, W. a. (1923), Founders of Oceanography and their Work. Edward .Arnold & Co., London, 340 pp. Moore, Ruth (1955), Charles Darwin. Alfred A. Knopf, New York, 206 pp. Murray, John (1895), A Summary of the Scientific Results. Editorial Notes. Report on the Scientific Results of the Vovage of H.M.S. Challenger, etc., VII-XII. Russell, F. S. and Yonge, C. M. (1928), The Seas. Frederick Warne & Co. Ltd., London and New York, 379 pp. Thomson, C. W. (1873), The Depths of the Sea. Macmillan and Co., London. 527 pp. Papers in Marine Biology and Oceanography, Suppl. to vol. 3 of Deep-Sea Research, pp. 92-109. The hydrography and the distribution of chaetognaths over the continental shelf off North Carolina By Dean F. BuMPUs*t Woods Hole Oceanographic Institution and E. Lowe Pierce Department of Biology, University of Florida Summary — Temperature, salinity and quantitative plankton data have been obtained from the continental shelf area and Florida Current off North Carolina in May and June 1953 and January 1954. Two water types, Virginian Coastal water and Carolinian Coastal water, and one water mass, Florida Current water, are identified. A breaching of the barrier at Hatteras between the two coastal water types was witnessed, and Virginian Coastal water was driven south-westerly across Diamond Shoals into Raleigh Bay by a north-east storm. The import of such an hydrographic event on the distribution of plankton is discussed. The distribution of the chaetognaths in the area was investigated and their association with the water type and water mass tabulated. Twelve species representing three genera were collected. All of these are found in tropical and sub-tropical waters. Chaetognaths fail as satisfactory indicators of the Virginian Coastal water intrusion into Raleigh Bay because of the absence, in our collection, of characteristically Virginian types in the southern limits of that faunal subprovince. INTRODUCTION Prior to the efforts of Pierce (1953), who described the distribution of the chaetog- naths over the continental shelf off North Carolina in relation to the hydrography of the area, and Bumpus (1955), who considered the circulation of these waters, little was known about the effect of the circulation system on the distribution of planktonic elements of the flora and fauna in the Hatteras area. Parr (1933) noted in winter a temperature barrier at Cape Hatteras corroborated by Bigelow (1933) and a fairly large cold-water temperature zone southwest of Cape Hatteras. This temperature barrier is well illustrated in the surface temperature charts of Fuglister (1947). Bigelow and Sears (1935) pointed out the abrupt transition in salinity which occurs at Cape Hatteras, occasioned by the wedge of pure oceanic water (>35*5%o) which presses in close across the shelf in Raleigh Bay and entirely separates the shelf and slope water bands to the north from the low coastal salinities farther south. The data available at that time suggested that the situation exists throughout the year. The existence of a barrier at Cape Hatteras has been postulated in defining the Carolinian and Virginian faunal subprovinces (Johnson, 1934). Many other zoologists have separated the cold-water and warm- water fauna on the continental shelf at this cape. Williams (1948, 1949), Sutcliffe (1950) and Pearse and Williams (1951) * Contribution No. 760 from the Woods Hole Oceanographic Institution. t Part of this research was sponsored by the Office of Naval Research under contract No. 27701. 92 Hydrography and distribution of chaetognaths over the continental shelf off North Carolina 93 found bottom living fauna, algae and plankton in Onslow Bay having in general tropical affinities. However, they noted in their winter and spring collections, in addition to the usual populations, a few individuals from a small number of species with northern affinities. Our recent collections provide material for a better understanding of the hydro- graphic influence on the distribution of the chaetognaths in the area than has been available heretofore and indicate how the hydrographic barrier at Cape Hatteras is occasionally breached, thus permitting the temporary establishment of the anomalous communities noted by Williams, Pearse and Sutcliffe. ACKNOWLEDGEMENT While the many outstanding investigations of Henry B. Bigelow do not strictly pertain to the area under discussion, the authors are especially grateful for his studies of the waters of the Gulf of Maine and the continental shelf from Cape Cod to Chesapeake Bay. His efforts have been a constant influence as we have attempted to extend the knowledge of shelf waters slightly beyond the area he so carefully and completely considered. THE DATA The data comprise a series of hydrographic sections together with plankton col- lections made in May and June 1953 (Caryn Cruise 64) and January 1954 (Atlantis Cruise 196) (Fig. 1). At each station serial temperature-salinity depth determinations were made to the bottom or to nearly 500 metres, whichever was less. At most stations quantitative oblique plankton tows, using Plankton Samplers (Clarke and Bumpus, 1950) fitted with #2 silk nets, were made from near the bottom or 100 metres. When feasible two Plankton Sampler tows were made dividing the water column in two. The chaetog- naths were picked out of the plankton samples and identified (Tables I and II, Fig. 12). These plankton data are more nearly quantitative and extend closer to the shore than those of Pierce (1953). HYDROGRAPHY Bumpus (1955) has shown that, south of Cape Hatteras, a southerly flowing coastal current, such as is common north of Cape Hatteras, is a transient afi"air. Such a current, when present, is restricted to a very narrow portion of the continental shelf. The dynamic pressure gradient resulting from the combined effect of the runolf and the cross-shelf thermal gradient together with the prevailing wind and the frictional drag of the Florida Current provide for a northeasterly drift over a broad pari o\' the Carolina continental shelf. In addition to the water of the Florida Current there arc two types of water in this region which we have named Virginian Coastal water and Carolinian Coastal water.* There are also mixtures of each of these with the Florida Current water. Virginian refers to the shelf water from Cape Cod to Cape Hatteras. Carolinian refers \o the * We have used here two names apparently new to oceanography inasmuch as we have been unable to find names concisely describing the separate continental shelf regions north and south of Cape Hatteras. Because the terms Virginian and Carolinian are used to describe the faunal subprovinces of the continental shelf, we shall introduce these terms to identify the water types on the shelf. 94 Dean F. Bumpus and E. Lowe Pierce 36° 9^^^^^^ 78° 77° 76° 75° 74. 1 {^^■■^■KlMP^iv' 36' ^^H^^^BM|^^^^^B' -'HV ^^^^^BB^^I^H^BHi^r^ ^^^^^^^^^^^^^^^^^^^^^S .^^ ^■■- "^-o ^^^^^^^^^^^^^^^^^^^^^^^^^^^^^^^^^^B^^^^^^^^^^^r ^r^ 35' ^^^^^^^ O' ••91 OlS8 169 O qITO I 35' 34° W^ml r- ^79 C ■• 176 / -186 X. I77T'»I75 ^185 ■■■•■-"■■■■■•••3-\ /' •'" /' •l74 _I8J lY 34' 33* ill 1 1 1 1 33° 7 -^ 1 1 1 1 4» 9* 78» 77* 76» 75' 7 Fig. 1 (A). Location of stations, on Caryn Cruise 64, May and June 1953. Open circles indicate no plankton tows were made. shelf water from Cape Hatteras southwards. UnpubUshed data indicate that Caro- linian Coastal water, as described below, extends to the offing of Daytona Beach and possibly at certain times to the offing of Cape Canaveral. The Virginian Coastal water is freshened by river water entering close to the surface inshore and salted by indrafts of slope water over the bottom from offshore (Bigelow and Sears, 1935). There is no widespread contribution to this coastal water from the south, nor flooding of the surface water with pure oceanic water of high temperature, nor upwelling onto the shelf of cold abyssal water (Bigelow, 1933). The Carolinian Coastal water is composed of Florida Current water and river effluent. This mixture is in general more saline than most coastal waters because the river runoff is less than for other sections of the coast ; the effluent from the sounds is more saline than from river mouths ; and the highly saline Florida Current frequently makes broad invasions over the continental shelf. There is no regular communication between the Virginian and Carolinian Coastal waters, although the frequent northeast storms from November to May (Miller, Hydrography and distribution of chaetognaths over the continental shelf off North Carol ina 95 1 — 7 5130 .5131 /^SISS 5140 .5154. ..-.y Yiii o 5(32 Y 35* 34* 33' 79° 78' 75* Fig. 1 (B). Locations of stations on Atlantis Cruise 196, January 1954. Open circles indicate no plankton tows were made. 1946; New York University, 1954) provide the energy for transient indrafts of Vir- ginian Coastal water into Raleigh Bay. Three events may take place following such an indraft: A. If the storm lasts for only a short period, one or two days, this water will eventually become absorbed within the Carolinian Coastal water, modifying it in proportion to the mixture. B. A violent meander of the Florida Current may completely sweep the indrafted Virginian Coastal water out of Raleigh Bay toward the northeast. Such an event would be aided by a southeast storm. C. If the northeast storm lasts for a period longer than two days or if another northeast storm follows within a few days, the indrafted water may eventually be driven around Cape Lookout into Onslow Bay. Such an occurrence could account for the presence there of the winter-spring species with northern atlinitics described by Williams (1948, 1949), Sutcliffe (1950), and Pearse and Williams (1951). Should this occur at a time of substantial runoff when a southerly flowing coastal 96 Dean F. Bumpus and E. Lowe Pierce current is being maintained in Onslow Bay and southward, such water movements may distribute planktonic elements of Virginian fauna for some distance along the coast. We have postulated here a sporadic occurrence which may contribute appreciably to the finding of anomalous species in the inshore waters of the northern part of the Carolinian subprovince. These species will find themselves in waters compatible to their existence until vernal warming traps them. Vernal warming or reduction in runoff with a consequent increase in salinity w^ill kill those species least resistant to higher temperatures and salinities. THE DISTRIBUTION OF TEMPERATURE, SALINITY AND DENSITY IN MAY AND JUNE, 1953, AND JANUARY, 1954 Typical spring conditions were encountered on the May and June cruises (Figs. 1, 2, 3, 4 and 5). These are the weak cross-shelf temperature gradient, moderate vertical temperature gradient, and the penetration across the shelf of highly saline Florida Current water along the bottom into Raleigh and Onslow Bays. In contrast the temperature and salinity in the Wimble Section are lower with weaker vertical tempera- ture gradients and stronger salinity gradients. Typical winter conditions for the Wimble, Onslow and Long Bay sections were observed in January 1954 (Figs. 1 , 6 to 10). Note the cold coastal water in the Wimble section and the nearly as cold, highly saline water in Onslow Bay and Long Bay. But Raleigh Bay was filled in its inner part with coastal water from north of Cape Hatteras, in contrast to the typical winter conditions of January and February 1950 (Bumpus, 1955) and February 1931 (Bigelow and Sears, 1935), when the 36°/oo isohaline (Florida Current water) was pressed well in across the continental shelf. This anomalous condition was due to a north-east storm several days earlier. This intrusion of Virginian Coastal water into Raleigh Bay is further discerned in the temperature-salinity relation (Fig. 11). The water at Stations 5141, 5142 and 5143 is clearly Virginian Coastal water. The temperature-salinity relations also provide a clue to the sources in the Florida Current contributing to the composition of Carolinian Coastal water. The water over the inner middle part of Long Bay (Stations 5149, 5150, 5151), Onslow Bay (Stations 5134, 5135, 5136, 5146) and the southern part of Raleigh Bay (Station 5145) in January appears to be from depths in the Florida Current, i.e. from depths perhaps as great as 150 metres. This water has been forced up onto the shelf in the course of current meanders and chilled by the colder air temperatures encountered there. Stations 5133 in Onslow Bay and 5147 and 5148 in Long Bay, the closest inshore stations, indicate appreciable dilution of this Florida Current water with river effluent and greater chilling as a result of the colder air temperatures near the coast. In contrast, the water over the outer parts of the shelf (Stations 5144 in Raleigh Bay, 5137, 5138 and 5139 in Onslow Bay and 5152, 5153, 5154 in Long Bay) is Florida Current water which has moved in over the shelf with no change in depth and has mixed only slightly with the water inshore of it. The intrusion of Florida Current water along the bottom in Onslow and Raleigh Bays in May and June (Figs. 3, 4, 5 and 1 1) was probably along surfaces of equal density. It has been forced onto the shelf by the meanders of the current as it passes Hydrography and distribution of chaetognaths over the continental shelf olT North Carolina 97 ■3- >% ca o j: E c g D >. c v g C c 'E ■r. 3 u a. E ij o c 3 t/1 Q so o E 98 Dean F. Bumpus and E. Lowe Pierce 196 STA NBR 197 5 10 15 20 25 NAUT MILES Fig. 3. (A) Distribution of temperature in ^ C, (B) salinity in °/^^ and (C) density (at), in Section II, Raleigh Bay, June 1953. 162 5 10 15 20 25 NAUT MILES ZOO Fig. 4. (A) Distribution of temperature in " C, (B) salinity in "/^^ and (C) density (at), in Section III, Onslow Bay, May 1953. Hydrography and distribution of chaetognaths over the continental shelf ofT North Carolina 99 A 190 STA Nan 188 18 7 Fig. 5. (A) Distribution of temperature in "' C, (B) salinity in ""[^^ and (C) density (at), in Section IV. Onslow Bay, June 1953. SI29 STA MBR SIV> Fig 6 (A) Distribution of temperature in C, (B) salmily in ,, and (C) density (at), in Section V. Wimble Shoals. January I "^54. 100 Dean F. Bumpus and E. Lowe Pierce 5142 STA.NBR 5143 I 7^ 5144 Fig. 7. (A) Distribution of temperature in "' C, (B) salinity in °l^^ and (C) density (ot), in Section VI, Raleigh Bay, January 1954. O 5 K) 15 20 25 NAUT MILES Fig. 8, (A) Distribution of temperature in ° C, (B) salinity in °/oo and (C) density (at) in Section VII, Onslow Bay, January 1954. Hydrography and distribution of chaetognaths over the continental shelf off North Carolina 1 1 5148 5149 5150 5151 ' ^ I I 5152 515 3 STA NBR 5154 Fig. 9. (A) Distribution of temperature in ° C, (B) salinity in ''/^^ and (C) density (o,), in Section VIII, Long Bay, January 1954. along the continental slope. During other parts of Caryn Cruise 64, the position of maximum current was observed to shift as much as 14 miles either side of the mean axis of flow (which off Onslow Bay normally lies about 18 miles south-east of the 100 fathom line. Associated with these shifts of position were onshore and offshore deflections in the direction of the current (von Arx, Bumpus and Richardson, 1955). This provides the energy to push water up the slope across the shelf. It also appears from salinity measurements that as the meanders move offshore they draw low salinity water (from the surface over the shelf) into the current. The proximity of the Florida Current to the continental shelf precludes the occur- rence (between the Current and the Carolinian coastal water) of slope water such as is found north-east of Cape Hatteras between the continental shelf and the Gulf Stream. DISTRIBUTION OF THE CHAETOGNATHS The chaetognaths collected in this area include 12 species representing three genera: Sagitta bipunctata, S. enflata, S. helenae, S. hexaptera, S. hispida, S. lyra, S. minima, S. serratodentata, S. tenuis, Krohnitta pacifica, K. subtilis, and Pterosagitta draco. The quantitative composition of the plankton tows is recorded (Tables 1 and ID. The species are listed in the order of their association with water of low or high salinity from left to right. The stations are arranged in the order of distance from the coa^.t. The ranges of individual species overlap others considerably and in many cases completely. Nevertheless the tables show that there is a general correlation between salinity tolerance and the distance from the coast at which various species were found. The most euryhaline-^urythermal species was 5". enflata (Pierce, 1953), although it is doubtful if it can tolerate for long periods salinities as low as those in which 5. hispida are usually found. S. hispida, S. helenae, and S. tenuis are species which are largely restricted to water of the continental shelf below Hatteras. The remainder o^ 102 Dean F. Bumpus and E. Lowe Pierce Fig. 10. Distribution of surface temperature in ° C (upper), salinity in ^/oo (middle) and density (m) (lower) on North Carolina Shelf, January 1954. Dash-dot line indicates track of ship. the species were found principally in the Florida Current. The increase in number of species is evident as one proceeds offshore over the shelf into Florida Current water. Sagitta bipunctata. The distribution and abundance of this species was similar in the area covered by spring {Caryn 64) and winter {Atlantis 196) cruises. S. bipunctata was collected in small numbers at several of the stations beyond the continental shelf and in a few instances they ranged shoreward about midway over the shelf (Tables I and II). With one exception (Station 5143) it was taken only in water where salinity was greater than 35 -5° j^^. It therefore appears to be restricted in this region to Florida Current water or water recently mixed with such water. Bigelow and Sears (1939) did not find this species between Cape Cod and Chesapeake Bay. Sagitta enflata was taken in almost every plankton sample on both cruises (Fig. 12). It was the most abundant chaetognath as well as the most widely distributed. The greatest concentrations were found near the edge of the shelf where as many as 160 per ten cubic metres were recorded. That they are sensitive to extreme conditions which are occasionally encountered in this area is borne out by their absence in the winter cruise from the three inshore stations north of Cape Hatteras. Here in water of less than 10° C and 32°/oo not a specimen was found. Moreover only one specimen was present at the two inshore stations south of Hatteras, where the water had been derived from Virginian Coastal water driven around the cape. Hydrography and distribution of chaetognaths over the continental shelf olV North Carolina |()3 1 r WIMBLE J L WIMBLE SAUNITY y„ "1 1 r y lee- -^•. ; RALEIGH ■^ ^ I9S-I95 " V ■. L_Li -I 1 I L J L « II -J 1 1 L_ _J4 ~i r ONSLOW JlM- rr7 / . - J. ,.