o-^T^^. ]930 Gift of Elaine Gomberg February^ 1984 In Memory of Her Sister Judy Joye ru oi 5 wD : CD I ° I r^ i CD ! m □ MARINE LIFE CARLETON RAYand ELGIN CIAMPI '^€tuide ie MARINE LIFE BIOLOGICAL LABORATORY LIBRARY WOOK HOLE, MASS. W. H. 0. L A. S. BARNES and COMPANY NEW YORK © 1956 BY A. S. BARNES AND COMPANY, INC. All rights reserved. No part of this book may be reproduced in any form, either wholly or in part, for any use whatsoever, including radio and television presentation, without the written permission of the copyright owner, except by a reviewer who may quote brief passages in a review printed in a magazine or newspaper. Manufactured in the United States of America. PUBLISHED ON THE SAME DAY IN THE DOMINION OF CANADA BY THE COPP CLARK COMPANY, LTD., TORONTO LIBRARY OF CONGRESS CATALOG CARD NUMBER 56-5558 ACKNOWLEDGEMENTS The comprehensi\'eness of this book, by its nature, leaves us greatly in debt to sexeral authors and publishers for their help and gracious cooperation in grant- ing permission to reproduce drawings. The figures of plants, chapter 6, were largely adapted from Marine Algae of the N orthwestern Coast of INorth America by W. R. Taylor, University of Michigan Press and Marine Algae of the Mon- terey Peninsula, California by G. M. Smith, Stanford University Press. The drawings of sharks, chapter 8, were partly reproduced from Part I of Fishes of the Western North Atlantic by H. B. Bigelow and W. C. Schroeder through the courtesy of the Sears Foundation for Marine Research, Yale Uni\'ersity, and from The Plagiostoma by Samuel Garman through the courtesy of the Museum of Comparative Zoology, Harvard University. Many of the drawings of fishes, chapter 9, were redrawn from The Fishes of North and Middle America by D. S. Jordan and B. W. Evermann, a Bulletin of the United States National Museum. The figures of turdes, chapter 10, were based on illustrations from The Handbook of Turtles by Archie Carr, Comstock Publishing Associates. Those of mamm.als were pardy based on illustrations from The Field Book of North American Mammals by H. E. Anthony, Putnam's Sons. All of the photographs except two by John A. Moore of Columbia University were taken by the authors with the aid of Russell Kinne. It would not have been possible to procure these photographs without the very kind cooperation of Jim O'Doherty of Robinson-Hannagan Agency, New York, Don McCarthy of the Bahamas Development Board, Nassau, and Abercrombie and Fitch Co., New York, for the loan of the compressor shown in figure 15. We also wish to thank Peggy and Dennis Hickman of the Pilot Flouse club, Nassau, and their surprised guests who were most understanding when they discovered that we were using their swimming pool as a specimen tank. Nora Wheatlv and Jean Wilmot were a great help in the typing of manuscript and the correction of galleys. The text was improved through the advice of the following: animal behavior, chapter 2, Dr. Helmut E. Adler and Dr. Evelyn Shaw, both of the American Museum of Natural History in New York; photography, chapter 3, Peter Stack- pole of Life Magazine; evolution, chapter 5, Dr. John A. Moore of Columbia University; plants, chapter 6, Dr. F. G. Lier of Columbia University; reptiles, chapter 10, Dr. Richard Zweifel of the American Museum of Natural History; mammals, chapter 10, Dr. H. E. Anthony and Dr. J. C Moore, both of the American Museum of Natural History. CONTENTS INTRODUCTION ix SECTION I. ^/le .9^ Chapter 1. ZONES OF THE SEA- Where They Live 1 Chapter 2. BIOLOGY OF THE SEA-How They Live 26 Chapter 3. MAN AND THE SEA I-Photography and Equipment 53 Chapter 4. MAN AND THE SEA II-Dangers and Chumming 62 Chapter 5. EVOLUTION, NAMES, AND CLASSIFICATION 68 SECTION II. ^lo^a a^d SSc^t^e^ ^^uuna Chapter 6. PLANTS OF THE SEA 76 Chapter 7. THE INVERTEBRATE LEGIONS 86 SECTION III. fjd^imii/6 nfil^t ^acA^tte^ : ^te ^f^^^eS^aie^ Chapter 8. THE LOWER FISHES-Lampreys, Sharks, Rays, and Ratfishes 132 Chapter 9. MASTERS OF THE WATER-Bony Fishes 177 Chapter 10. THE RE-ENTRANTS 303 BIBLIOGRAPHY 323 INDEX 329 vii INTRODUCTION The sea for thousands of years has been the least explored part of the earth. Now, with increased interest in the sea and the development of new and growing technology for its study and utilization, great esthetic and material rewards have been made a\'ailable to mankind. The sea has become for many men a new source of esthetic enrichment because of its primiti\'eness. It is also a vast reservoir of food and minerals, and this important part of the earth's resources is becoming more so because of the ever-growing problem of over- population—a problem which se\'eral atomic scientists have called more serious than the threat of the atomic bomb. If the sea is to continue to plav its important role in solving this problem, world-wide and local conservation programs must be enacted. Man's relationship with the sea is the same today as that of the early American pioneers with the undiscoveired wilderness. They believed that the land had unlimited productivity. But the rapid consumption of land resources without any thought of conserving the supply has resulted in great decimation of one-time plentiful forests and wildlife. Even James Audubon did not believe that the passenger pigeon of our plains, formerlv one of the world's most numerous birds, could ever become extinct. Flocks of millions once filled the skies, yet today not one remains. Similar stories of decimation have been repeated in the cases of the American bison, pronghorn antelope, bighorn sheep, beaver, many species of ducks, and valuable timber trees such as southern long-leaf pine, sugar pine and several hardwoods. These examples are onlv a small fragment of a long list of man's abuse of the land. We must avoid the careless and tragic exploitation of the sea which we have