« "-< -^ ONSLOW J,,j_ ' ». * • • . I - Ill J I I IV ' I 1 1 r RALEIGH 5l44f ! 1 r ONSLOW VI 5IJ7-fc i SI59 » ■ /». SlJt 146 LONG J4 ~i r SI55 * VII J L ►Sl»4 9191 • X1I9I *>ISO "♦''^ ••9.49 9146 I vm J I Fig. 11. Temperature-salinity diagrams, upper for Caryn Cruise 64, May and June 1953; lower for Atlantis Cruise 196, January 1954. The stippled line in the diagrams is the left side of an envelope describing the T-S characteristics of the Florida Current, Parr's (1937) stations in the Straits of Florida. Sagitta helenae was found all over the continental shelf south of Cape Hatteras but it disappeared at the stations beyond the edge (Fig. 12). The greatest concentrations were found in the middle of the shelf where as many as 94 specimens per ten cubic metres were collected. This species is able to tolerate a rather wide range of salinity, varying from approximately 32 to 36^o,. Although present in the edge of the Florida Current at times, it was never found far inside the current proper, and it does not appear from these data that it is a true inhabitant of such water. This species has been closely associated with continental shelf water from the Gulf of Mexico (Ritter-Zahony, 1910; Pierce, 1951, 1953) to Cape Hatteras. Apparently it disappears jusi north of the cape where the colder, less saline water is encountered over the shelf. Only a few specimens were taken near the edge of the shelf above Hatteras in the spring and winter cruises. Bigelow and Sears (1939) do not record this species in their plankton studies from Chesapeake Bay to Cape Cod. Because of its close relation to continental shelf waters from North Carolina southward, this species appears to be of special interest in its relation to the hydrography of this area. 1 he movement of Virginian Coastal water around Hatteras in January displaced this species from the inshore stations jn Raleigh Bay where it usually occurs (Fig. 12). 104 Dean F. Bumpus and E. Lowe Pierce Table I The distribution of chaetognaths by station during Caryn Cruise 64. Species arranged in approximate series with respect to saUnity, those which normally occur in coastal water on the left. Figures indicate number per 10 cubic metres of water strained 4: ■^ «i i 1 c^2 <5 ^ ^ -s: S ■4. .5 5 S ^5 3 .§. i 1 1 ^ Q ^ ^ ^ oj '>j ^ ^ c/j ^ k i< a^ t2.s :x SECTION I— -WIMBLE SHOALS 165* 014 82 48 51 8 134 71 166 019 23 14 19 3 1 1 60 98 0-40 111 6 44 3 3 120 72 SECTION II —RALEIGH BAY 193 0-20 18 47 56 140 113 194 010 72 2 16 98 111 195 015 7 2 8 8 6 3 1 39 114 15-30 83 3 79 11 5 1 1 1 163 89 196* 0-30 10 1 1 11 90 30-60 38 4 35 4 1 1 3 1 244 270 197 0-80 46 2 8 18 3 100 131 80-160 11 6 1 1 30 158 SECTION III — ONSLOW BAY 182 012 9 1 7 73 181 0-20 4 5 27 84 152 180 0-15 15-30 35 40 1 1 10 46 65 140 141 161 179 0-31 94 4 52 23 314 180 178 015 52 1 13 41 3 172 158 15-30 54 2 32 10 164 168 177* 0-50 67 1 6 62 1 1 11 208 138 50-100 29 1 29 5 1 2 184 287 176 0-30 4 2 54 2 2 2 110 163 30-62 43 15 8 41 5 138 122 175 0-90 137 3 12 121 3 16 150 51 90-180 11 3 7 2 1 31 122 174 0-80 30 39 4 2 3 18 198 206 80-160 3 2 5 1 1 2 7 44 278 173 0-80 21 1 25 1 1 7 136 244 80-160 1 1 4 1 1 17 300 172 0-75 3 1 5 4 1 1 1 24 207 75-150 3 3 6 1 2 3 59 318 SECTION IV — ONSLOW BAY 192 0-10 10-20 2 5 11 130 140 191 0-13 54 1 25 65 81 13-26 29 2 38 1 93 135 190 016 23 30 3 39 70 16-37 23 41 70 20 98 98 189 0-20 102 6 10 16 2 114 84 20-40 48 8 94 7 194 124 188* 0-20 35 4 1 2 35 83 20-40 25 18 3 3 4 53 98 186 0-50 9 19 4 106 337 50-100 1 1 1 2 1 1 23 385 185 0-65 30 24 2 2 3 184 297 65-130 5 10 8 1 5 111 332 184 0-70 10 16 1 3 8 90 235 80- 140 5 6 1 1 1 1 1 2 60 297 183 0-80 9 19 2 1 7 89 231 80- 100 1 2 3 1 1 3 1 26 297 * One hundred fathom contour lies between this station and the next. Hydrography and distribution of chaetognaths over the continental shelf off North Carolina 1 05 Table II The distribution of chaetognaths by station during Atlantis Cruise 1 96 Species arranged m approximate series with respect to salinity, those which normally occur in coastal water on the left. Figures indicate number per 10 cubic metres of water strained "g. --, i .2 ^cpth of sai (metres) 5; 1 s i 5! 3 1 1. .2 lal No. sample 1. ^ Q ^ ^ ^ ^ ^ ^ ^ ^ < ■< ^ ^.s — SECTION V— -WIMBLE SHOALS 5126 0-30 47 72 1 S4 5127 0-20 21 2 1 34 1 _'*T 141 5128 0-28 1 1 3 47 1^1 159 167 5129 0-45 1 7 1 19 5130* 0-25 34 5 2 45 2 14*^ 165 5131 100 7 3 43 SECTION VI —RALEIGH BAY 5141 015 1 98 • 113 115 5142 016 13 1 4 19 108 12-24 1 8 1 8 29 169 5143 020 78 16 14 2 30 6 3 241 161 20-40 49 15 4 2 10 1 2 1 2 152 178 5144 0-50 9 3 1 1 17 1 1 2 64 193 50-100 8 3 1 21 1 1 4 79 208 5145 015 15-30 33 2 1 1 11 4 1 13 5 3 3 95 5 142 120 5146 015 16 1 4 1 8 4 1 40 111 15-30 11 3 7 1 4 26 97 SECTION VII — ONSLOW BAY 5133 13 21 14 183 8 267 118 5134 019 59 30 66 7 2 401 185 5135 0-25 56 2 5 11 6 1 1 124 152 5136 0-28 3 2 2 2 2 1 14 120 5137 0-39 27 5 12 6 21 1 4 156 204 5138* 0-20 50 5 38 13 2 3 145 132 20-40 9 1 3 1 TV 147 5139 0-50 166 3 6 2 31 1 3 6 174 156 50-100 25 6 23 3 3 78 129 5140 0-50 9 12 1 2 1 3 50 181 50-100 6 4 6 1 5 40 192 SECTION VIII — LONG BAY 5147 015 5 1 minute unidentified chaetognaths in ; sample 5 167 5149 015 83 10 68 3 97 59 5150 0-20 17 4 20 1 48 113 5151 0-24 13 3 50 2 89 133 5152 0-30 122 13 44 T 9 1 1 265 137 5153 5-30 89 1 22 1 5 19S 168 5154 0-50 50-100 39 29 2 I 1 1 13 5 1 1 3 1 69 S4 155 226 * One hundred fathom contour lies between this station and the nc\t. Sagitio hispida is an inshore species which was taken at Stations 182. 5133, 5141 in water with a salinity of 31 to 35°/oo- No specimens were collected as far offshore as the ten-fathom lino. Other records (Pierce, 1951) indicate that this species is present in bays and at points close to shore south of Cape Hatteras. Sagitta minima was taken many miles on either side of the continental slope (Fig. 12). It was not taken at the stations within the ten-fathom curve or at the stations farthest in the Florida Current. 106 Dean F. Bumpus and E. Lowe Pierce - NUMBER PER 10 CUBIC METERS = • = 50 . NUMBER PER 10 CUBIC METERS 0=0 • = between the air and the bicar- bonate of the sea water. If this procedure is not adopted, there may be a serious loss, and variable results will be obtained (Calvin, et ai, p. 123). The lens tissue should not be silicone treated, since this type tends to curl up on drying. There was no tendency to curl with the photographic Ions tissue used, so that the addition of collodion was unnecessary. To ensure that no change in absorption of the radiation was caused by variations in salinii\. the same sample of sea water was used throughout the experiments. The activity of each sample was measured by exposing it in a holder at a constant distance from the mica end-window of a G.-M. counter, and the counts were recorded on a scaling unit. The results are expressed as counts per minute, but since we know the concentration of cells in the culture and the activity of both whole culture and filtrate, we can also express them as cell equivalents. The bodies and faecal pellets of the Calanus have an appreciable self-absorption, and even after they had been torn up by needles the losses were considerable. Good duplicates were obtained by 1 ] 2 S. M. Marshall and A. P. Orr the use of a small disintegrator. This consisted of a narrow tube (diameter 9 mm) about 7 cm long, into which was fitted a perspex piston of almost the same diameter as the tube, and shaped at the foot to fit the bottom of the tube. The Calanus or the faecal pellets were put in the tube in about 0-5 ml of sea water, and the piston rotated by a small motor. By raising and lowering the tube the water and the Calanus or faecal pellets were forced past the rotating piston and thus disintegrated. After washing down the piston and tube and making the volume up to 3 ml, five aliquot samples were taken for measurement of activity. Microscopical examination showed that disintegration was almost complete. No fragments of a Calanus body could be recognized, and there were very few recognizable bits of faecal pellet. The disadvantage of the disintegration method is the dilution of the activity, since only 0-2 ml samples were used from the 3 ml of fluid. This can be countered by taking more sub-samples, or by making much longer counts. In a feeding experiment a number of bottles of about 70 ml capacity were filled with the diluted culture prepared as described, and sampled for a count of cell number and activity. Into each bottle was introduced a single Calanus. Since females feed better than either Stage V or male Calanus they were always used. Each bottle was then tied in a black cloth bag (because Calanus feeds better in the dark than in the light), and attached to a wheel revolving about once every three minutes in a vertical plane. This keeps the culture cells from sinking to the bottom and so giving the Calanus an accumula- tion to feed on. Control bottles containing filtrate from the culture used were also set up to measure any uptake of ^*C from solution. Other control bottles, containing culture but no Calanus, were used to measure the^*C returned to solution by the respiration of the plant cells. After leaving them to feed for a suitable time, usually 15 to 18 hours, each Calanus was removed, washed three times to free it from radioactive water, disintegrated and sampled. The contents of the bottle were then poured into a flat-bottomed perspex dish with the inside angles bevelled, and the faecal pellets picked out under a binocular microscope. These too were washed, disintegrated, and sampled and their activity was measured. The activity of the Calanus body added to that of the faecal pellets gives a measure of the total i*C removed from the culture. From these and the culture reading can be calculated the number of cells ingested, the percentage digested and the volume of water swept clear. The results can be expressed either as counts per minute or as cell equivalents. The cell equivalent is a useful figure when consider- ing the total amount taken up, but if we express the activity of the faecal pellets as cell equivalents, it must be remembered that each " cell equivalent " really represents several cells, the number varying according to the percentage digested. EXPERIMENTAL WORK Feeding experiments with female Calanus were made, using cultures of the diatom Skeletonema costatum, one of the more important spring diatoms in the sea, and the flagellates Cryptomonas sp. (Plymouth strain 23) and Syracosphaera carterae. The cultures when used were only a few days old, and were probably in the exponential growth phase. It was thought that some of the "C might be present in the inorganic form, either adsorbed on the cells or in solution inside. This was tested with Skele- tonema by exposing samples on planchettes to the fumes of hydrochloric acid (Steemann Nielsen, 1952), and comparing the activity before and after exposure. About 7 % of the total disappeared with this treatment. With Syracosphaera, which possesses large numbers of calcareous coccoliths, the loss seemed to be greater but accurate measurements were not made. The control bottle containing culture but no Calanus showed that, as a result of the respiration of the plant cells, the i*C content of the filtrate had risen by 2-5%. This will cause a slight underestimate of the amount taken up by the Calanus. A preliminary experiment was done with a culture of Skeletonema costatum in two different concentrations. It was thought that the activity of the faecal pellets and, in the lower concentrations, of the bodies also, would be too weak for the disintegra- tion method, so they were torn up as finely as possible with needles and put in 0-2 ml Experimental feeding of copepod Calanus finmarchicus (Gunner) on phytoplankton cultures 1 ] 3 Of sea water on a planchette. Owing to self-absorption, this was not a satisfactory method, giving results which were too low, and it was therefore impossible to calculate the percentage digested. If we assume that this is the same as in a later experiment (see Table I), a figure can be obtained for the volume filtered which will be approximately correct for the richer concentration but minimal for the weaker. The estimated volume filtered in 24 hours in a concentration of 106,000 Skeletonema cells per ml varied from 1-6 to 40 ml in 24 hours, and in a concentration of 10,600 cells per ml from 5-7^2-5 ml. It is usual to find that in high concentrations of food cells filtration falls off, and the figures obtained compare well with our earlier experiments using '^?. Table I Feeding experiments with Calanus using cultures grown with "C Species Concentratiuns Cal- anus o Time of expt. in hours iFaecal pellets I No Skeletonema costatum Culture Filtrate Cryptomonas Culture Filtrate 1615 counts ml min 144000 cells ml 001 counts cell 35 counts/ml /min 1 T 3 4 5 A B I7i 17-1 \1\ Ml 17* 1275 counts /ml /min 12600 cells ml 010 counts cell 60 counts/ml/min 1 2 3 4 5 A B in in in in 21i m in 54 17 47 10 4 Syracosphaera carterae Culture Filtrate 840 counts/ml/min 1 13500 cells/ml 2 06 counts 'cell 3 8 counts ml min A 17 65 17 38 17 45 17 38 51 80 56 57 2 Lost Body Total I less removed Countsfiltrate, c min | Imin cimin ' ml °o filtered used in 24 hr lis 192 301 59 36 170 465 449 220 516 817 283 673 119 }«. 1095 475 974 178 74-6 59-6 69- 1 090 0-38 0-81 66-9 14 289 86 392 585 755 77-5 0-66 780 1245 62-7 110 811 1260 64-4 113 454 674 67-4 0-58 589 1105 53-3 0-98 1 930 I2I9 76-3 2 07 180 266 67-7 . 0-45 885 1277 69-3 216 30 The results of a second experiment with Skeletonema are shown in Table I. In this and subsequent experiments all the bodies and faecal pellets were disintegrated. As is usual in feeding experiments, individual variation was considerable; the number of faecal pellets varied from 10-54, and one Calanus did not feed at all. The amount digested varied from 60-75%, a figure very similar to that obtained with cultures labelled with ^^P. Unfortunately the Calanus v/ere in poor condition and the volume filtered, less than 1 ml in 24 hours, was low. The activity of the culture was not high enough to give very accurate results. In both experiments with Skeletonema, the cell concentration was high compared with what is found in the sea. In an experiment with Cryptomonas, which has cells about 20ix long, all the females fed well, although they filtered on an average only about I ml in 24 hours (Table 1). The digestion ranged from 53-78%, as compared with 51-89% in cultures labelled with 3 2p. Finally, the experiment with Syracosphaera was carried out using three Calanus in a concentration of 13,500 cells/ml. Judging by faecal pellet production, they fed 114 S. M. Marshall and A. P. Orr well, but the maximum volume filtered was just over 2 mi in 24 hours (Table I). The digestion (68-76%), was decidedly lower than that found using Syracosphaera labelled with ^'^p^ in which it was usually over 90%. This may be due to the presence on its surface of a layer of coccoliths which would contain "C in the form of calcium carbonate. The rate of production of faecal pellets was not high in any of the experiments with ^*C, the most rapid being found in a Calanus in Skeletonema culture, which produced an average of one every twelve minutes. It has been found that even with a much more rapid production, digestion of the phosphorus-containing fraction remains high. ACKNOWLEDGEMENTS The experiments with ^*C labelled cultures were carried out at The Laboratory, Plymouth, and we are very grateful to the Director and staff for their help. More especially we wish to thank Dr. H. W. Harvey and Dr. B. C. Abbott for their advice and assistance, and Dr. M. W. Parke and Miss D. Ballantine for supplying us with cultures. REFERENCES Calvin, M., Heidelberger, C, Reid, J. C, Tolbert, E. M. and Yankwich, P. F. (1949), Isotopic carbon. New York and London, 376 pp. Clarke, G. L. and Bonnet, D. D. (1939), The influence of temperature on survival, growth and respiration of Calanus finmarchicits. Biol. Bull., 76, 371-383. Clarke, G. L. and Gellis, S. S. (1935), The nutrition of copepods in relation to the food cycle of the sea. Biol. Bull., 6S, 231-246. Dakin, W. J. (1908), Notes on the alimentary canal and food of the Copepoda. Int. Rev. HydrobioL, 1, 772-782. EsTERLY, C. O. (1916), The feeding habits and food of pelagic copepods and the question of nutrition by organic substances in solution in the water. Univ. Calif. Pubi, Zool., 16, 171-184. Fuller, J. L. (1937), Feeding rate of Calanus finmarchicus in relation to environmental conditions. Biol. Bull, 72, 233-246. Fuller, J. L. and Clarke, G. L. (1936), Further experiments on the feeding of Calanus finmarchicus. Biol. Bull., 70, 308-320. Gauld, D. T. (1951), The grazing rate of planktonic copepods. /. Mar. Biol. Assoc, U.K., 29, 695- 706. Gest, H. and Kamen, M. D. (1948), Studies on the phosphorus metabolism of green algae and purple bacteria in relation to photosynthesis. J. Biol. Chem., 176, 299-318. Goldberg, E. D., Walker, T. J. and WmsENAND, A. (1951), Phosphate utilization by diatoms. Biol. Bull., 101, 274-284. Kamen, M. D. and Spiegelman, S. (1948), Studies on the phosphate metabolism of some unicellular organisms. Svmp. Quant. Biol., Cold Spring Harb., 13, 151-163. Lebour, M. V. (1922), The food of plankton organisms. /. Mar. Biol. Assoc, U.K., 12, 644-677. Marshall, S. M. (1924), The food of Calanus finmarchicus during 1923. /. Mar. Biol. Assoc, U.K., 13, 473-479. Marshall, S. M. and Orr, A. P. (1955), Studies on the biology of Calanus finmarchicus. VIIL Food uptake, assimilation and excretion in adult and Stage V Calanus. J. Mar. Biol. Assoc, U.K., 34, 495-529. Nielsen, E. Steemann (1952), The use of radio-active carbon (C") for measuring organic production in the sea. /. du Cons., 18, 117-140. Rice, T. R. (1953), Phosphorus exchange in marine phytoplankton. U.S. Fish and Wildlife Serv., Fish. Bull., 54 (80), 75-89. Papers in Marine Biology and Oceanography, Suppl. to vol. 3 of Deep-Sea Research, pp. 115-133. The hydrography of the Gulf of Venezuela By Alfred C. Redfield Woods Hole Oceanographic Institution and Department of Biology Harvard University One of the very distinguished contributions of Henry B. Bigelow to the subject of oceanography is his study of the hydrography, plankton, and and fishes of the Gulf of Maine. It is the most complete description and analysis which exists of any circumscribed body of coastal water. Because of its breadth of outlook and high technical standard, this work is a model for studies in marine ecology. It is appropriate to contribute to this volume in Professor Bigelow's honour a sketch of the hydrographic conditions which are found in another gulf of somewhat similar pro- portions, but under far different influences than those which dominate the Gulf of Maine. Summary — The distribution of salinity, temperature, oxygen, and total phosphorus in the Gulf of Venezuela is described. The physical circulation appears to consist of two estuarine cells. The first is generated by the outflow from Lake Maracaibo, which terminates in a mixing zone over the sill off Calabozo Bay. where the water which occupies the deeper basin of the Bay is formed. The second is fed by water formed in this mixing zone which escapes seaward after mingling with more saline water drawn in from subsurface layers of the Caribbean. The semi-diurnal components of the tide are augmented by resonance in the Gulf of Venezuela, and with the wind account for the vertical mixing which occurs over the sill of Calabozo Bay. The trade winds, which predominate in winter, produce large seasonal differences in mean sea level across the Gulf, and control the distribution of the brackish water as it moves seaward from the outlet of Lake Maracaibo. Upwelling, which occurs in the lee of the Peninsula of Paraguana. is accompanied by an accumulation of phosphorus and a depletion of oxygen in the deep water near the coast. Similar conditions are found in the basin of Calabozo Bay. The influence of countercurrents on the biochemical circulation is discussed. The Gulf of Venezuela lies in the seaward extension of the syncline which fornix the Maracaibo Basin. It opens directly on the deep water of the Caribbean Sea, and carries into it the entire outflow from Lake Maracaibo. This is the most substantial accession of fresh water along the Venezuelan coast of the Caribbean, and is estimated at 21 billion cubic metres per year. The northeast trades blow steadily along the axis of the Gulf from December through April. The annual range in temperature is small, being about 2° C at Maracaibo. Being situated in 12^ N latitude the Corioli parameter is weak. Differences in salinity and the winds thus dominate the hydrography ol' the Gulf. * Contribution No. 776 of the Woods Hole Oceanographic Institution 115 116 Alfred C. Redfield The Gulf may be divided into two parts: the Outer Gulf and Calabozo Bay. These regions are separated by a sill with depths of 18 metres, extending along the 71st meridian. West of the sill a basin 28 metres in depth occupies the northern half of Calabozo Bay. East of the sill the bottom of the Outer Gulf slopes downward to provide depths of 40 to 80 metres over a considerable area. Access to the Gulf is probably limited to Caribbean water from depths not greater than 100 metres (Fig. 1). The only earlier observations on the Gulf of which I am aware are measurements of 72° 71° 70" Fig. 1. Bathymetric chart of Gulf of Venezuela. Based on H.O. No. 5520 — Depths in metres surface salinities across the Gulf in December 1953 by Gessner (1953 b, 1955), and a few records of chlorinity off the entrance to Lake Maracaibo by the Corps of Engineers, U.S. Army (1938). Undocumented statements relative to the water of Lake Maracaibo and its approaches, and on tide levels in the Gulf, are based on information secured by the Woods Hole Oceanographic Institution in the course of studies which it is expected to publish subsequently. These studies were made for the Creole Petroleum Corporation, which has graciously consented to the use of this information. The outflow of Lake Maracaibo was estimated from climatological data by Douglas B. Carter of the Johns Hopkins University Laboratory of Climatology (Carter, 1954). The greater part of the data to be discussed was secured by the Atlantis between December 7 and 9, 1954. Three sections across the Gulf and two extending seaward from the adjacent capes provide information on the distribution of temperature, The hydrography of the Gulf of Venezuela 117 salinity, oxygen, and total phosphorus content (Fig. 2). I am greatly indebted to L. V. WoRTHiNGTON and W. G. Metcalf who secured this data and to Nathaniel CoRWiN who analyzed the samples for total phosphorus, using the method of Harvey (1948). The data secured by the Atlantis are presented in Figs. 3 to 6, which show the distribution of the variables in the five sections occupied. Fig. 7 shows their distribu- tion in a section along the axis of the Gulf. ir 71" 70" Fig. 2. Position of hydrographic stations, sections and tide stations THE PHYSICAL CIRCULATION The Axial Circulation The more general features of the circulation are shown by the distribution of properties along the axis of the Gulf (Fig. 7). The distribution of salinity is especially informative, since the fresh-water outflow from Lake Maracaibo serves as an indicator of the water movements. In Calabozo Bay the circulation is estuarine. Along the western shore the outflow from the Lake produces a thin layer of brackish water of salinity less than 15\,^. As this water moves seaward the salinity of the surface layer increases gradually to about 30°/„^ over the sill. A rather sharp halocline at about 15 metres separates the surface layer from water having salinities greater than 307oo which occupies the deeper basin. 118 Alfred C. Redfield Over the sill, and on its seaward slope, there appears to be a zone of eflfective mixing, in which the surface water from the Bay mingles with the Caribbean water from the Outer Gulf. In this zone the salinity increases from 30 to >35°J^^ and does not vary greatly with depth. On one side this mixing zone produces the bottom water of SW 5238 5237 5236 NE NW 5253 5252 5251 5250 5249 5248 S E 10 20 - ' 40 60 70 80 90 - 100 I 35»75 35«62^v^5»56 1 35.6 l,35»75 35»62 p^\ 35 8 - Uje ^- .^ 36.0 \ 36.2 ~ 1 36.4 - 1 36*32 - \ "36.6 - 1 7e*M - 1 36*66 - I I 136 71 1 * 10 2 2 30 X lu ■40 - 60 , SW 5254 5255 5256 NE 20 30 - 40 o 50 70 - 80 ] 1 35*37 1 1 35*73 35*69 136,36 /' 35-6 - Tr_vr ' 35*62 - Wsi^ 36*71 - 35*70 ^36.0 - \ .36.2 36*36 - \ 36*56 _,,36.4 - \ ___.—— 36.6 ^ 36*68 11 ^"^ NW 5243 5244 5245 5246 5247 SE N 5242 5241 5240 5239 S in 10 DC 20 o 30 40 I P6*84 / V — '20 ^3.7, 23*03 23*67 \ \ ^ 12*90 j 23*87 ■*■ ^26j^ \ 33.21 -30 -y 32*66 ^^ "^ - V SALINITY , 7oo Fig. 3. Distribution of salinity in Sections 1-V, Gulf of Venezuela. See Fig. 2 for positions The hydrography of the Gulf of Venezuela 19 Calabozo Bay which, flowing westward as a countercurrent, suppHes the salt consumed in diluting the outflow from the Lake. On the other side it produces a mixture so similar to the superficial layers of the Caribbean that its identity can be recognized only with difliculty. SW 5238 5237 5236 NE 10 20 30 a: " UJ V- (Ij z 50 X t- o. UJ °60 80- 90- 100 ' 27«56 27»5e^^ 27««3 \ zimiO 27«59 27. 5 126 7^\ ^X 1 * ^v ^ \ ^\. 27»2I - V>\ \27 - r\ - 1 Z4.e9\^ _ 1 ^25 - 1 23«5T - I 1 ^23 |22 00 1 • NW 5253 5252 5249 5248 SE NW 5243 5244 5245 5246 5247 SE 1 2r»97 1 27*69 i I I I 1 1 1 2T«8T ^v j-«7a 27»7 y 10 20 - 1 V \ \ 27»70 \ 27 8-' 1 1 1 1 ^ 27«90 27#B5 """^ y 1 30 - IV - SW 5254 5255 5256 NE 30- 40- Q 50 - 60 - 70 - 80 \ 27.I8 27»3i 27»35 y^'A - \ \ 27.37 - 127 (5 '^ 27»53 \ 27.2 \ \ \27,3i 27»29 \ ^27 "~ V— _26 25»67 - V— \ 2«»7I 25 - N^ 24 25«49 n ^ N 5242 5241 5240 5239 S •S- 10 z 20 ' 30 1 27*63 27«69^^' 27«56 ; 27«2 f \27 74 ^ ——m % • ! z'." ,--''".y \ -.i'!L*_ -276— " ,_- - -V *■ ^^ 2T,9»^ ^^^ - V " TEMPERATURE , "C Fig. 4. Distribution of temperature in Sections I-V, Gulf of Venezuela. See Fig. 2 for positions 120 Alfred C. Redfield The distribution of temperature supports the above interpretation. The coolest surface water is found in the brackish outflow from the Lake. The surface water of Calabozo Bay warms as it moves toward the sill, where the warmest water and an almost uniform vertical distribution of temperature is found. The water of Calabozo SW 5238 5237 5236 NE 100 NW 5253 5252 5251 5250 5249 5248 SE 1 \ 4»3« 4»34 4*34 ,4.41 ««46 ^>.^ 4«3r / 10 - \ k4*29 4.36 4*36 \ 4«38 "^ >* N \ t 4.4 4«4I 4*44 ^1 f- « 20 UJ *- Vv^ 4*24 4*40 I Z z 30 - \ ~ "~~ — -42- \^ - I i- a. - /t ^ / Ad ^ / M y / m ^ / m ^ i m 5 / / - °40 'f 3. 3 y 50 III \3»72 3«5S 3*62 / ^%v}^^r . NW 5243 5244 5245 5246 5247 SE SW 5254 5255 5256 NE \ 4«4r 4.43 \4u36 4»36 10 V '' 4*37^ 1440 \ \ \ \ 4.4S 20 _ % \ ■".^ 4.43 30 ^ I 4.39 ""^--44 40 A 2 \ 4.06 50 \ 5.94 4 60 \^ -— --3.8 N »«s^ 3.61 70 on n ^v ""^ N 5242 5241 5240 5239 S OXYGEN , mVl Fig. 5. Distribution of oxygen in Sections I-V, Gulf of Venezuela. See Fig. 2 for positions The hydrography of the Gulf of Venezuela 121 Bay below the halocline is warmer (>28° C) than any other water in the region. This water cannot have arisen by direct advection from the deeper water of the Outer Gulf, which is colder and more saline. Data from the files of the U.S. Hydrographic Office show that the mean monthly SW 5232 5237 5236 NE 20 30 ui 40 z 50 SO - 70 - 80- 90 100 1 • 23 •29 •25 1 •2« •29 \ — I .sK I \'" •" I ^^ y X ,,0.3'' •w^ ^^ \ •29 •2« %ti "* / fl - \>--~- „....-- / / / l\ - \ ^04 \ .05' 0.6 III 08/ - ^k •«» •^ ^^"^ m^t ^ .y\^ - ni \:::_::>-"^ SW 5254 5255 5256 NE 10 - 20 - „30- lE bl »- bl Z 2 40 r I- n. UJ Q 50 60 70 SO n • 55 • 34 V *" VvJ" 1 Y»42 0.4 y • 31 • 32 0.3 - • Sr \ \ ♦«4 \ • 39 \ ^ #45 u ^^"^ NW 5243 524 S 5246 5247 SE H 10 - Ui z X 20 — 1 ^39 •39 J •43 ^^•52 • 59 1 V"*-^ / i r - \ 04 ^v^y - -»•-'-- •** ^ y ~ ^^--^^""""^ - IV - N 5242 5241 524'0 5239 S o 30 40 . .44 • 42/ •52 1 ^OB'^ - \ ( •«' 3« # • # Ir 0.6 o.8/'^>^ y^ ,^^ •3236 X" ~ • 36.09 • 36 19 30 CALABOZO BAY \ - 40 ~ \ - 50 - N. »'*''-',^-^ • 36 92 - 60 SALINITY . %o OUTER GULF \ - 70 - \ • 36 69 - 20 30 40 50 60 70 z I 7^3 / •27,56 •27.78 ~-J>J7.53.. 276 TEMPERATURE , *C 45 — 3.5--^IZ— -^; OXYGEN ml /I •*41 «43i • 4. 59 • 4.39 - \ •4.39 - -l • 4.25 - k >v •S-SZ ^«,. \ • 336 - \ • 3.69 10 20 30 40 50 60 70 ^523" ra!F" TOTAL PHOSPHORUS .^"g'^/ I • 0.30 Fig. 7. Distribution of salinity, temperature, oxygen,[and total phosphorus in Section A-A along axis of Gulf of Venezuela. See Fig. 2 tor positions 124 Alfred C. Redfield westward from the Peninsula of Paraguana. It provides a component which, when mixed with water from Calabozo Bay, forms a mass of water slightly colder and less saline than the water of the Caribbean. This mass may be traced as it flows west- wards close to shore around the Peninsula of Guajira (Fig. 8). The Sailing Directions report a westerly set of 1 -75 knots off this headland during the greater part of the year. In summary, the physical circulation of the Gulf of Venezuela appears to consist of two estuarine cells, separated by a transition zone of vertical mixing. The surface outflow of the inner cell is fed by water escaping from Lake Maracaibo, and is con- sumed in the mixing zone over the sill. This zone is the source of the deep counter- 72" 7r 70" Fig. 8. Salinity at 1 metre depth in Gulf of Venezuela current which provides salt to the surface outflow. The outer cell, less clearly defined, is fed by the water formed in the mixing zone, which escapes seaward at the surface after mingling with water from a countercurrent drawn in from the subsurface layers of the Caribbean. In both cells the surface drift is displaced to the left, under the influence of the wind. The estuarine cell of Calabozo Bay finds its counterpart in many bays of the eastern coast of North America. Aside from the quantitative effects of the controlling topography, the more general difference in the circulation arises from the dominant effect of the prevailing wind on the horizontal flow in Calabozo Bay, as compared to the Corioli effect in the estuaries of higher latitudes. The hydrography of the Gulf of Venezuela 125 Along the Atlantic coast the bays and estuaries discharge into, and lose their identity in producing, a broad band of coastal water of reduced salinity which separates the coast from the full sea water of the ocean. Along the Venezuelan shore a distinct band of coastal water is lacking, and Caribbean water in full strength penetrates the Gulf of Venezuela. The front separating the bay water from the full sea water is the mixing zone over the sill of Calabozo Bay. The water in this zone may be considered to be the rudimentary counterpart of the coastal water of the Atlantic coast. If this view be accepted, the outer cell of the circulation of the Gulf of Venezuela finds its counterpart in those processes taking place along the margin of the Gulf ir 7r Fig. 9. Temperature at 1 metre depth in Gulf of Venezuela Stream and in the slope water, by which the coastal water of the Atlantic coast becomes incorporated and lost in the general circulation of the ocean. Tides The tides at Aruba, like those of the Caribbean in general, are of the mixed type in which the diurnal constituents predominate. The diurnal range of tide is 0-8 feet. In contrast, at Zaparita Island, off the mouth of Tablazo Bay, the tide is predominantly semi-diurnal and the mean range is 3 feet. The dimensions of the Gulf of Venezuela are such that the semi-diurnal constituents are augmented by resonance to a high degree, while the diurnal constituents are relatively unaltered. 126 Alfred C. Redfield A comparison of the harmonic contents of the tides (Table I) shows that the M2 component at Aruba precedes that at Zapara Island by 123° or more than one-quarter period. High water occurs at Las Piedras only one-half hour earlier than at Zaparita Island, and 1 -3 hours earlier than at Zapara Island. These relations place the antinode 72" 71" 70" Fig. 10. Total phosphorus at 1 metre depth in Gulf of Venezuela Table I Tidal harmonic constants, U.S. Coast and Geodetic Survey (1951) Constituent Greenwich Epoch {G) Aruba Zapara I Amplitude {H)feet Aruba Zapara I Diurnal Oi Pi Qi 241° 228° 241° 247° 246 243° 244" 0-3 0-2 01 0-24 015 08 02 Semi-diurnal M2 S2 K2 161° 84° 282° 284 217" 248 268" 013 07 03 104 Oil 0-32 003 Sum — Diurnal amplitudes Semi-diurnal amplitudes 0-6 0-23 0-49 1-50 Ratio of sums 2-7 0-33 The hydrography of the Gulf of Venezuela 127 for the semi-diurnal tides across the entrance to the Gulf at a longitude intermediate between Aruba and Las Piedras. The amplitude of the M, component is increased eightfold, and the sums of the semi-diurnal components sixfold as the result of the resonance. 72" 71° l(f Fig. 11. Oxygen under saturation at 1 metre depth in Gulf of Venezuela Since enough water must cross the sill of Calabozo Bay, having a depth of 18 metres, to raise and lower the level of the Bay nearly one metre twice daily, it is evident that strong tidal currents are to be expected over the sill. These currents, combined with the wind, account for the mixing which appears to occur in this region. Water Levels The monthly mean tide levels at Las Piedras change by as much as 0-8 feet in the course of the year (Fig. 12, A). The pattern is similar to that observed at positions on the Atlantic coast south of Chesapeake Bay and in the Gulf of Mexico (Marmer, 1951), and evidently is part of widespread phenomena. The monthly mean tide levels at Zaparita Island follow a similar pattern, but the annual variation in level is only 0-4 feet. The mean annual difference in water level across the Gulf of Venezuela, between Las Piedras and Zaparita Island, is 0-92 feet in a distance of 93 nautical miles. The 128 Alfred C. Redfield difference in level varies with the season, being highest (1-1 ft.) in January, and lowest (0-6 ft.) in October (Fig. 12, B). The seasonal difference in levels across the Gulf is clearly related to the prevalence of north-easterly winds, as shown by wind data from Maracaibo (Fig. 12, D). The difference in levels may be explained in part by the reduced density of the water at the head of the Gulf. The magnitude of the difference is, however, so great as to raise a question as to the precision of the levelling between Las Piedras and Zaparita Island. There can be little doubt, however, that the mean tide level at Zaparita is one-half foot greater, relative to the level at Las Piedras when the north-east trades are at a maxi- mum, than it is when their influence is reduced. Clearly winds which affect the slope of the sea surface so greatly are adequate to produce the effects on the circulation which have been deduced from the hydrographic observations. 