not avoided in the case of the land. Behind this exploitation lies one of the commonest beliefs of modern man— that the sea is inexhaustible. It is absolutelv vital to realize that the sea, like the land, has its limits and that the exploitation of the sea beyond its ability to replenish itself can onlv result in the situation that exists on much of the land. Already there are signs of the exhaustibilitv of the sea. Whales, thought to be innumerable at one time, were so scarce before World War II that manv whaling fleets ceased operations. Since the war, international control has resulted in a slow increase in their numbers. Sea otters, fur seals, sea turtles, striped bass, salmon, shad, and others all were, or still are, exceedingly important economically and have all suffered severely from exploi- tation. Commercial catches of manv important species of schooling fishes are IX X INTRODUCTION smaller todcty than previously in spite of the introduction of scientific fishing methods. The sea is also used as a gigantic dumping ground for industrial chemicals, sewage, garbage, oil, and atomic wastes. There is a widespread feeling that these things thrown into the sea are safely and completely disposed of. But they return to plague us in the form of reduced commercial fisheries, exterminated game fishes, littered beaches, poisonous shellfish, and polluted waters. For example, several New England rivers at one time had runs of Atlantic salmon which compared favorably with the runs of Pacific salmon on the west coast, but now, because of overfishing and pollution, there are few salmon south of Maine. These are the abuses that conservationists work to prevent in order to main- tain the long-term esthetic and economic resources of the earth. Conservationists are not the wide-eyed dreamers they are frequentlv thought to be, interested merelv in preserving remnants of rare fauna and flora. Although preservation of rare species is part of conservation, they are mainly concerned with the theme first popularly advocated by Theodore Roosevelt— "conser\'ation through wise use." There are two basic concepts per\'ading the philosophy of western man that have been serious barriers to the development of a realistic conservation pro- gram. The first is the economic conviction that the only possible healthy econ- omy is one which is constantly expanding— more and more production with little thought gi\'en to resource limits and replenishment. Second is the primarily theological dualistic view of the world wherein man and nature are somehow separated— man rules the world by a sort of egotistical "divine right of kings." Both of these concepts are in direct contradiction to the known facts that man and nature are one and that man cannot abuse nature without harming himself in turn. I do not see a planet concreted over its land surfaces to provide living and factory space for an immense human population nourishing itself and sup- plying all its needs synthetically. Human life is going to depend on plant life for a long time yet, and in far more ways than its value as an efficient conversion agent . . . man is still adolescent as a conscious species and he is faced, again in full consciousness, with the choice of whether he shall mature into a species loving his world or remain irresponsibly wielding his new toy of an expanding economy. F. Fraser Darling American Scholar, Winter 1956 >(■ H- 1- 1- Spearfishing and Skin-diving: A Progra)); for the Putiire Such is oftenest the young man's introduction to the forest, and the most original part of himself. He goes thither at first as a hunter and fisher, until at last, if he has the seeds of a better life in him, he distinguishes his proper objects, as a poet or naturalist it may be, and leaves the gun and fishpole behind. H. D. 1 horeau — Walden INTRODUCTION xi Wc have noticed a gradual transformation in some divers which invoh'es a change from a spearfisherman to a naturahst, photographer, or explorer. As the spearHsherman moves about the reefs looking for his prev, he becomes aware of the beauty and variety of marine life. It soon becomes unimportant whether or not he spears anything. His reward is in viewing nature and in gaining an understanding of it and of himself. He also gains an appreciation and respect for the order of things in nature and develops a feeling for not wanting to disturb it. Uncontrolled spearlishing, on the whole, has the two major disadvantages of rapidly depleting the number of fishes in an area— even more rapidly than the net or hook in most cases— and of driving fishes that are not killed into hiding at the sight of man. It doesn't take lono for a diver to recognize an area where spearfishermen have been. There is an air of desolation as evidenced by the empty reefs, and the few large fishes that are present are exceedingly wary. In sharp contrast, reefs which have not been invaded with spears have yielded some of our most interesting experiences underwater. Most of the fishes show little fear and have given endless opportunities for study of their fascinating behavior and for photography. The same contrast exists between areas where the over-enthusiastic shell or coral collectdr has been and those reefs which are untouched and unbroken. We feel there is a definite need for a program which will maintain and conserve marine life so that it will not be depleted beyond its ability to replenish itself. Only through a sound conservation program can skin-diving, either for esthetic or sporting values, be maintained as an expanding and long-range acti\'ity. A reef can not be cleaned out of fishes in one summer and be expected to ha\'e replenished itself the following year. It takes some species many years to reach maturity and large size and if the sport diver wants to see fish, he can not take them