3.0 2 5 2 "I — I — r "1 — I — r ZAPARITA - LAS PlEOPAS -A MEAN TIDE LEVEL J I I i I I \ I I L J I L I 2 10 0.8 06 04 I I I I r 1 — I — r 1 I 1 r B DIFFERENCE IN LEVEL J I I I I I I I I I I [ I L -4 T — r 2 — - C WATER BALANCE l. maracaibo - J I I I l__l I I I I I I I L "I — I — I — I — I — r 100 — 80 — 1 — I — r • D WIND AT MARACAIBO I I I I I I I I I I I L J L M A M J S N M A M J A S N Fig. 12. A. Monthly mean tide levels. Average at Las Piedras for 1942-49 and at Zaparita Island for 1940-53. Data from Bar Survey, Maracaibo B. Difference in average monthly mean tide level at Las Piedras and Zaparita Island C. Average monthly net water balance of Lake Maracaibo. Estimated by Carter (1954) D. Per cent of time wind blew from N through ENE at Maracaibo, 1952. Data from Bulletin Bimensureal, Servicio de Meteorologia y Communicaciones, Ministerio de la Defensa , Fuerzas aereas, Maracay The hydrography of the Gulf of Venezuela 1 29 Seasonal Influences on the Hydrography The available data on the waters of the Gulf were collected in December. While there is no information on the conditions at other seasons, the annual variation in the two factors which control the circulation is known, and may be used to indicate the direction, if not the degree, in which the pattern will change with the season. The rainy season in the Maracaibo Basin extends from April to December. The observations were consequently made at a time when the quantity of fresh water in the Gulf is maximal. During the following four months the outflow from the Lake is reduced and may come to an end (Fig. 12, C). The salinity of Calabozo Bay may consequently be expected to increase, and the hydraulic forces which drive its estuarine circulation to weaken. These tendencies should be reversed beginning in April. A rough estimate indicates that the quantity of fresh water present in Calabozo Bay in December is equivalent to about two years' outflow from the Lake. It is probable consequently that in the four months of the dry season the change in the mean salinity will be limited. The effect of the north-east trade winds in maintaining the anticyclonic circulation in Calabozo Bay should continue through the winter. With the reduction in outflow from the Lake, the continued wind-induced motion will mix the water, and reduce the vertical and horizontal differences in salinity. After April the winds become more variable and slacken in intensity. As the summer progresses, increasing outflow from the Lake should restore the salinity stratification. With the weakened effect of wind, the brackish water should spread more diffusely across the Bay. Possibly the Corioli force overcomes the wind influence and causes the escaping water to follow the southern shore to the eastward in the latter part of summer. THE BIOCHEMICAL CIRCULATION The distribution of the major elements in sea water, which determine the salinity, is the result of purely physical processes of advection and mixing. It may be used, consequently, to trace the general circulation of the water. In contrast, the distribution of those elements which are present in limited quantity, and which enter into the composition of organisms, may be profoundly influenced by biological activity. In particular, elements such as phosphorus and nitrogen, which become incorporated into the substance of the phytoplankton growing in the surface layers of the sea, tend to be carried downvvards by the sinking of organisms, and are liberated at depth by the ultimate oxidation of the organic matter. There is thus a circulation of elements of biochemical importance which is different from that of the water itself, and of its biologically inert components. In the case of the Gulf of Venezuela, data on the total phosphorus and oxygen content are available for an examination of the biochemical circulation. Total Phosphorus The total phosphorus content of water includes the phosphorus present as inorganic phosphate ions, as organic compounds present in solution, and as components of suspended organisms. The total phosphorus content of the Gulf is higher than that of the source waters from the Caribbean. This is due primarily to the high phosphorus content of the F 130 Alfred C. Redfield water from Lake Maracaibo, which is on the average about 1 -4 [agA per litre, whereas the Caribbean water contains about one-quarter this amount. The general pattern of distribution of salinity and phosphorus in the surface waters (Figs. 8 and 10), and in the axial section (Fig. 7), are so similar as to suggest that both properties vary as the result of the system of circulation which mixes waters derived from the Lake and Sea. In the deeper parts of Calabozo Bay and the eastern side of the Outer Gulf, con- centrations of phosphorus occur which are too great to be accounted for by the physical circulation (Figs. 6 and 7). In the former case the high salinity excludes an origin from lake water; in the latter the high phosphorus content precludes Caribbean water as the source. It is concluded that organisms and particulate matter have sunk into these basins, carrying down from the surface layers the phosphorus which has accumulated at depth. Oxygen In the Gulf of Venezuela the surface water, as well as that at greater depths, was everywhere under-saturated, in amount varying from 1-4 to 9-6 per cent. Physically this implies that the oxygen pressure was positive across the surface, and that oxygen was diffusing downward into the water. This condition cannot be explained by the cooling and consequent under-saturation of the surface, as is the case in higher latitudes in winter (Redfield, 1948), because the annual range in temperature is too small. It must be attributed to biochemical effects, arising from the excess of respiration over photosynthesis in the water. The consumption of oxygen by respiration, in excess of its production by photo- synthesis, can only persist if there is available some external source of organic matter, as is the case in waters polluted by sewage. The undersaturation of the surface waters of the Gulf of Venezuela may be the result of the high organic content of the outflow from Lake Maracaibo. In the lake water two-thirds of the phosphorus (1 [J.gA/1) is present in organic form, and while in the Lake much of this is probably present in living organisms capable of photosynthesis. On introduction into the Gulf, however, these organisms, which are fresh water or brackish species (Gessner, 1953 a), may be killed by the higher salinity, and thus contribute to the quantity of organic matter immediately available for oxidation. In support of this suggestion are the observations that the greatest under-saturation of surface water occurs in the immediate offing of Tablazo Bay (Fig. 1 1), and that the water of that bay is generally under-saturated with oxygen. In the deeper parts of Calabozo Bay and the eastern parts of the Outer Gulf, where the total phosphorus was found to be anomalously high, the oxygen concentration is reduced to less than 3 ml per litre. (Compare Figs. 5 and 6). The deficiency in oxygen increases with the phosphorus content, and is about that to be expected from the increase in phosphorus if it arises from the oxidation of planktonic material (see Fig. 13). It may be concluded that the deficiency of oxygen in the deeper parts of the Gulf is due to the accumulation of organic matter as the results of the sinking of organisms and particulate matter from the surface layers of water, and that the greater part of this material has undergone oxidation. At no place was the deeper water found to be completely devoid of oxygen. The deep circulation appears to be sufficiently rapid to prevent the accumulation of enough The hydrography of" the Gull" of Venezuela 131 organic matter to exhaust the oxygen dissolved in the water. In the Strait of Maracaibo the water is flushed out each season by escaping lake water, to be replaced again with more saline water when the outflow slackens in the dry season. The oxygen content of the deeper water of the Strait is reduced to values comparable to those in the Gulf in the course of two or three months. Thus it is needless to assume that much more time is required to produce the conditions observed in the deeper parts of the Gulf. >- 3 X o \ \ \ \ \_ O GO ° \ \ \ • \ \ \ \ • \ \ \ \ \ \ \ \ \ \ • \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ • \ \ \ \ \ \ \ \ \ \ \ \ \ o • - • SECTION III - o SECTION V \ \ \ \ \ \ \ J_ \ 0.2 04 06 0.8 TOTAL PHOSPHORUS , /^^A/ \ \2 Fig. 13. Relation of oxygen and total phosphorus content of samples collected in Sections ill and V. The slope of the envelopes is that required by the complete oxidation of organic matter from plankton of average composition; i.e., 2-35 ml Oo is consumed in oxidizing organic matter containing 1 ;x gA phosphorus COUNTERCURRENT SYSTEMS AND BIOGENETIC PROPERTIES The major ions of sea water exist in all parts of the ocean in very nearly equal proportions. It is a remarkable fact that, in contrast, most of those components which enter into the structure and chemical activity of living organisms are distributed in very different proportions in different parts of the oceans. Phosphate and nitrate, for example, are more than twice as abundant in the deep water of the Pacific and Indian Oceans than in the North Atlantic, although the concentrations of the major ions are approximately the same. 132 Alfred C. Redfield The concentrations of the organic derivative of sea water (phosphate, nitrate, and carbonate), and of oxygen, have been shown to vary from place to place in proportions related to the statistical composition of the plankton (Redfield, 1934). It is evident that some broad-scale process of biological origin is responsible for the distribution of these biogenetic properties in ways which are anomalous in respect to the purely physical character of the circulation. The obvious mechanism for separating the organic derivative from the dissolved materials in one mass of sea water, and transferring it to another, is the sinking and subsequent decomposition of organisms into a deeper layer. Redistribution leading to accumulation or attenuation is then dependent on the horizontal movement of the respective layers. If such movements are consistently in opposite directions, great differences in concentrations may be developed. The counter-current principle is commonly employed in such physical mechanisms as heat exchangers. Its applications in physiology have been discussed by Scholander (1954). The Gulf of Venezuela affords examples of two somewhat different types of counter- current system which lead to the local accumulation of phosphorus, with attendant depletion of oxygen in the deeper water. In the Outer Gulf the winds appear to produce an offshore movement of the surface water, which is compensated for by an onshore counter-current at depth. This is referred to as upwelling. Organic matter sinking from the surface layer is carried landwards in the deep counter-current. The process leads to an attenuation of phosphorus content of the surface layer with distance from shore, and its augmenta- tion in the deeper water which increases as the coast is approached. The degree of accumulation finally developed depends, of course, on a balance between this process and the dissipating effects of the circulation, which mixes the water vertically or trans- ports it horizontally across the region of upwelling. The situation in the Outer Gulf provides a small-scale example of the mechanism of enrichment of ocean water which occurs wherever trade winds give rise to upwelling along the continental coasts. In Calabozo Bay the counter-current system of the estuarine circulation is engendered by the inflow of fresh water from Lake Maracaibo rather than by the wind. Otherwise the process leading to the attenuation of phosphorus in the surface water and its accumulation at depth is the same as in the upwelling system of the Outer Gulf. The degree of accumulation attained is limited by the rate of circulation of the deep water, which appears to move toward the head of the Bay, where it is most actively incorporated into the surface outflow. The highest phosphorus and lowest oxygen concentrations were found immediately off the outlet of the Lake, The estuarine circulation of Calabozo Bay provides an example of a mechanism which must operate to varied degrees in many coastal embayments, fjords, and estu- aries, and which may account in part for the greater fertility common to such enclosed waters, when compared to the off-lying sea. REFERENCES Carter, Douglas B. (1954), Report on water balance of the Maracaibo Basin. Unpublished manu- script on file, Woods Hole Oceanographic Institution. Corps of Engineers, U.S. Army (1938), Model study of channel improvements at outer bar, Lake Maracaibo, Venezuela. U.S. Waterways Experiment Station, Corps of Engineers, U.S. Army, Tech. Memo. No. 106-1, 3 vols, December 1, 1938. The hydrography of the Gulf of Venezuela I 33 Gessner, Fritz (1953 a), Aufdcn Spuren Alexander von Humboldts in Venezuela. Sciiur u. Wissen- schaft, 138 neue Zeitung, 20, 13, 14, Juni 1953. Gessner, Fritz (1953 b), Investigaciones hidrograficas en el Lago de Maracaibo. Acta Cientifica Venezolona, 4 (5), 173-177, 4 figs., 4 tables. Gessner, Fritz (1955), Die limnologischen Verhiiltnissc in den Seen und Fliissen von Venezuela. Verhandl. lutcnuit. Verein.Jiir theoret. unci atii^ewandle LimnoL, 12. 284-295, 12 Hgs. Harvey, H. W. (1948), The estimation of phosphate and of total phosphorus in sea water. J. Mar. Biol. Assoc, U.K., 27 (2), 337-359, 11 figs., 3 tables. Livingstone, D. A. ( 1954), On the orientation of lake basins. Amcr. J. Sci., 252 (9), 547-554. Marmer, H. a. (1951), Tidal datum planes. U.S. Coast ami Geodetic Survey, Spec. Pun. 135,, 142 pp. Redfield, Alfred C. (1934), On the proportions of organic derivatives in sea water and their relation to the composition of plankton. In: James Johnstone Memorial Volume, Univ. Press, Liverpool, 176-192, 5 figs., 2 tables. Redfield, Alfred C. (1948), The exchange of oxygen across the sea surface. J. Mar. Res., 7 (3), 347-361, 4 figs., 3 tables. Scholander, p. F. (1954), Secretion of gases against high pressures in the swimbladder of deep-sea fishes. II. The rete mirabile. Biol. Bull., 107 (2), 260-277, 5 figs., 4 tables. U.S. Coast and Geodetic Survey (1951), Tidal harmonic constants — Atlantic Ocean including Arctic and Antarctic Regions, 137 pp. U.S. Navy Hydrographic Office (1949), Sailing directions for West Indies, Vol. II, The Lesser .Antilles and the coast of Venezuela. H.O. Pub. 129 (6th ed.), 335 pp. Papers in Marine Biology and Oceanography, Suppl. to vol. 3 of Deep-Sea Research, pp. 134-138. The specific characters of the coral Sty/aster roseus By H. BoscHMA Rijksmuseum van Natuurlijke Historic, Leiden In the collections brought together by Dr. P. Wagenaar Hummelinck in the Leeward Islands and other parts of the West Indian region there are a few small colonies of a stylasterine coral with the following data: Curasao, Plaja Djerimi, North corner, December 11, 1948 (rock, sand; tidal and lower zone). Examination of these corals showed that in all their salient characters they prove to correspond with Stylaster roseus (Pallas) as far as the eighteenth-century data on the species are concerned, while differing from the specimens identified with this name in recent literature. The corals from Curasao are four or five colonies of a more or less fan-shaped growth, occurring together on a fragment of dead coral rock. The colonies are up to 3 cm high and not over 3 cm broad. Each colony consists of a few stems of a breadth of about 5 mm, rapidly tapering upwards while giving off side branchlets which in their topmost parts have a thickness of about 1 mm. The stems and the thicker branches are slightly compressed in the flabellar plane of the colony. On the upper parts of the branchlets the cyclosystems occur alternately on the two lateral surfaces, whilst on the thicker branches the cyclosystems are distributed without apparent order; they are more numerous on the anterior than on the posterior surface. The number of dactylopores in the cyclosystems varies from 7 to 12; this number was counted in 100 cyclosystems with the following result: 5 cyclosystems with 7 dactylopores; 12 with 8; 28 with 9; 37 with 10; 14 with 1 1 ; 4 with 12; yielding an average of 9-55. Except on the tops of the smaller branchlets the cyclosystems extend very little above the surface of the coral; their diameter varies from 0-5 to 0-7 mm. The gastro- pores have a depth of about 1 mm, and the gastrostyles have a length of 0-4 mm and a thickness of 0-15 mm, so that they are rather slender, occupying about one-third to one-half of the lower part of the gastropore (Fig. lb). The gastrostyles are covered with small spines on the whole of their surface (Fig. la). Feebly developed dactylo- styles (not drawn in Fig. lb) are present in the dactylopores. The colonies bear numerous ampullae of a hemispherical shape, scattered among the cyclosystems on the larger branches, or occurring in clusters densely covering large parts of the surface; as a rule the ampullae are numerous, especially on the posterior surface of the colonies. The ampullae have a smooth surface, without warts or spines, and their diameter varies from 0-6 to 0-8 mm, the size indicating that probably they are female ampullae. The colour of the corals from Curasao is yellowish with an irregular admixture of pink, occasionally turning to a light purplish tint in the topmost parts of some branch- lets or on and around some cyclosystems on the larger branches. Pallas (1766) gave a description of the coral named by him Madrepora rosea, mentioning some characters which at present still may be regarded as typical of the 134 The specific characters of the coral Stylaster roseus 135 species; this description was based on an examination of specimens in Dutch collec- tions. In Houttuyn's (1772) account of the coral, the salient data are mentioned as noted by Pallas; Houttuyn's work is here cited because it contains the first figure of the species (1772, PI. 129, Fig. 4, copied in Fig. 2 of the present paper). This figure r\ Fig. 1. Stylaster roseus (Pallas) from Curasao, (a) gastrostyle, cyclosystem, 60 250; (b) longitudinal section of a Fig 2. Copy of the figure of Madrepora rosea Pallas in houttuyn (1772, PI. 129, Fig. 4). Original size again appeared in a treatise by Muller (1775, PI. 23, Fig. 4), which practically is a translation of Houttuyn's work with a few additional remarks. Esper (1794) gave an elaborate description of the species, based on a specimen from a German collection, his figures (1794, Madrep., PI. 36) represent a coral of a shape similar to that of 136 H. BOSCHMA HouTTUYN. The data noted here contain the original contributions towards a know- ledge of the species ; remarks in other eighteenth-century publications were copied from the cited works. According to Pallas, the colonies of his Madrepora rosea •are " semi-palmares " (half a hand high, about 5 cm); Esper records the height of large specimens as " zwey bis drey Zollen " (about 5 to 1\ cm). The two authors further state that the basal parts of the stems are comparatively thick, and that they gradually taper to very thin terminal branchlets while giving off numerous side branchlets. At that time the only known locality of occurrence of the species was " Mare circa Insulam St. Domingo " (Pallas), " an den Kiisten der Insel Domingo " (Esper), the quoted passages referring to the eighteenth-century name of the island Haiti in the West Indies. It stands to reason that the corals which were collected nearly two hundred years ago came from shallow water. It is interesting to note that the corals from Curasao, on account of their small size and of their gradually tapering branches, closely correspond with the specimens described and figured by Pallas, Houttuyn, and Esper, thereby differing from the other species of Stylaster known to occur in the West Indian region, which attain a larger size and have branches of a much slenderer growth. Lamarck (1816) placed the species Madrepora roseus in the genus Oculina ; Gray (1831) erected the genus Stylaster, in which he placed the two species Oculina rosea and O. flabeUiformis \ in 1850 Milne Edwards and Haime (1850-1854) selected Stylaster roseus as the type of the genus Stylaster. In another paper (Milne Edwards and Haime, 1850) there is a rather elaborate description of Stylaster roseus, in which the authors state that they have never seen colonies of a larger size than a few centi- metres, the thicker branches of which have a thickness at the base of 5 to 6 millimetres. These data correspond with those of the corals from Curagao; in other details, however, there are slight differences: Milne Edwards and Haime remark that the number of dactylopores in the cyclosystems is from 10 to 14, most commonly 12, while the diameter of the cyclosystems is nearly 1 mm, these figures being somewhat higher than in the specimens from Curagao. The measurements of the ampullae in the specimens of Milne Edwards and Haime (not over \ millimetre) perhaps indicate that these were male ampullae. PouRTALES (1871, p. 83) remarks that Stylaster roseus is " abundant on the under surface of blocks of coral rock, on the reef at Cruz del Padre, north coast of Cuba, a couple of feet below low-water mark ". Moseley (1880, p. 79) refers to this occur- rence of the species in almost the same words, and on another page {loc. cit., p. 77) remarks that " ampullae are especially well developed in the shallow water Stylaster roseus; those in the female stocks being very large and prominent ". A coral of rather common occurrence in depths of 120 to 324 fathoms (220 to 592 metres) off the Florida Reef was described by Pourtales (1868) as Stylaster erubescens; with some slight changes this description again appeared in a later paper (Pourtales, 1871); the following details are taken from the latter, the terminology partly being altered to a more modern wording: younger branchlets slender, with rather close-set alternate cyclosystems, these slightly prominent, 1-2 to 1-5 mm in diameter, deep; nine to twelve, most frequently eleven dactylopores in a cyclosystem; gastrostyle deeply sunk, rounded and hirsute; dimensions, height and breadth of flabellum, 10 cm. On Plate 4 of the cited work is figured a colony of Stylaster erubescens in The specific characters of the coral Stylaster roseus \ 37 natural size (Fig. 10), and the terminal part of a branch, 4 times enlarged (Fig. 11). The figures show that the tendency for a lateral arrangement of the cyclosystems on the branches is much more obvious than in the specimens from Curasao, that the branches are much less pronouncedly tapering, and that the cyclosystems definitely extend over the surface of the branchlets, the smaller branchlets thereby becoming distinctly zigzag-shaped, in contradistinction to the corresponding parts of the specimens from Curasao. The cyclosystems of Stylaster erubescens (diameter 1 -2- 1 -5 mm) are about twice as wide as those of the specimens from Curasao (diameter 0-5-0-7 mm), and distinctly wider than in the specimens of Stylaster roseus examined by Milne Edwards and Haime (diameter nearly I mm). The most important difference of the two forms is that of the entirely different shape of the gastroslyle, which in Stylaster erubescens is rounded (almost spherical), in the specimens from Curasao conical, more than twice as high as broad (Fig. I). The Stylasteridae of the North Atlantic region remained imperfectly known till 1914, when Broch's important paper appeared dealing with the specific characters of Pliobothrus symmetricus Pourtales, Allopora norvegica (Gunnerus), Stylaster gem- mascens (Esper), and a species named by broch Stylaster roseus (Pallas). The material of the last named came from depths between 263 and 1,400 metres; Broch notes that the colonies display a marked difference between small branches, main branches, and stem, that the cyclosystems are placed laterally and alternately on the small branches, that the cyclosystems show from 8 to 17, generally 9 to 1 1 dactylopores, and that the gastrostyle is almost spherical, with equal height and breadth. The characters here cited from Broch's description are almost exactly those of the species Stylaster erubescens as described and figured by Pourtales (1871). Broch's figures of colonies in natural size (1914, PI. 1, Figs. 8 and 9, PI. 2, Figs. 10 and 1 1) represent corals with an entirely similar form of growth as the specimen of Stylaster erubescens of Pourtales (1871, PL 4, Fig. 10); moreover, the figures of enlarged terminal branches (Pourtales, 1871, PI. 4, Fig. 11 ; Broch, 1914, PI. 2, Fig. 17) are strikingly similar. Unfortunately Broch gave the name Stylaster roseus to the corals here dealt with, while placing the name Stylaster erubescens in the synonymy of the species. In Broch's paper there is an instructive figure of a longitudinal section (1914, PI. 3, Fig. 22) showing in two of the cyclosystems the spherical gastrostyles, which, judging by this figure, have a height and a breadth of about 0-3 mm. This figure has been copied in other publications (Broch, 1924; Kuhn, 1939) as a longitudinal section of a branchlel of Stylaster roseus (Pallas). Broch (1914, p. 15) remarks " As the species is the commonest Stylasterid in the Atlantic north of the equator, it is probably the same form that served as a basis for Pallas' description of Madrepora rosea "; this is right as far as the specific identity of the North Atlantic corals with Stylaster erubescens is concerned, but it was an error to identify this species with Stylaster roseus. Pourtales (1868, p. 136, footnote; 1871, p. 83) examined the two forms and referred to them as separate species, the one from shallow water, the other from deeper water only; it is to be regretted that he did not give a description of the specific characters of Stylaster roseus. The conclusion of the data dealt with above is that Stylaster roseus (Pallas), the type species of its genus, has a rather complicated history. The descriptions and figures of the various eighteenth-century authors contain some characters which may be considered typical of the species. In later years these older data were overlooked 138 H. BOSCHMA and the name S. roseus was given to corals which in reaUty belong to the species Sty/aster erubescens Pourtales. An examination of specimens from shallow water of the island Curagao could lead to additional characters, especially those of the gastro- style, proving that the species Stylaster roseus is distinct from all the later described species of the genus. REFERENCES Broch, Hj. (1914), Stylasteridae. Danish Ingolf Exped., Copenhagen, 5 (5), 1-25. Broch, Hj. (1924), Hydroida. In: Handbuch der Zoologie (Kiikenthal-Krumbach), Berlin & Leipzig, 1, 422-458. EsPER, E. J. C. (1794-1797), Fortsetzungen der Pflanzenthiere in Abbildungen nach der Natur mit Farben erleuchtet nebst Beschreibungen. Niirnberg, 1, 1-230. Gray, J. E. (1831), Description of a new genus of star-bearing corals. The Zoological Miscellany, London, 1-80. HouTTUYN, M. (1772), De Zee-Gewassen. Natuurlyke Historic . . . volgens het Samenstel van den Heer Linnaeus. Amsterdam, 1 (17), 1-613. KuHN, O. (1939), Hydrozoa. In: Handbuch der Pal aozoologie (Schindewolf). Berlin, 2A, 3-68. Lamarck, J. B. P. A. de (1816), Histoire naturelle des animaux sans Vertebres. Paris, 2, 1-568. Milne Edwards, H. and Haime, J. (1850), Recherches sur les Polypiers; 5me mem. Monographic des Oculinides. Ann. Sci. Nat., (3), ZooL, 13, 63-110. Milne Edwards, H. (1850-1854), A monograph of the British fossil corals. London, i-lxxxv, 1-322. Moseley, H. N. (1880), Report on certain Hydroid, Alcyonarian, and Madreporarian Corals pro- cured during the voyage of H.M.S. Challenger, in the years 1873-1876. Rept. Sci. Res., Challenger, ZooL, 2 (1), 1-248. MiJLLER, P. L. S. (1775), Von den Corallen. Des Ritters Carl von Linne . . . voilstandiges Natur- system. Nurnberg, 6 (2), 641-960. Pallas, P. S. (1766), Elenchus Zoophytorum. Hagae-Comitum, i-xxviii, 1^51. Pourtales, L. F. de (1868), Contributions to the fauna of the Gulf Stream at great depths (2nd series). Bull. Mus. Comp. ZooL, Harvard Coll., 1 (7), 121-141. Pourtales, L. F. de (1871), Illustrated catalogue of the Museum of Comparative Zoology at Harvard College, No. 4. Deep-sea Corals. Cambridge, 1-93. Papers in Marine Biology and Oceanography, Suppl. to vol. 3 of Deep-Sea Research, pp, 1.19-148. External metabolites in the sea By C. E. Lucas Marine Laboratory, Aberdeen With the exception of a hint or two before the turn of the century, by such prophets as Brandt and Nathansohn, it was not until the inter-war period tiiat marine workers began to turn their minds to the possible existence of more subtle ecological relation- ships than those imposed on organisms by their inanimate environment and by predators. It is relevant that Knorrich and Putter had postulated the direct food value of dissolved organic food substances in the sea, although this view appeared to be finally rejected by Krogh (1931); however, such possibilities are not the main subject of this paper, although they may once again demand investigation. But those who drew our attention once more to the other possible significance of dissolved organic matter deserve our gratitude. Quite the most senior of those now living was Henry Bigelow who said in his famous book Oceanography, among many other stimulating things, " as yet we know little of the inter-relationships of different species or groups of animals in the sea beyond the obvious fact that some prey on others, but we may be certain that in many cases inter-relationships of less obvious sorts are vital links in the animal economy " (Bigelow, 1931, p. 131, quoted by Russell, 1936). Before many years elapsed, it seemed clear that he was right and we now know that he was! For the volume in his honour, it is fitting to pay tribute to his foresight. It would not be proper on this occasion, however, to ignore the others: from Johnstone, Scott and Chadwick (1924)* to Allee (193l)t and Hardy (1935):}:. All were feeling towards the certainty of a new type of relationship in ecology, and particularly marine ecology, for which there was then all too little evidence (although it may now seem striking that it was around this very time, in 1929, that the late Sir Alexander Fleming was making his first observations on the antibiotic influence o'i Penicillium). It was the stimulus of such hints, and particularly that of working with Hardy when he was evolving his idea of " animal exclusion ", that led Lucas to gather an odd collection of references in support of a speculative theory of " non- predatory " relationships — based on the release and biological influence of metabolites ranging from " toxins to vitamins and hormones " (Lucas, 1938). This evidence ranged from the effects of external metabolites in the field of bacterio- logy, through some of the " mass physiology " experiments of Allee's school, to the * " Also, we are pretty sure that the plankton communities influence each other — that there are what we may call group symbioses on the great scale so that the kind of plankton which we may expect to be present in a certain sea-area must depend, to some extent, on the kind o{ plankton which was previously present." t " Once formed, aggregations of aquatic organisms condition the medium surrounding them by the addition of secretions and excretions, the nature and biological effect of which form one of the important problems of mass physiology." X Hardy postulated a " presumably chemical " basis for his theory of " animal exclusion ", and also speculated upon the '" biological history " of waters, e.g. the changes which may occur in water passing from regions of predominantly "free" phyto-plankton to regions of "■ imprisoned" phyto- plankton. 139 140 C. E. Lucas speculations of Akehurst (1931) about the temporal alternation of oil-producing and starch-producing algae in ponds, and some tentative experiments (Lucas, 1936) regarding " animal exclusion ". With these were coupled Allen and Nelson's pioneer demonstrations (1910) of the need for some accessory substances in diatom culture; the use of " soil solution " for more effective growth; and Gran's observa- tions (1931) on the relatively intensive growth of phytoplankton at the junction of two bodies of water. But more evidence was needed. More evidence did in fact exist, scattered widely through the field of biological research, and work during the war brought even more, particularly that associated with antibiotics. It thus became possible to review a wider field and to formulate the concept more precisely (Lucas, 1947, and from a more general ecological viewpoint, 1949). In the aquatic field reference was made, among others, to the very different growth-rates of diatoms cultured in different natural sea waters (Matsudiara, 1939; Harvey, 1939), and Harvey's associated investigations of the effects of various accessory substances on growth; to other examples of the influence of waters pre- viously containing living organisms (or their by-products) upon different organisms in culture (e.g. Levring, 1945); and to Fox's demonstrations of the occurrence of carotenoids free in natural waters and their deposits (e.g. 1944). When supported by the rapidly increasing knowledge from the various fields of microbiology, together with a wide range of other biological references, it seemed not unreasonable to come to conclusions along the following lines: " (1) It is characteristic of cells to liberate certain metabolites, and these are known in a variety of instances to have great influence as endocrines. " (2) It is now well known that a number of these potent metabolites are eliminated as secretions or excretions by the organisms themselves, and many other chemicals are eliminated which are not yet known to have any specific effects within the body. "(3) Particularly insofar as any of these metabolites are . . . parts of the environ- ment of other organisms, they may be expected to have immediate potency for many of them. . . . The term ' ectocrines ' has been suggested for such metabolites. "(4) More generally, however, . . . the capacity for adaptation of most organisms suggests that further differentiation between beneficial and antagonistic relationships would be likely to have developed between the producers and those affected. In the extreme instances escape, exclusion, or death must be expected on the one hand, and obligatory association (parasitism, symbiosis) on the other. " (5) Such processes are believed to be important in evolution, and they are con- sidered