without any thought of conser\'ing the supply. We suggest the following program: 1. Underwater Parks Some of the richest areas should be set aside and protected as are "wilderness " areas on land. The taking of any marine life, animal or plant, by any means, should be prohibited. The areas selected should be distributed widely so that they could act as reproductive centers from which non-protected areas would be replenished with life. These areas, would be open to skin divers, naturalists, photographers, and all those who come to enjoy the wonders of nature. The benefits derived from undersea wilderness areas are as follows: a) Areas of replenishment. Game and commercial fishes can replenish their numbers and stock non-protected, surrounding areas, insuring a constant supply. This guarantees continued and growing interest of sportsmen upon whom large sporting industries and tourism are dependent. b) Sanctuary. Common and rare species would be protected from extinction. c) Esthetic values. The natural, virgin beauty of sea gardens unspoiled by broken corals or fishes made scarce and wary by spearing is preserved. Interest in natural beauties of the sea is maintained and expanded and these areas would become valuable tourist attractions. xii INTRODUCTION d) Science and education. Parks serve as natural outdoor laboratories for accumulating knowledge of the sea, for investigating methods of increasing fisheries' productivity, for formulation of better conservation policy, and for basic scientific research. 2. Regulation of Spearfishing The pressures on fish populations exerted by increasing numbers of spear fishermen have rendered many areas not worth visiting by the sightseer or spearfisherman. Controlled spearing, however, will not diminish the population of an area because it serves to take off some of the individuals that would normally have died of natural causes. a) Licensing. Spearfishermen should be licensed in order to maintain control. Funds derived should be used solely for conservation and enforcement. b) Regulating catch. Game fishes should be designated and for each of these, season, bag, and minimum size limits should be established. The spearing of small, non-game fishes such as angelfishes, butterflv fishes, etc., should be pro- hibited. c) Designation of areas. Spearing should be restricted to outlying areas be- cause, in general, spearfishermen are better divers than sightseers are, and because any amount of spearing tends to make fishes become warv or drive them off to deep waters. This would leave easily accessible waters for non-spear- fishermen such as skin-divers, sightseers, naturalists, and photographers. Whether our inshore waters remain filled with life and a source of enjoyment for present and future generations, or, whether they go the way our now com- paratively barren forests and plains have gone, depends upon the instigation of a conservation program immediatelv. How to use this Book This book is divided into three sections. The first section consists of chapters on the sea as an environment for life, how life adapts to this environment, dangers that exist in the sea for man, underwater photography, and evolution. These chapters are included in order that the reader will better understand the sea and its life and should be read before attempting to identify species or groups or to interpret behavior. The last two sections are a guide to the identi- fication and habits of groups of marine plants, invertebrates, and vertebrates. Because of the huge numbers of species in the sea (400 basses alone, for in- stance), each group (family, order, etc.) is represented by only a few of its commonest species which characterize the group as "types." Therefore, the reader will not always be able to identify a particular species with this book, but, by becoming familiar with the characteristics of a group, he can categorize any species of that group he may come across anywhere in the world. Thus, the ulua of Hawaii can be identified as a species of jack, the merou of the Medi- terranean as a grouper, and Japan's tai as a porgy. Identification of groups is made by characteristics recognizable in the field— silhouette, pattern, mo\'ement, etc.— and not by laboratory methods such as fin counts or fine anatomy. Though this book emphasizes groups rather than species, it is important that the reader refer to a particular species in his observations of behavior. All of the species given as examples of groups are found in North American INTRODUCTION xiii waters chiefly o\'ei" the continental shelf. Many of the species given, however, range to deep waters and bevond North American seas. Many of the pelagic species are world-wide in distribution. Because of the tremendous scope of the subject of the sea, the reader is asked to refer to the bibliography for detailed information. There is a great body of unknown facts that only the swimmer can procure because of his intimate contact with the sea. In 1933, Beebe and Tee-Van wrote that ". . . not one complete life history of a Bermuda fish is known." That statement is larpcly true today of marine fishes exervwhere. This guide has been written in response to the growing interest in the sea and its life in the hope that better knowledge of the sea and its relationship to man will result. Carleton Ray and Elgin T. Ciampi April, 1956 SECTION ONE CHAPTER /J ZONES OF THE SEA— Where They Live The study of mankind— his behavior, his diseases, his. history— could not progress very far nor be very meaningful if careful attention vere not paid to his environment— where he grew up and under what circumstances, the climate, culture, geographical location, and economic conditions. The same is true of wild life. Whatever aspect we care to study, scientific or aesthetic, a considera- tion of the environment must enter our minds. If one tries to imagine a plant or animal without environment, it becomes almost impossible to obtain much insight into its nature, whether it be a one-celled animal or a complex animal such as man. For instance, the odd shape of a sargassum fish has little meaning by itself, but when it is viewed from the perspective of self-preservation in its habitat of sargasso weed, its meaning as a concealing, protective form becomes clear. The study of the relationship between life and its environment is scientific natural history or ecology. Ecology might be called the crossroads of the life sciences, where such fields as genetics, paleontology, anatomy, physiology, embryology, animal behavior, oceanography, evolutionary study, and others find common ground. But let none assume that ecology is for the scientist only. The diver who has learned to associate groupers with their homes in coral or rock has made an ecological observation. Ecology is the subject of the first two chapters of this book in the hope that as insight into the sea's environment increases, so will our ability to appreciate the beauties and wonders of tfte sea increase. In this chapter the main character- istics of the sea and its zones will be given. The ways and means that life has adapted to this environment will be discussed in the chapter which follows. EARTH AND THE SEA The perspectives and visions of mankind are often not as broad as we would like to believe. Much of the narrowness of man's particular world is a con- sequence of the evolutionary place he has come to occupy— that of a giant land 2 UNDERWATER GUIDE TO MARINE LIFE animal. His size has made it extremely difficult for him to project himself intc the worlds of the small living things that swarm about him and has restrictec his vision to include mostly the forms of life that are, like him, giant. fiis terrestrial point of view has caused him to make a fundamental mistake in his conception and naming of the planet on which he lives, for this planet is not one characterized by dry land nearly as much as by the sea. No other planet, as far as is known, has a sea. Had man been aware of this from earliest time, he might well have named this planet "Sea" rather than "Earth." The sea is not only the most unique characteristic of earth, but it clearly dominates the land in surface area covered. About 70 per cent of the earth's surface is covered by the sea, an area encompassing about 139 million square miles. By contrast, all of the lakes and rivers add up to only about one million square miles. The sea averages about two and one-half miles in depth, whereas the land rises an average of only slightly more than four-tenths of a mile above sea level. The volume of sea water is about 334 million cubic miles, a volume so great that it has been estimated that if all the earth were made level, the sea would everywhere cover it to a depth of one and one-half miles! PHYSICAL FACTORS OF THE SEA Water The remarkable compound to which all life is responsible and which gives the sea most of its important characteristics is water. First, water is most common on earth in its liquid state, and it is fortunate that the temperature range of most of the earth lies between water's freezing point and boiling point, that is, between 32° and 212° Fahrenheit (salt water freezes at about 29°). It is not merely happenstance that the temperature limits of life lie between the temperature limits of liquid water. Second, water is one of the few compounds that expands, becoming less dense, when it changes from the liquid to the solid state. This enables ice to float, a fact of the greatest significance, for if ice sank, the depths of the sea, and eventually the whole of polar seas, would freeze solid. Third, water is the most universal of all solvents, able to dissolve large amounts of a wide variety of substances that are absolutely necessary for life. In fact, protoplasm is 50 per cent (flour beetles) to 98 per cent (jellyfish) water. This liquid virtually bathes every fiber of living bodies, carrying food and oxygen to cells and carting off metabolic wastes such as carbon dioxide and urea. Fourth, water has a very high capacity to store heat. The great body of ocean water is the storehouse of the earth's heat. Without the oceans, the dark side of our planet would come close to freezing nightly. Fifth, water is dense and gives support and buovancv to the animals that live in it. This has the important consequence of solving the problem of support for marine animals. Strong legs, arms, and skeletons, necessary for support on land, are lacking in the sea. Instead, the bodies of marine animals are modified for locomotion and for offense or defense, and not usually for support. Plants in the sea have no strong supporting stem or trunk. Because of water's buoyancy. ZONES OF THE SEA-WHERE THEY LIVE 3 many marine animals attain large size. Whales reach measurements that are impossible on land; if they themselves are beached, they are unable to breathe because of the great weight of their own bodies. Dissolved Salts For all of its remarkable qualities, pure water would not afford much of a home for living things. The salts are the fertilizers of the seas, and without them animals and plants could not grow. The seas of the world average about 35 parts per thousand of salt with extremes of 15 parts per thousand at the poles, where melting ice dilutes the waters, and 46.5 parts per thousand in the Red Sea, where there is a high evaporation rate and practically no dilution by rivers. Most of the salts are sodium chloride (table salt), potassium chloride, and magnesium chloride, but these are not the really vital ones. Nitrogen is vital in the building of protein. It is rather odd that animals and plants are not able to utilize atmospheric