to mediate communal relationships in ecology, which is the contemporary aspect of evolution. They should be seen as part of the nexus which also includes physical and chemical relationships as well as those of prey and predator " (Lucas, 1949, pp. 353-354). If these seemed then to be rather premature speculations, they now have much more specific support. It is not possible to review the whole field here and this note is simply intended to bring together a few references indicating some of the lines along which progress is now being made. However, the suggestions could properly be regarded as stemming from, and partly supported by, the theory of " animal exclusion ", and evidence (Bainbridge, 1952) has recently been brought to bear against the only laboratory experiments (Lucas, 1936) specifically made to investigate that theory. It seems necessary, therefore, to mention Bainbridge's experiments first, External metabolites in the sea 141 and it is particularly appropriate that they were made by one who has worked with both Hardy and Lucas. Lucas concluded that his very preliminary experiments were not inconsistent with Hardy's theory and, indeed, appeared to offer support for it. The work had to cease and they went no further than that. Bainbridge's work was not only more detailed but much more precisely executed, and it represents a real contribution to our knowledge of the prey-predator relationship. His criticism of Lucas's " light and dark " experiments is that they simply reflected the tropism of the animals — a possibility discussed by Lucas. It is and always was relevant. For all that, the very weaknesses in Lucas's work and the greatly improved techniques in Bainbridge's are very relevant, and they may still mean that Bainbridge's results are not necessarily so critical of the theory of animal exclusion as they seemed at first sight. Bainbridge was careful to use only algal cultures aged not more than " a week or so " (his p. 391) after inoculation, whereas Lucas probably seldom used such fresh cultures in his work. Indeed, Lucas's cultures may, in the light of modern ideas (many of his experiments were made in 1934), have been approaching senescence at times and, whatever may be their potency during the early phase of a culture, we can now see that metabolites released during the later stages (e.g. Pratt, 1943) may well have been harmful and have acted more as deterrents than as attractions. This is, of course, far from certain and exploratory experiments on these lines would be useful. In any event, it is also relevant that, whilst Bainbridge found that most of his plankton animals were attracted by, or at least did not appear to avoid, the majority of the denser phytoplankton cultures, several were either neutral or were markedly avoided or even lethal. Two of the more " harmful " were flagellates. It is also significant that not only did most of his cultures influence the animals but, in several experiments, so did the culture fluids in the absence of the plant cells. The main conclusion is that his experiments demonstrated the release by plant cells of substances which were frequently attractive and occasionally repellent to many of his animals (Bainbridge, 1952, p. 429). Like Lucas, also, he found evidence of an optimum density of plant cells below and above which the animals were presumably either starved or poisoned. However, other evidence was accumulating. In the first place, there was the work of Lwoff's school (1943), which demonstrated convincingly the release of vitamins " into their environment by some micro-organisms, and the vital need for such vitamins by some other forms which are unable to synthesize them. Lwoff saw this as evidence of a progressive loss of physiological function during evolution, but it provided also a mass of evidence of mainly beneficial inter-relationships, mediated by the release ot metabolites potentially of communal significance. Next there is the work of Lefevre and his school (1952) which, while collectively reviewed under the title of " Auto, et heteroantagonisme chez les algues d'eau douce ", demonstrates also quite clearly the very real influence, both favourable as well as unfavourable, which one micro-organism may exert upon others through the mediation of its secretions or excretions. In brief, these workers concluded that the three groups of bacteria, algae and fungi, all have " Faculte d'elaborer des substances actives autoantagonistes, heteroantagonistes ou favorisantes. Specific.te des substances actives produites . . . Decharge rapide des substances accumulees par les cellules quand on les replace dans un milieu neuf Solubilite des substances actives dans 142 C. E. Lucas I'eau ou dans certains solvants organiques. . . . D'autre part, il n'est pas impossible que les substances actives produites par des Aigues se developpant massivement dans une collection d'eau aient une influence directe sur la multiplication et le developpe- ment des animaux aquatiques: Entomostraces, Insectes, Mollusques, Vermes et peut- etre meme Poissons " (Lefevre, Jakob and Nisbet, 1952). The limitations implied by the title of their paper and the wider relevance of the text (summarized on their pages 173-181) are not without interest, for there seems to have been a strong tendency on the part of various workers to anticipate unfavourable reactions rather than favourable ones in given circumstances; e.g. the review by McCoMBiE, 1953, mentions only the possibility of harmful effects of free metabolites,* while one or two workers, in referring to Lucas' papers, have noticed only his ref- erences to harmful effects. The fact is that one organism's " meat " may be another's " poison " in the ecological nexus, and terms such as " harmful " and " beneficial " can only be used in an immediate and limited sense. In this sense, much of the evidence de- monstrates the development of " favourable " relationships, although admittedly some of the most striking are unfavourable. One of the latter is instanced by the phenomenon of" red tide ", with its harmful effects on marine animals and unfortunate repercussions on man. Lucas instanced such effects of the secretions of plants or animals, and much more evidence has been accumulated since (e.g. Br0ngersma-Saunders, 1948). There is now no doubt that, even though in their more striking forms such phenomena can be regarded as abnormal, they are, in fact, far from unusual in a lesser degree and, further, they are mediated by the release of " toxic " substances, frequently by flagellates. Their nature and the more precise conditions which lead to their production in a mild or extreme form, are being intensively investigated in several laboratories. The major task now is to determine the nature and effects of some of the more significant metabolites in aquatic ecology, to trace their probably variable distribution in some natural waters, and to determine the conditions leading to their production. Several lines of work have recently been developed. In the United States, Pratt (1943) has produced clear evidence that Chlorella cells in culture release a growth inhibiting substance, whilst Rice (1954) has grown Chlorella vulgaris and Nitzschia frustulum (both fresh water algae) together, and demonstrated clearly that neither grows so satisfactorily in the company of the other as it does in pure culture (depending upon the size of the populations used). Each was similarly inhibited when grown in the culture medium of the other, after its cells had been removed by filtration, and both were also inhibited when grown in a culture medium prepared with pond water which had supported a dense growth of Pandorina before filtration. Again, the metabolites in solution could be absorbed in charcoal and removed by autoclaving, suggesting volatile substances. Harvey, at Plymouth, following on his pioneer culture experiments with growth substances, is now attempting to review all the evidence available so as to define more * Lucas drew attention to this tendency (1944 and later) in respect of the term " antibiotic ". He pointed out that not only were antibiotics necessarily favourable to those organisms, such as man, which are preyed upon by the object of antibiosis, but that the antibiotic might well prove beneficial to those organisms succeeding its producer in the ecological chain (just as Akehurst, 1931, had suggested the autotoxic secretions of one algae may be beneficial for its successors). Indeed, the ecological successors of the producers of antibiotics can only succeed by virtue of being adapted to the presence of the antibiotics or their degradation products, and there is already some evidence that this may be true even for a potent substance such as penicillin. Here is a possible theoretical basis for ecological succession. i External metabolites in the sea 143 rigorously than has previously been possible the precise requirements of phytoplankton organisms in nature and in culture. Meanwhile, Provasoli, Hutner, and their colleagues at the Haskins Laboratories, have also very rightly undertaken fundamental and precise experiments on the basic requirements for the growth of marine and fresh water micro-organisms. Provasoli and Pintner ( 1 953), in their review of the ecological implications of the nutritional requirements of algal flagellates, have brought together a large number of references to the need of various micro-organisms for growth factors and trace elements. To these they have added the striking results obtained in their own laboratory. The evidence ranges from the early demonstration by Hutchin- son (1943) of the actual existence of thiamin in natural waters, to the significant requirement of various forms for cobalamin (vitamin Bio) in pure culture within chemically defined media. Of particular importance is the rigorous technique, necessary and adopted, in their work. All the results to date demonstrate aspects of " non-predatory " relationships. As they say: " It is a reasonable assumption that if an organism requires a growth factor in vitro, then this metabolite or its physiological equivalent should be found in significant amount in the environment " (p. 845). " The water environment is the one in which metabolites are interchanged most efficiently. It is to be expected that the interdependent growth of the different groups of water organisms should sensitively reflect the excretion and consumption of metabolites. Undaunted by new intricacies, we should envisage all the possibilities in these relationships, and not hesitate to follow Lucas's lead in constructing theoretical frameworks upon which to hang data. In the present paper, only a few aspects of the nutrition of phytosynthetic forms are considered. It is possible, nevertheless, to state more definitely some of the interdependencies based upon ' external meta- bolites ': (1) the interchange of growth factors; (2) the lowering of inhibitory con- centrations of several major mineral nutrients, especially PO4; and (3) the preferential utilization of minerals, including trace metals, may condition waters, bringing their concentrations into the optimal zones for succeeding forms. The practical aim — to predict algal successions and blooms— may be achieved through a comprehensive knowledge of vitamin cycles as well as mineral cycles. An immediate problem is to trace the thiamine and cobalamin cycles " (p. 849). Droop (for example, 1954) is pursuing a similar course at Millport, in Scotland, and has shown the need of several marine flagellates* for vitamin Bj... LEW'iN,in Canada, has also found B12 essential for the growth of the alga Stichococcus in sea water (1954). The presence of free B12 in natural sea water has been demonstrated, and its distribu- tion is being examined in more than one laboratory. Droop is now developing an assay for it in sea water. Along a rather diff'erent line Fogg (1952) has suggested that some at least of the external metabolites (for instance, the polypeptides of blue-green algae) may further communal growth, via chemical linkage, by making relatively insoluble nutrients available in a form more suitable for assimilation. Turning now to the animal field, several workers have followed Allee in linking aggregations with the release of metabolites (for example. Cole and Knight-Jones, 1949, and Knight- Jones, 1950). Again, there was the demonstration by Allison and Cole (1935) that the feeding movements of barnacles can be correlated with the * Droop has now demonstrated this need in the diatom Skeletonema costattim (Droop 1955). 144 C. E. Lucas abundance of dinoflagellates in the water (with the postulate that this is mediated by a by-product) and the subsequent hint (Miazaki, 1938) of the effect of an alga upon the spawning of the male oyster. Now, Collier and his colleagues, at Galveston, Texas, have moved into this field from a rather different angle. Whilst studying the effects of industrial wastes on oysters, they deduced a generalized influence upon their pumping rates, which was found in due course to be correlated with the presence or absence in the water of a carbohydrate-like substance; this is either truly soluble or colloidal, and may attain densities in neritic waters of up to 25 mg/1 (Collier, Ray, Magnitski and Bell, 1953). During further work (Collier, 1953) they found that, along with tyrosine-tryptophane, these carbohydrates have a marked diurnal variation in abundance, and their production is associated with light and aeration — so that they are probably the by-products of plant growth. In addition, a rhamnoside (Wangersky, 1952) has been isolated from oceanic water, and particularly from " red tide " water up to quantities of 50 mg/l, whilst minute quantities of ascorbic acid and some other carbohydrates of very low molecular weight are being isolated for identification. The significance of such substances to the oyster may be two-fold, but it is at least clear that one of the substances in question is both widespread and very variable in quantity, and that it acts as a remarkably effective (and almost instantaneous) pumping stimulus. It would appear that the substance is also absorbed by the oysters (up to 50 mg per hour), although it remains to be seen whether such substances are of positive and substantial food value (see, for example, Korringa, 1949, and Jorgenson, 1952) — a possibility which would greatly have interested both PiJTTER and Krogh — or whether their role is limited to providing sensory stimuli and perhaps growth factors. At the moment, the chief point is that they " found a biologically active compound (or group of compounds) to which an organism would respond quantitatively ", and drew attention to the link between this work and the probable significance of external metabolites or " ectocrines ". Work being undertaken by Wilson, at Plymouth, is also relevant. For many years he has been concerned with the problems of breeding and growing planktonic larvae, particularly polychaetes. On the one hand he noticed that sands from some areas were more suitable for the settlement of polychaete larvae than others (e.g. Wilson, 1948 and 1953) whilst, on the other hand, he found that, with the passage of years, his success with rearing these and other larvae was tending to decrease (Wilson, 1951). Continued and painstaking investigations in various directions now seem to make it clear, however, that the suitability of sands for larval settlement must be determined by the existence on them of other forms of life, probably micro- organisms — " Organic material, living or dead, on the sand grains plays an important role in rendering a sand attractive or repellent to the larvae " (Wilson, 1954). Whilst we may not be so obviously concerned here with a free metabolite (although that is quite possible), we have to deal once again with the significance for living larvae of organic remains, and with their detection by these larvae. The other instance is of even more direct interest. Wilson associated his decreasing success with the now familiar change known to have taken place whereby the waters of the English Channel have been much less productive since 1930 than in the 1920s. Superficially, this was due to reduced phosphate content, but the possibility of more subtle factors remained, and comparative experiments were made by rearing poly- chaete and echinoderm larvae in (a) " local " Channel water collected in the Plymouth External metabolites in the sea 1 45 neighbourhood (as had been usual) and (b) water collected from the Celtic Sea, in which the typical plankton community is normally similar to that of the " local " water before 1931. The results were striking; good growth of polychaete and echinoderm larvae was obtained in the latter and only poor or deformed growth in the former. Appropriate combinations of experiments suggested the presence of a " beneficial " substance in the Celtic water, and its lack in the local water, rather than the existence of a harmful substance in the latter. More recent work has shown that Clyde water also tends to be more suitable than " local " Channel water and, although there are variations, the inferences seem to be unambiguous. In association, Wilson and Armstrong (1954) have shown that heating the waters for shorter or longer periods generally had an adverse effect on the larvae bred in them, thus suggesting the existence of a beneficial substance of a more or less volatile nature. Their analyses have not yet been carried to the stage of demonstrating any particular component or fraction which is responsible for the biological difference. Recently, at Aberdeen, Johnston has extracted various fractions of the organic matter in natural sea waters, and made some preliminary biochemical analyses. Certain of these were tested in bio-assay on a number of phytoplankton diatoms and flagellates (with results summarized by Johnston in his Table I, 1955, from a paper read to the International Council for the Exploration of the Sea in 1954). It was found that the growth of many was favoured and of some others hindered relative to controls, although, " since the tests were limited to one concentration, further (and perhaps different) instances of these effects would probably have been observed by testing a range of concentrations. One particular fraction was found to promote greater growth in 9 of the 1 1 species tested." The initial information so gained is providing a basis for the bio-assay of sea water samples from different areas, with a view to a preliminary labelling of such waters according to their physiological effects. Then, further attention will be given as far as possible to identifying some of the substances thus found to be of biological significance. Johnston tentatively discusses the possible value of such information to the fisheries worker. In all such experimental work, tribute should be paid to the surveys made by Russell (e.g., 1939) and Fraser (e.g., 1952). By distinguishing on biological evidence between water masses, which have often been indistinguishable on familiar hydro- graphical criteria, they are providing the experimenter and biochemist with invaluable clues. In conclusion, it can be said that, before the war, a few people were moving towards a conception of ecological inter-relationships which seemed likely to be an important complement to the already familiar relationships existing between the organism and its physical environment, and those between prey and predator. Limited observations in diverse fields suggested that these non-predatory relationships would be mediated by the release of metabolites of varying potency for other members of the community. Further work during the war made it possible to gather much more support for the suggestion, and it is now quite clear that dissolved or colloidal organic matter may be present in natural fresh and marine waters in greater quantities than was originally envisaged. Indeed, it is still uncertain whether these may not in themselves provide, for some forms, nutrients in the ordinary sense of the word*. It is also clear that within *See, for example, Morris, 1955. 