nitrogen, even though nitrogen composes about 80 per cent of air. That task falls to the incredibly numerous nitrifying bacteria in the soil of land, which manufacture salts called "nitrites" and "nitrates" from the nitrogen of air. These are then used by plants to build protein. Eventually, however, many of these nitrites and nitrates find their way into the sea by way of river systems. Allee, and others (1950) estimate that the Mississippi River alone brings 792,000 tons of nitrate into the Gulf of Mexico each year. This is the principal but not the only way that nitrogen reaches the sea. Some nitrifying bacteria exist in shallow sea waters, but how much they contribute is not known. Once in the sea, nitrites and nitrates are used by plants in the so-called shallow-water "zone of nitrogen utilization." Animals eat the plants, and nitrogen in the form of protein is distributed in the tissues of living things of the sea. When these things die, they sink to the depths. Bacteria cause decay, and nitrogen compounds are released in a deep-water "zone of nitrogen regeneration." Nitrites and nitrates accumulate in the depths until they are swept to the surface by currents in places where upwelling occurs. On the surface this rich fertilizer once again becomes available to plant life. So we see that nitrogen in the sea, is subject to an endless cycle, a cycle intimately connected to that of life and death. The story of that other important fertilizer, phosphorus, is similar to that of nitrogen. Once again, it is brought from the land to the sea by rivers, and again it is used bv plants in a zone of phosphorus utilization. The plants are eaten by animals. These living things sink to the deep sea when they die. Decay releases phosphates in a zone of phosphorus regeneration and upwelling of deep waters brings them to the surface. Immense as are the amounts of nitrogen and phosphorus compounds in the sea, we must not think that they are inexhaustible. Just like any other fertilizers, they can be used up. Every summer, as the sea warms and the plankton turns the sea into a living soup, these fertilizers become progressively scarcer as the abundant life uses them up. In some places the amount of such fertilizers limits the amount of life that can grow in the water. When cold weather returns, much of this life dies and the fertilizer returns to the depths. So the cycle of 4 UNDERWATER GUIDE TO MARINE LIFE nitrogen and phosphorus is one of the seasons as well as of life and death. Calcium is an important substance in the life of the sea but for a different reason than was the case for nitrogen and phosphorus. It is also brought to the sea by fresh water, existing in sea water as calcium sulfate largely, but it does not build tissues. Instead, it builds skeletons. About 1.6 per cent of the salts of the sea are calcium compounds. Many groups of animals, notably corals, sponges, crustaceans, worms, molluscs, foraminifers, and some coralline plants are able to precipitate soluble calcium compounds into relatively insoluble calcium carbonate to build their skeletons. Since this reaction proceeds fastest at about 75° to 80° Fahrenheit, a common temperature range for tropical waters, it is no surprise that coral reefs and the largest shelled animals are found in the tropics. Silicon is similar to calcium. It is a skeleton builder, but it is used by only a few groups, notably radiolarians, diatoms, and sponges. It exists in the sea in the form of silicates. Silicates are the principal component of common glass, and most siliceous skeletons have a decidedly glassy look. Nitrogen, phosphorus, calcium, and silicon are the four biologically most important elements that are present in the sea in the form of dissolved salts. There are other elements of lesser importance, such as iron and magnesium, with similar case histories. Aside from their importance to life, salts have two important effects on water. One is that they depress the freezing point of salt water from 32° Fahrenheit to about 29°. The other is that salts cause water to be more dense. Various salt concentrations are partly responsible for varying water densities. Every year about 660 million tons of earth are washed into the sea, and with this dirt come great quantities of nitrogen, phosphorus, calcium, and silicon. Therefore, it would seem that the sea is constantlv growing more and more salty. Some scientists have assumed that the sea was originallv fresh water and have even tried to compute the age of the sea by dividing the total tonnage of salts in the sea by the yearlv deposition rate, but they have arrived bv this method at very low figures for the age of the sea. The reason for this is that deposition of these salts in the sea occurs at a varying rate, and salts are often deposited permanently on the bottom. As salts are added by rivers, so they are taken out of the water by various means. In fact, the sea has probably remained about as it was with respect to salt content for the last billion years. There is some correlation between river deposits of salts and abundance of life. The Atlantic and Arctic receive most of the North American river drainage and have continental shelves to hold the deposits. Less river water flows into the Pacific, it has no shelf to speak of. The comparative richness of life in the Atlantic is due to this fact. The richness of some Pacific areas is due to another factor, upwelling. Dissolved Gases Oxygen enters the sea primarily by being dissolved at the water's surface and is distributed by water currents. Disturbed surfaces dissolve oxygen best simply because more surface is exposed and because rough water acts as a good mixing agent. ZONES OF THE SEA-WHERE THEY LIVE 5 Plants also furnish oxygen to the sea by the process of photosynthesis. This reaction requires light, however, so it occurs only in approximately the first 250 feet of water. As the light diminishes with depth, plants become oxygen users, not producers. At about 100 feet, plants use about as much oxygen as they release, and below that depth the few plants that exist become dependent upon oxygen, releasing very little. The 100-foot depth is called the "compensa- tion level," and it xaries with light conditions, latitude, time of day, and water claa'ity. Oxygen is extracted from water by the respiration of all forms of life and bv oxidation and decay of dead bodies through the agency of bacteria. Allee, and others (1950) state that bacteria probably use more of the sea's oxygen than all other forms of life combined. The concentration of oxygen in the sea is highest at the surface, where solution and photosynthesis take place and where the water may even become supersaturated. This is particularly true of brightly lit surface waters in mid- afternoon when photosynthesis is at its peak. At a depth of about 1,500 feet, oxygen falls to its lowest level because it is not produced there, yet enough life is present to use what there is rapidly. Oxygen increases below the 1,500-foot level because it is carried downward by sinking cold vvaters from polar seas. In the comparatively few deep spots where overturn of water cannot occur and where sinking cold water cannot reach bottom, the water may become stagnant. All of its oxygen is used, and the poisonous gas hydrogen sulfide accumulates from the decay of dead bodies that sink there. The Black Sea is the most famous body of water that accumulates hydrogen sulfide. Practically no life at all occurs there from a depth of 600 to 6,000 feet. The Bay of Naples and several deep fiords of northwestern Europe also tend toward stagnancy. Carbon dioxide is the third major gas in the sea. It is a fertilizer similar to the salts of nitrogen and phosphorus. It is released into the sea by the respiration of all animals and by plants deprived of light. It is dissolved by surface waters in the same way that oxygen is dissolved from the atmosphere. It is also released at the bottom of the sea by decomposition of dead bodies and, like nitrogen and phosphorus compounds, accumulates in deep waters. Upwelling brings this carbon dioxide to the surface. Carbon dioxide is used in two very important ways. First, it is used by plants in photosynthesis to build carbohydrates (sugars and starches). Second, it is used in the converted form of carbonates and in combination with calcium to build the limy skeletons of many invertebrates. Pressure and Depth Next to the yvetness of water itself, the most noticeable effect of water on the diver is that of increased pressure as he descends. The pressure of air at sea level is one atmosphere or about 15 pounds per square inch. For every 33 feet of descent into the sea, the pressure increases one atmosphere. At the deepest part of the ocean, animals are subject to about 1,000 atmospheres of pressure or 15,000 pounds per square inch. This immense pressure would be quite disastrous if applied unevenly to a body, but it has no effect when it is exerted in all directions inside the body and out. To illustrate: each square inch 6 UNDERWATER GUIDE TO MARINE LIFE of our body is subject to 1 5 pounds of pressure at sea level. Yet we feel nothing. The pressure outside us is balanced by pressure in our tissues. However, if the pressure were exerted unevenly, there would be a startling difference. The area of the palm of a hand is about 10 square inches. Very few of us could move our hands if they were held down by 150 pounds of weight. At a depth of 100 feet this weight would become 600 pounds. But, as many aqua-lung divers know, there is little difference in swimming at 100 feet or near the surface because the pressure is equal in all directions inside the body and out. This is not to say, however, that dives to 100 feet are as simple as those at the surface. Pressure at any depth is due to the weight of water in a column above that depth. As is well known, added weight on a body will tend to compress it, but water is not very compressible. If water were incompressible, the surface of the ocean would rise about 100 feet, but 100 feet in an average depth of two and a half miles is a very small fraction. This comparative incompressibility of water has the effect of keeping buoyancy about the same at all depths, so that incom- pressible objects sink in the sea at an even rate all the way to the bottom. Com- pressible objects, such as fishes with a swim bladder, would tend to sink faster and faster with increased depth since added pressure reduces their volume which, in turn, decreases buoyancy. To combat this, fishes increase the pressure in their swim bladder to keep their body volume constant and to equalize buoyancy at all levels where they swim. The depth to which any fish can safely descend might depend in part on its ability to increase pressure in its swim bladder to match water pressure. Hesse, Allee, and Schmidt (1951) state that surface fishes with empty swim bladders can stand pressures of 100 atmospheres without harm. Plankton is similar to fishes in being able to withstand rather drastic pressure changes. By day, much plankton lives just at the edge of the lighted sea, at a depth of 1,200 to 1,500 feet. By night, this plankton rises to the surface, a change of 30 to 40 atmospheres of pressure, without harm. Though very little is known about pressure relations in the deep sea, one thing seems fairly certain. The pressure itself plays a relatively minor part in determining what forms of life live there. More important are temperature, which is near the freezing point in the deep sea, lack of light, and relative scarcity of oxygen and food. But pressure does have special effects to which