146 C. E. Lucas this general heading of organic matter are included, sometimes in very minute quantities, substances whose effects within the community may not unreasonably be compared with those of endocrine metabolites within the body. It now seems necessary to believe that such substances play a considerable part in the growth of aquatic communities of bacteria, algae and fungi (we still have much to learn of the activities of the last in natural waters), so that the success of various organisms, and consequently their ecological succession, may be largely determined in this way. The next point is that we now have a number of instances in which such free metabolites affect not only the lives of protozoans (whose communal life may be expected to be very similar to that of the micro-plants, except in so far as their need for external metabolites can be expected to be much greater), but also those of higher animals such as worms, echinoderms and molluscs and probably crustaceans. We can confidently expect this list to be extended, probably even to include fish. For example, Hasler and Wisby (1951) have obtained most interesting evidence of the influence of different natural waters upon the movements of fish. They found that minnows were able to discriminate between the waters of two Wisconsin creeks after as little as two months conditioning, while cautery of their olfactory epithelia rendered them unresponsive to conditioning. It appeared, therefore, that olfaction was the principal, if not the sole, means of discrimination. They further demonstrated that the chemical response was not to carbon-dioxide in the creek waters, for example, but that the significant fraction was probably organic, in the usual sense, in that the minnows reacted to the distillate rather than the residue (vacuum distillation at 25° C), so that the existence of a volatile substance in the water can be anticipated. Here then is the essence of one reaction system on which " homing ", and perhaps other aspects of migration, might reasonably be based. Indeed, Hasler's preliminary tests suggest that salmon can detect such " odours " of streams and discriminate between them. In that instance, it would be necessary to imagine the salmon being conditioned during its early fresh water life to the " odours " of the " home " tributary. Hasler has also been able to demonstrate that minnows could respond to such odours after " forgetting periods " which were longer in fishes trained when young than in old ones. It may not therefore be so unreasonable to think of such reactions as even applying to the move- ments of other and wholly marine fish. Lastly, when mentioning this particular possibility as one of the possible uses to which such fundamental research might ultimately be put in a fisheries laboratory, Johnston (1955) also referred to the possibility that " fertilization " of natural waters, normally by enrichment with phosphates and nitrates, might come to include the addition of minute amounts of critical metabolites. Ideally, at least, these would be selected as those likely to mediate the succession of '' desirable " algae in relation to the favoured crop. For example, it has seemed at times that the risks of fertilization being followed by blooms of " undesirable " algae are considerable, but it does not seem unreasonable to imagine that such a succession could, to some extent, be con- trolled if the fertilizer included not only normal manures but a specific metabolite antagonistic to these algae and preferably favouring other forms. Indeed, it may not be too speculative to suggest that some natural waters, normally producing little in the way of a desirable crop, might be induced to undergo a complete ecological change by the addition of a critical metabolite alone. The apparent scarcity of nutrients in some of these waters may not be so significant as it may seem. Not only may there External metabolites in the sea I47 be great reserves of nutrient in the bottom deposits but, as the Haskins Laboratories have demonstrated, many algae can tolerate only quite dilute nutrient solutions, and can thrive on them, so long as the necessary free metabolites are present. Perhaps particularly in some " barren " fresh waters, it may be that certain types of ecological development are blocked, just as in some laboratory experiments, by the natural absence of a specific metabolite essential to the life of an essential organism in fish management. Such possibilities as these various lines of work indicate may make this field an attractive one to fisheries worker and biologist alike. Certainly, considerable develop- ment can be expected during the next few years. A number of those working in the various fields, however, may possibly have missed something of the community of interest they share with many others. With a few striking exceptions, what now seem to be very relevant cross references are frequently missing from bibliographies. Perhaps this paper may serve to emphasize, where necessary, some of the fundamental features thought to be common to the various investigations. REFERENCES Akehurst, S. C. (1931), Observations on pond life, with special reference to the possible causation of swarming of phytoplankton. J. Roy. Micr. Soc, 51, 237-265. Allee, W. C. (1931), Animal aggregations. Univ. Chicago Press, 431 pp. Allen, E. J. and Nelson, W. E. (1910), On the artificial culture of marine plankton organisms. J. Mar. Biol. Assoc. U.K., 8, 421-474. Allison, J. B. and Cole, W. H. (1935), Behavior of the barnacle Balanus balanoides as correlated with the planktonic content of the sea water. Bull. Biol. Lab., Mt. Desert Is., 24-25. Bainbridge, R. (1953), Studies on the interrelationships of zooplankton and phytoplankton. J. Mar. Biol. Assoc., U.K., 32, 385^47. BiGELOW, H. B. (1931), Oceanography. Houghton Mifflin Co., 263 pp. Cole, H. A. and Knight- Jones, E. W. (1949), The setting behaviour of larvae of the European flat oyster, Ostrea ediilis L., and its influence on methods of cultivation and spat collection. Min. Agric. and Fish., Fish. Invest., (2), 17 (3), 1-39. Collier, A. (1953), The significance of organic compounds in sea water. Trans. N. Amer. Wildlife Conf., 18, 463-472. Collier, A., Ray, S. M., Magnitski, A. W. and Bell, J. O. (1953), Eff'ect of dissolved organic substances on oysters. U.S. Fish and Wildlife Serv., Fish. Bull., 54 (84), 167-185. Droop, M. R. (1954), Cobalamin requirements in Chrysophyceae. Nature, 174, 520-521. Droop, M. R. (1955), A pelagic marine diatom requiring cobalamin. J. Mar. Biol. Assoc, U.K. 34, 229-231. Fogg, G. E. (1952), The production of extracellular nitrogenous substances by a blue-green alga. Proc Roy. Soc, (B), 139, 372-397. Fox, D. L. (1944), Fossil pigments. Sci. Mon., 59, 394-396. Eraser, J. H. (1952), The Chaetognatha and other zooplankton of the Scottish area and their value as biological indicators of hydrographical conditions. Scottish Home Dept., Mar. Res., 1952 (2), 1-52. Gran, H. H. (1931), On the conditions for the production of plankton in the sea. Rapp. Proc. Verb.. Cons. Perm. Int. E.xpl. Mer, 75, 37-46. Hardy, A. C. (1935), Part V. The plankton community, the whale fisheries, and the h\pothesis of animal exclusion. In: Hardy, A. C. and Gunther, E. R. (1935), The plankton of the South Georgia Whaling Grounds and adjacent waters, 1926-1927. Di.scovery Repis., 11, 1-456. Harvey, H. W. (1939), Substances controlling the growth of a diatom. J. Mar. Biol. Assoc, U.K.. 23, 499-520. Hasler, a. D. and Wisby, W. J. (1951), Discrimination of stream odors by fishes and its relation to parent stream behavior. Amer. Nat., 85, 223-238. Hutchinson, G. E. (1943), Thiamin in lake waters. Arch. Biochem., 2, 143-150. Johnston R (1955), Biologically active compounds in the sea. J. 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Vetenskaps- och Vitterhets-Samhdlles Handi, 3 (12), 1-18. Lewin, R. a. (1954), A marine Stichococcus sp. which requires vitamin Bio (Cobaiamin). J. Gen. Microbiol., 10(1), 93-96. Lucas, C. E. (1936), On certain interrelations between phytoplankton and zooplankton under experimental conditions. J. du Cons., 11, 343-362. Lucas, C. E. (1938), Some aspects of integration in plankton communities. /. du Cons., 13, 309-322. Lucas, C. E. (1944), Excretions, ecology and evolution. Nature. 153, 378. Lucas, C. E. (1947), The ecological effects of external metabolites. Biol. Rev., 22, 270-295. Lucas, C. E. (1949), External metabolites and ecological adaptations. Svmp. Sac. Exp. Biol., 3, 336-356. LwoFF, A. (1943), L'evolution physiologique. Actualites Sci. Industr., 970, H. Hermann, Paris. McCoMBiE, A. M. (1953), Factors influencing the growth of phytoplankton. /. Fish. Res. Bd., Canada, 10 (5), 253-282. Matsudiara, T. (1939), The physiological property of sea water considered from the effect upon the growth of diatoms, with special reference to its vertical and seasonal change. Bull. Japan. Soc., Sci. Fish., 8, 187-193. MiYAZAKi, I. (1938), On a substance which is contained in green algae and induces spawning action of the male oyster. (Preliminary note.) Bull. Japan. Soc, Sci. Fish., 7, 137-138. Morris, R. W. (1955), Some considerations regarding the nutrition of marine fish larvae. J. du Cons. 20 (3), 255-265. Pratt, R. (1943), Studies on Chlorella vulgaris. VL Retardation of photosynthesis by a growth- inhibiting substance from Chlorella vulgaris. Amer. J. Bot., 30, 32-33. Provasoli, L. and Pintner, L J. (1953), Ecological implications of w vitro nutritional requirements of algal flagellates. Ann. N. Y. Acad. Sci., 56, 839-851. Rice, T. R. (1954), Biotic influences aflfecting population growth of planktonic algae. U.S. Fish and midlife Serv., Fish. Bull., 54 (87), 227-245. Russell, F. S. (1936), A review of some aspects of zooplankton research. Rapp. Proc. Verb., Cons. Perm. Int. Expl. Mer, 95, 5-30. Russell, F. S. (1939), Hydrographical and biological conditions in the North Sea as indicated by plankton organisms. J. du Cons., 14 (2), 171-192. Wangersky, p. J. (1952), Isolation of ascorbic acid and rhamnosides from sea water. Science, 115, 685. Wilson, D. P. (1948), The relation of the substratum to the metamorphosis of Ophelia larvae. /. Mar. Biol. Assoc, U.K., 27, 723-760. Wilson, D. P. (1951), A biological difference between natural sea waters. J. Mar. Biol. Assoc, U.K., 30, 1-20. Wilson, D. P. (1953), The settlement of Ophelia bicornis Savigny larvae. J. Mar. Biol. Assoc, U.K., 31,413-438. Wilson, D. P. (1954), The attractive factor in the settlement of Ophelia bicornis Savigny. J. Mar. Biol. Assoc, U.K., 33, 361-380. Wilson, D. P. and Armstrong, F. A. J. (1954), Biological differences between sea waters: experi- ments in 1953. J. Mar. Biol. Assoc, U.K., 33, 347-360. Papers in Marine Biology and Oceanography, Suppl. to vol. 3 of Deep-Sea Researcli. pp. 149-168. A revision of Ernst Haeckel's determinations of a collection of Medusae belonging to the Zoological Museum of Copenhagen By P. L. Kramp Zoological Museum, Copenhagen Summary— The collection of medusae sent from the Zoological Museum of the University of Copen- hagen to Ernst Haeckel for identification comprised 231 numbers; from 166 of these the specimens are still in our collection and are the subject of this revision. Specimens from 12 localities were not identified with certainty by Haeckel; they belong to 9 different species, 6 of which are described in previous literature, whereas 3 species have been described after the publication of Haeckel's monograph. In the collection and the accompanying list 54 species are provided with generic and specific name in Haeckel's hand-writing. After the revision the actual number of species is reduced to 37. 26 species are designated as new and are represented by their type-specimens or cotypes; 5 of these may be retained as valid species, 3 of them with unaltered generic name; the remaining 21 of Haeckel's new species belong to 15 species previously described. Among the 28 species, which are not marked as new, 10 are synonyms, and 8 are erroneously identified. While Ernst Haeckel prepared his famous monograph, " Das System der Medusen ", which was published in 1879-80, he borrowed the whole collection of medusae belonging to the Zoological Museum of the University of Copenhagen. The Danish zoologist Japetus Steenstrup had organized a fruitful collecting of marine animals, particularly pelagic ones, by officials in Greenland (inspector Olrik and others) and captains and physicians on Danish merchant vessels (Andrea, Bang, Holboll, Hygom, etc.) on their journeys to Greenland, the West Indies, South America, India and China. The result was that, until the far-going oceanographical expeditions began with the cruise of the Challenger in 1873-76, the museum in Copenhagen possessed one of the greatest collections of marine animals in the world. The collection of medusae was sent to Haeckel in Jena accompanied by a detailed list written by Chr. Lutken with (1) provisional determinations by Steenstrup and LiJTKEN, (2) localities, (3) names of collectors, (4) an empty column, in which Haeckel wrote his own determinations of the species. The list, which contains 231 numbers, is still in our museum and is a document of great value, giving us the names of all the species in Haeckel's own hand-writing (see Fig. 1). The majority of specimens are also still in our collection. Many of them are in a fairly good condition, though they must be handled with care, because they are more or less brittle in consequence of their being preserved in alcohol. Many of the species described or recorded by Haeckel have been the subject of much discussion, and it has often been desirable to re-examine the original specimens. It is not to be denied that Haeckel's artistic temperament and fertile imagination sometimes led him to construct a detailed description and beautiful drawings from an ugly and mutilated specimen. But he also had excellent powers of observation, and when his descriptions and figures are based on living or tolerably well-preserved 149 150 P- L- Kramp Specimens, they are usually reliable, though sometimes one must wonder at what he has overlooked (e.g. in " Hippocrene platygaster " and " Thaumantias eschscholtzii ", see below). Occasionally some of the specimens in the Copenhagen collection have been re- examined, partly by me (Kramp, 1919; 1926; 1947, specimens from the northern Atlantic), partly by Cl. Hartlaub (1913, northern Pandeidae) and by G. Stiasny (1922 a, some East- Asiatic Scyphomedusae). Remarks on some tropical Atlantic forms will also be published by me in the near future {Atlantide Reports and Discovery Reports, in press). A revision of the whole collection may, however, be useful to future workers on medusae, and in the present paper I shall give a complete list of that part of the old collection which is still in existence. Fortunately, as seen from the list, none of the specimens, which have disappeared, belonged to species which are not represented in the preserved collection. The succession of the species follows the order in which they are mentioned in Haeckel's monograph and with Haeckel's generic and specific names as head-lines, with the exception of the Narcomedusae, none of which were finally determined by him. In square brackets [ ] is added the correct name of each species as found by the revision, in so far as it differs from the name given by Haeckel or in cases where this latter turned out to comprise two or more species. The figures in common brackets ( ) refer to the numbers in the list. It is very understandable that Haeckel, in his worship of beauty, was attracted by these pretty animals. His " System der Medusen ", followed immediately by his work on deep-sea medusae in the Challenger Reports (1881), is a mile-stone in the progress of our knowledge of the medusae, and this progress soon became rapid, especially when one great expedition after another was sent out to explore the oceans and their inhabitants. No wonder that the knowledge contained in Haeckel's works was soon considerably augmented, and the reliability of his apprehension of the observations, or even of the observations themselves, were considered open to doubt. In the present paper I hope to contribute to the removal of some of these cases of doubt. When the next monograph of medusae appeared in A. G. Mayer's " The Medusae of the World " (1910), H. B. Bigelow^ had just published his outstanding work on the medusae of the eastern tropical Pacific (1909) to be followed {provisionally until 1940) by many other papers, to which the student of medusae must continually refer for valuable information. ANTHOMEDUSAE Codonium princeps Haeckel. 