animals must adapt. The swim bladder problem is one that deep-sea fishes must solve. Pressure also causes colloid gels (such as protoplasm) to take on more water, and for this reason deep-sea animals are more gelatinous in texture than shallow-water ones. Light Light is important to life in several ways. First and foremost, it supplies the energy necessary for plant photosynthesis. Second, it is necessary for sight, without which the behavior of many forms of life would be radically changed. Third, it is responsible for color itself and for the adaptive coloration of animals and plants. The nature of light and color are vitallv important concepts to grasp if we ZONES OF THE SEA-WHERE THEY LIVE 7 are to understand the way animals and plants live. Light as we see it is actually composed of a mixture of colors: red, orange, yellow, green, blue, indigo, and violet. These spectral colors have been seen by anyone who has seen a rainbow or looked at sunlight refracted through a prism. These colors take the form of waves of varying length. As the length of the waves varies, so does the color of the light vary. The longest visible wave lengths are red and the shortest are \iolet. Light waves that are invisible to our eyes exist, and these are called "infrared" (longer than red) or heat waves, and "ultraviolet" (shorter than violet) or burning rays. Light has two important characteristics that are important to the photographer (Chapter 3) and to the life of the sea. These must be distincdy separated in the mind as two factors which may act either separately or together. They are interisity and quality. Intcnsitv simply means brightness. Any wave length or combination of wave lengths can exist over a wide range of intensities. As progress into the depth of the sea is made, intensity of light decreases. Some animals have photoreceptors that react solelv or chiefly to light intensity. Intensity regulates the behavior of a great number of animals. Some plankton rises to the surface by night. Squirrel fish, conchfish, coral polyps, and big-eyes become active at dusk. Plants cannot photosvnthesize below a certain light intensity. Quality of light, or color, depends chiefly on differences in wave length from red to violet. The color of an object depends on differential absorption of some wave lengths of light and the reflection of others. For example, the squirrel fish appears red because it absorbs all wave lengths of light except red, which it reflects. Quality of light has important consequences for life. The different groups of algae photosynthesize in response to different wave lengths. Adaptive coloration of many animals is a response to light quality. It is now possible to descend from the surface to the abyssal depths of the sea and to see what happens to light as progress is made downward. The most immediate effect of sea water on light is reflection from the surface. In calm waters about 3 to 4 per cent of light is reflected and never enters the sea, in average waters 15 per cent, and in rough waters about 30 per cent. From the surface to depths, the quantity of light diminishes constantly due to the opacity of water. It is possible to divide the sea into three zones on the basis of light intensity alone. 1. Euphotic or Well-Lit Zone. This is the shallow-water zone with a depth range of about zero to 250 feet. It is rich in plankton, especially the photosvnthetic types, and in herbivorous or plant-eating animals. Photo- synthetic plankton is abundant to 150 feet and almost disappears at the deepest part of this zone. The depth of this zone is largely determined by the water's clarity. The Sargasso Sea is probably the clearest sea water found anywhere. Sea water is generally more transparent than fresh water. 2. Dysphotic or Dimly Lit Zone. From 250 feet to the level at which all light disappears, the life of the sea is adapted to dim light. The level at which all light disappears varies from 600 feet to about 5,600 feet depend- 8 UNDERWATER GUIDE TO MARINE LIFE ing on water clarity and the angle of incident light. The extent of this zone is not as deep near the poles as near the equator. Animals of this zone frequently have large eyes in order to be able to see in weak light. Animals with bioluminescence, that is, possession of organs that can emit light, become common. Herbivores, or plant-eating animals, become scarce or absent. In clear water, the intensity of light at 3,000 feet is one three-millionth of its intensity at 3 feet. 3. Aphotic or Lightless Zone. Below the dysphotic zone, no light at all exists. Many, if not most, animals possess luminescent organs, either for food- finding or defense. Eyes may be totally lacking or reduced in size. Herbivores are completely lacking, and only detritus eaters (those who eat debris) or carnivores are left. Quality of light is also affected by water, but there are no zone names to describe this. In clear waters, the long waves or reds are absorbed first and the short waves or violets are absorbed last (Chapter 3). Silty or plankton-filled waters reflect red and yellow and therefore appear greenish. By the use of colored photographic plates exposed for long periods, it has been determined that all wave lengths are still present in clear water at 300 feet, but the reds are exceedingly weak. At 1,500 feet red and green are gone completely. William Beebe, in his dives in the bathysphere, observed that a strangely bright, blue light could be perceived by his eye up to about 1,800 feet. After that he saw no light at all. Blue light not visible to the human eye persists quite a distance below this depth, however. The colors of animals and plants vary in response to these changes in light quality. It is not nearly as difficult to correlate the color changes in plants with depth (Chapter 6) as it is to correlate