1879, p. 13, PI. I, figs. 1, 2. [Sarsia princeps (Haeckel).] The description is based entirely on specimens in the Zoological Museum, Copen- hagen. Specimens are still retained from the following localities: (8 and 13) Greenland? Mus. zootom. Hafn. 3 specimens. (9) Davis Strait and Baffin Bay; Borch, 1859. 2 specimens. (10) Greenland; 1865. 7 specimens. (12) Umanak, Greenland; Fleischer, 1865 (Neotype). 1 specimen. (137) Godhavn, Greenland; Olrik, 1860. 12 specimens. (138) Greenland. 4 specimens. A revision of Ernst Haeckel's determinations of a collection of Medusae 1 5 1 No. 22 in the list, Greenland, H. P. C. Molllr, is determined by Haeckel as Sarsia g/acialis; this specimen really belongs to S. princeps. Haeckel's statement of the colour of the species was based on sketches made by H. P. C. Moller. These sketches are in our museum and they were drawn partly at Frederikshaab in 1839, partly at Godthaab in 1840; they evidently represent Sarsia tuhulosa and noi princeps, showing no trace of an apical canal. No. 11 in the list, taken north of the Faroe Islands by Steincke and identified by Haeckel as Codonium princeps, is unfortunately not in the collection. It is very improbable that this species should have been found in this southern locality. Since no type-specimen was pointed out by Haeckel, I designate No. 12, Umanak, Fleischer, 1865, as Neotype. Sarsia tubulosa 1879, p. 16. Specimens identified by Haeckel as Sarsia tubulosa are retained from the following localities: (15) Iceland; Steincke. (16) Bordeyri, northern Iceland; Steincke. 2 specimens. (17) Isafjord, Iceland; Mariboe, 1865. 2 specimens. (18) Faroe Islands; Steenstrup, 1844. 2 specimens. Steenstrupia galanthus 1879, p. 31. [Steenstrupia nutans (M. Sars)] No. 118 in the list, 49' N. 7" W., off the mouth of the English Channel, Hygom, 1857, is labelled by Haeckel Steenstrupia {rubra Forbes?). There are two specimens, belonging to S. nutans. Stomotoca pterophylla Haeckel. 1879, p. 52, PL IV, fig. 10. (168) 20' 36' N. 76" W., north of Cuba; Andrea, 1867. 3 specimens, one of which I designate as Neotype. Mayer (1910, p. 113) gives a new description of this species, slightly differing from that of Haeckel. Mayer presumes that certain errors in Haeckel's description are due to the state of preservation of the specimens. It is true that this may account for the absence of an apical projection and the finely serrate inner margin of the ring- canal. The so-called " Ocellarkolben " are rudimentary tentacle bulbs, and they have no ocelli; their number is as stated by Haeckel, and on the whole the description is in good accordance with the structure of the specimens. Pandea saltatoria 1879, p. 54. In the text of his monograph Haeckel only mentions the original specimen from Norway described by M. Sars (1835) as Oceania saltatoria {Pandea saltatoria Lesson 1843), a species which has never been identified with certainty, though Hartlaub (1913, p. 336) is inclined to think that it was an Aglantha. 152 P- L- Kramp In our collection is a specimen (No. 112 in the list) labelled by Haeckel Pandea saltatoria, 14° N. 25° W., west of the Cape Verde Islands, collected by Hygom in 1858. It belongs to Pandea conica Lesson. The number of exumbral nematocyst tracks corresponds to the number of tentacles. Conis cyclophthalma Haeckel. 1879, p. 55, PI. IV, fig. 1. [Oceania armata KoUiker.] (1 14) 36° 29' N. 2° 28' W., Mediterranean, near Gibraltar; Branner. (Haeckel as the result of misprints gives the longitude as 2° 23' W. and the name of the collector as Bramer.) This is the type specimen of C cyclophthalma, and we are fortunate to have it in the collection. Mayer (1910, p. 130) retains the species within the genus Conis beside C. mitrata Brandt. Hartlaub (1913, p. 342) has examined the type-specimen, and I can confirm his statement that it belongs to Oceania armata Kolliker. Thus Conis cyclophthalma is an obsolete name. Oceania sp. Haeckel (1879, p. 56) declares that the generic name Oceania is obsolete, but in the list some medusae are mentioned as Oceania sp. ? One of them, 28-33° N. 60-64° W., Hedemann, 1867, has disappeared, the others are in the collection and may be identified as follows: (88) 21° S. 57° E., east of Madagascar; Andrea, 1864, 2 specimens, 5 mm in height, with 8 tentacles ; they belong to Neoturris papua (Lesson). (126) 49° N. 7° W., off the mouth of the EngHsh Channel; Hygom, 1857. 1 specimen, belonging to Leuckartiara nobilis Hartlaub. Tiara pileata. 1879, p. 58. (Ill) 36° 17' N. 3° 27' W. Mediterranean near Gibraltar; Branner, 1869. 4 specimens, labelled by Haeckel: Tiara pileata = Oceania (coccinea). They belong to Pandea conica Lesson. Tiara conifer a Haeckel. 1879, p. 59. (142 and 143) Greenland; Olrik. 2 specimens. Haeckel's description is entirely based on these specimens; they belong to Catablema vesicarium (A. Agassiz) as already presumed by Hartlaub (1913, p. 315). Tiara reticulata Haeckel. 1879, p. 60, PI. Ill, fig. 11. [Pandea conica Lesson.] (104) 35° 31' S. 0° 51' W. Atlantic Ocean near Tristan da Cunha; Andrea, 1862. 2 specimens. These are the only specimens known, and they belong to Pandea conica Lesson, as already stated by Hartlaub (1913, p. 340). I MVI 1;M1 1 ! K|S , ^. , . ^ ^ a/ ^ -^V Z(IOl,(MW^M Ml ^i-.l M --^7^ y^-^ r *^* ' ^ KJ0BENHAVN. u. •J£l- ^-^ • /^^.A^ j^./i^r. '^^'-«-- u.<^^J..^^^ /^-^ .■ ■ f^.j Fig 1 Fascimile of one of the pages of the lisl ol medusae sent from the Zoological Museum of Copenhagen to Eknst Halckil. The right hall of the sheet contains the names of the species m Haeckkl s hand-wntmg. A revision of Ernst Haeckel's determinations of a collection of Medusae I 53 Turris digitalis. 1879, p.61. (44) 59° 20' N. 15"^ 47' W., between Scotland and Iceland; Rink, 1852. 7 specimens. (47) 58-59° N., between Iceland and Greenland; Rink, 1852. 1 specimen. (147) 59° 09' N. 16^ W., west of Scotland; Olrik, 1861. I specimen. These specimens were re-examined by Hartlaub (191 3. p. 329) and Kramp (1926, p. 94). They belong to Neoturris pHeata (Forskal). Catablema campanula. 1879, p. 63, PI. IV, figs. 4, 5. [Catablema vesicarium (A. Agassiz).^ Haeckel described this as a " nova species? ", indicating that it might be identical with Medusa campanula Fabricius, 1780. The description was based on the following specimens, which are still in our collection: (43) Greenland; Zimmer, 1856. 2 specimens. (144) Umanak, Greenland; Olrik, 1853. 7 specimens. Haeckel also referred to a coloured sketch by H. P. C. Moller; the sketch is in our museum, and it was drawn after a specimen taken near Frederikshaab in Green- land in 1839; like the preserved specimens it undoubtedly belongs to C. vesicarium. Catablema vesicarium. 1879, p. 64. In his book Haeckel only quotes the description of this species as given by A. Agassiz (1865) and says nothing about a specimen, which is in the collection, labelled by himself Catablema vesicarium : (45) Greenland; Moberg, 1857. Catablema eurystoma Haeckel. 1879, p. 64, PI. IV, figs. 6, 7. The description of this species was based on specimens in the museum of Copen- hagen and a coloured sketch by H. P. C. Moller, drawn at Qajartalik in Arsuk Fjord in southern Greenland. The species is undoubtedly identical with C. vesicarium (A. Agassiz), but the specimens are lost. They were taken in the following locality: (146) 67^ 35' N. 54' 10' W., in South Stromfjord, Greenland; Olrik, 1866. Cytaeis nigritina Haeckel. 1879, p. 74, PI. VI, figs. 2-5. [Cytaeis tetrastyla Eschscholtz." On a previous occasion (Kramp, 1953, p. 263) I have discussed the name o( this species. Haeckel applies the specific name tetrastyla only to the original form described by Eschscholtz, but refers the numerous specimens from the museum of Copenhagen to a new species,'c. nigritina. Mayer (1910, p. 133) tried to revive the name atlantica Steenstrup, 1837, but, as stated by me, this name was only found in Steenstrup's hand-written catalogue of the collection in our museum and was never published. ]54 P- L. Kramp The specimens in our collection are derived from the following localities: (72, 73) 14° N. 20° W., near Cape Verde Islands; Prosch. 15 specimens, type specimens of Cytaeis nigritina. (74) 8°30'N. 24° W., south of Cape Verde Islands; Andrea, 1872. 2 specimens, labelled Nigritina atlantica. (75) 31° 28' N. 29° 39' W., south of the Azores; Andrea, 1860. 5 specimens, labelled Nigritina sp. (76) 5° S, 28° W., off Cape San Roque, east coast of Brazil; Hygom. 2 speci- mens, labelled Nigritina sp. {polyblasta ?). (77) 23° 03' N. 31° 48' W., N.W. of Cape Verde Islands; Mathiesen, 1848. 2 specimens, labelled Nigritina atlantica. (78) 5° 31' N. 23° 15' W., south of Cape Verde Islands; Reinhardt. 1 specimen, labelled Nigritina atlantica ( ?). (80) 34° N. 34° W., S.W. of the Azores; Hygom. 1 specimen, labelled Nigritina {polyblasta'}). (81) 6° N. 22° W., south of Cape Verde Islands; Hygom. 3 specimens, labelled Nigritina (polyblastal). (89) Atlantic Ocean; Hygom, 1863. 1 specimen, labelled Cytaeis nigritina. Before the publication of his book Haeckel had evidently been much in doubt of the most convenient name of this species. The specific name polyblasta, applied to some of the specimens in the list, is found nowhere in the monograph; he may have thought about attaching this name to specimens carrying medusa buds. Cytaeis macrogaster Haeckel. 1879, p. 74, PI. VI, fig. 1. [Cytaeis tetrastyla Eschscholtz.] It is generally acknowledged that C. macrogaster is merely a synonym of C. tetra- styla, and an examination of the present specimens confirms this view. (83) 0° S. 29° W., Andrea, 1866. 2 specimens, designated as the types. (84) 1° 20' S. 26° 20' W. Andrea, 1863. 1 specimen. (85) 2°30'N. 24°W. Andrea, 1863. 4 specimens. (87) 1° 30' N. 24° W. Andrea, 1863. 3 specimens. All these localities are at a considerable distance N.E. of Cape San Roque on the east coast of Brazil. Margelis principis Steenstrup. 1879, p. 88, PI. VI, figs. 14-16. The specimens labelled Margelis principis belong to two different species: Bougain- villia principis Steenstrup, and B. superciliaris L. Agassiz. (3) Sandvaag, Faroe Islands; Steenstrup, 1844. The type-specimen of Margelis principis. (5) Davis Strait off Holsteinsborg; Olrik, 1859. 7 specimens. (6) Godhavn, Greenland; Olrik, 1860. 1 specimen. The specimens from the two last mentioned localities are Bougainvillia superciliaris. No. 50 in the list, north of Orkney Islands, collected by Olrik, 1859, is likewise identified by Haeckel as Margelis principis, but the specimens are not in the collection. A revision of Ernst Haeckel's determinations of a collection of Medusae 1 55 Haeckel's figures of Marge/is principis are said to be based on specimens from Iceland, but no Icelandic locality is mentioned in the list. Hippocrene platygaster Hacckel. 1879, p. 91. [Bougainvil/ia platygaster Haeckel.] (107) 25= 04' S. 27° 26' W., S.W. of the islands of Trinidad; Andrea, 1869. 2 specimens, one of which 1 have designated as the type; by Haeckel himself named H. platygaster n.sp. (1 10) 24° N. 33° W., about 700 miles N.W. of Cape Verde Islands; Iversen, 1871. 1 specimen, labelled Marge/is (platygasterl). In my report on the Hydromedusae of the Discovery expedition (in press), I have dealt with this species at some length. Numerous specimens were collected by the Discovery in the tropical parts of the Atlantic and off the east coast of Africa, and it was also taken by the Dana in some localities in West-Indian waters. In my opinion BougainviUia platygaster is a valid species, distinct from B. carolinensis (McCrady), B.fulva Agassiz and Mayer, and B. niobe Mayer. I found that B. platygaster exhibits a most peculiar form of asexual propagation and, as a matter of fact, this same form of propagation is also observed in one of Haeckel's own specimens (No. 107), but it seems to have escaped his attention; at any rate, he does not mention it in his description. Hippocrene superciliaris. 1879, p. 92. [BougainviUia superciliaris L. Agassiz.] (1, 2 and 4) 66° 13' N. 55 05' W., off South Stromfjord, Greenland; Moberg. 12 specimens. These specimens really belong to B. superciliaris, as also some of the specimens erroneously identified as B. principis (see above). Nemopsis heteronema Haeckel. 1879, p. 93, PI. V, figs. 6-9. [BougainviUia principis Steenstrup. , (7) Iceland; Steenstrup, 1839. 2 specimens. Haeckel described these specimens together with some others from Norway collected by himself. As previously pointed out by me (Kramp, 1926, p. 48; 1939. p. 6) the specimens from Iceland belong to BougainviUia principis (Steenstrup). Rathkea fasciculata. 1879, p. 97. [Kollikerina fasciculata (Peron & Lesueur).] (151) Mediterranean; Keferstein and Ehlers. In the list as well as on the label this specimen is named Lizzia kollikeri Gegenbaur, but in the monograph the name is altered to Rathkea fasciculata. [56 PL. Kramp LEPTOMEDUSAE Thaumantias eschscholtzii Haeckel. 1879, p. 129, PI. VIII, fig. 4. [Tiaropsis multicirrata (M. Sars).] (56) Greenland; Holboll, 1841. 1 specimen. (58) Greenland, 1865. 2 specimens. The description of this species was one of Haeckel's great mistakes. The beautiful figure is reproduced in several handbooks as a typical " Thaumantias ", a leptomedusa, without any kind of marginal sense organs. As a matter of fact, it has eight large, black ocelli adjacent to eight large, open marginal vesicles. When as a young man, in 1915, I commenced my work at the Zoological Museum of Copenhagen, I found the specimens mentioned above, and of course I was very interested to see the type specimens of the famous Thaumantias eschscholtzii, but while looking at them through an ordinary hand lens I saw eight black spots on their umbrella margin, and a closer examination revealed the fact that they belonged to the common and well-known Tiaropsis multicirrata. This surprising discovery was published by me in the reports of the Danish Ingolf-Expedition (Kramp, 1919, p. 78; see also my revision of the Mitrocomidae, 1932, p. 364). To my regret " Thaumantias eschscholtzii " with a reproduction of the usual figure reappeared in Kukenthal's " Handbuch der Zoologie ", Bd. I, 1924. The author of the article on Hydroida, my friend Hj. Broch, Oslo, told me that his manuscript was delivered before 1914, and when the printing of the " Handbuch " was resumed after the war, he was not allowed to alter anything in his original text. Since then, however, I think that " Thaumantias eschscholtzii " has been regarded as an obsolete name. Staurostoma laciniata. 1879, p. 130. [Staurophora mertensi Brandt.] (164) 43' N. 61° 30' W., near Nova Scotia, Canada; Hedemann. Fragments of about 5 specimens. The specimens were labelled Staurostoma laciniata Agassiz, and in the monograph it is placed among the Thaumantidae, whereas Staurophora mertensi Brandt, which is the same species, is mentioned (p. 149) under the Cannotidae. Laodice cruciata. 1879, p. 132. [Laodicea undulata Forbes & Goodsir.] (53, 54) 59' 07' N. 13° 32' W., north of Rockall; Moberg. 5 specimens, labelled Laodice cruciata. (140) 59° N. 18' W., between Iceland and Rockall; Olrik. 3 specimens, labelled Laodice cruciata. (165) 58-59° N. 13-15° W., north of Rockall; Rink, 1852. 5 specimens, labelled Thaumantias (pilosellal). As demonstrated by Browne (1896, p. 482) there is only one single name among Haeckel's 25 synonyms of " Laodice cruciata ", which really refers to a Laodicea, viz. " Thaumantias mediterranea " Gegenbaur (one of Haeckel's synonyms was A revision of Ernst HaeckeFs determinations of a collection of Medusae | 57 Thaumantias (Cosmetira) pilose/la Forbes, which is a Mitrocomid). I have previously discussed the specific name (Kramp, 1919, p. 21) and adopted the name umlulata Forbes & Goodsir, which was proposed by Browne (1907). Orchistoma steenstrupii Haeckel. 1879, p. 139, PI. XV, figs. 3-5. [Orchistoma pileus (Lesson).] (154) 20°N. 81° W., south of Cuba; Hygom. 3 specimens. Two of the specimens are in good condition, and they agree quite well with the new description and figures by Mayer (1910, p. 211, PI. 25, figs. 1-4), which were based on specimens from near the Bahamas and Tortugas. I agree with Mayer that O. steenstrupii is identical with Mesonema pileus Lesson (1843) and should be called Orchistom