these changes in animals. For instance, it is well known that many nocturnal animals are red (squirrel fish, big-eye, etc.) and that animals of the dvsphotic zone where red is lacking tend also to be red (especially fishes and crustaceans). At those depths these animals must appear black since there is no red left to reflect. Red animals give way to brown, black, or transparent animals in the aphotic zone. So it would appear that red coloration is an adaptation to dim light or to waters where red wave lengths are lacking and that it acts as protective coloring for these animals of nocturnal habits or dimly lit zones. Many people have wondered why so many animals are red at depths where red supposedly cannot be seen. The assumption is made that animals see as humans do. But many animals that live in waters of low light level have the capacity to see objects that humans cannot see. To these animals, red objects are protectively colored. Temperature Temperature determines to a very important extent where and how animals live. Sea water varies from 27° to 108° Fahrenheit depending on latitude and depth. The temperature of surface water varies the most, but deep water of over 1,000 feet in depth in all latitudes is extremely uniform at about 35° Fahrenheit. Temperature variation is not nearly as great anywhere in the sea ZONES OF THE SEA-WHERE THEY LIVE 9 as it commonly is on land. Rarely does the sea vary more than 70° Fahrenheit at any one place. By contrast, land temperatures at Fairbanks, Alaska, vary from 60° below zero in winter to 105° in summer. If a di\'er were to progress downward through the water, even in warm seas, he would reach, at a depth of 150 to 500 feet, a level where the water suddenly became much colder. This place is called the "thermocline" and marks the strong stratification between warm surface waters and cold deep waters. Temperature has several effects on water and on animals which are purely physical. Cold waters are denser and have a greater viscosity than warm water. Cold waters, therefore, sink below warm waters, and the temperature differences become partly responsible for water movements in the ocean. Increased viscosity of cold water means that planktonic animals sink more slowly in cold waters than in warm. This may be part of the explanation for the huge concentrations of plankton, particularly crustaceans, pteropods, and protozoans, in arctic or subarctic waters or in deeper waters of warmer seas. These planktonic animals are relatively scarce in shallow, warm waters. Features of the Sea It is easy to imagine the sea as a sort of featureless bowl, but since the advent of sonic soundings about fifty years ago, it is now known that the seas show all the features of the land— mountain ranges, canyons, valleys, vol- canoes, etc. There are differences between the features of the sea and those of land, however. The former are on a much grander scale but are barren and comparati\'elv immutable. The gigantic deformities in the suboceanic crust of the earth for the most part lurk in dark, silent, plantless depths, where erosion is almost at a standstill. Geologically speaking, the ocean may be divided into three regions Cfig. i): 1. Continental Shelf. This is a gradually sloping extension of the land under water. It is formed by the deposition of river sediments as they are brought from the land. As the ocean's water level rises and falls, the shelf is alternatelv part of the underwater sea and part of the land. In some places, such as the West Coast of North America, no continental shelf to speak of is present, but the edges of most continents have shelves extending outward from shore as little as. 10 or as much as 800 miles. The edge of the shelf is usually at a depth of 600 feet. The shelf fauna and flora is the most varied in the sea. 2. Continental Slopes. At the edge of the continental shelf the downward pitch of the ocean bottom increases rapidly so that the water gets rapidly deeper. These clifflike slopes and escarpments form intermediate depths of the sea where protozoan oozes cover most of the bottom (Chapter 7). The size of these escarpments so dwarfs anvthing of similar form on earth that it is hard to comprehend them. They may descend from 600 to 18,000 feet, taking 100 miles of downward slope to do so. This repre- sents a maximum size, but in some places the escarpments are even more spectacular, being not as long but much steeper. 10 UNDERWATER GUIDE TO MARINE LIFE 3. The Deep-Sea Bottom. This lies below the great majority of the sea's area. It is here that the most spectacular canyons on earth sink and the greatest mountains on earth rise. OCEANIC PROVINCE OCCANIC TRENCH Fig. I. Veatures and zones of the sea. In the western United States are many well-known mountains, one of which is Pike's Peak. It rises to 9,000 feet from a plateau of 5,000 feet high, a total of 14,000 feet. Mt. St. Elias in Alaska is one of the very largest of mountains on earth, rising from sea level to 18,000 feet, twice the height of Pike's Peak. Mt. Everest rises to over 29,000 feet, but its base is very high, so it actually is not as large in sheer volume as is St. Elias. By contrast, Mauna Kea of Hawaii rises a total of 31,000 feet directly from the sea bottom (almost 14,000 feet from sea level) and is so huge that it forms the greater part of the largest of the Hawaiian Islands. There are many huge mountains under the sea at least as large as Pike's Peak that never even reach the surface. Mountain ranges of the sea are no less spectacular. By far the longest range on earth is undramatically called the "Mid- Atlantic Ridge," which is 10,000 miles long. It begins off the coast of southern Africa and runs unbroken to Iceland, showing a few times above the surface in such islands as the Azores and Ascension. Perhaps the most spectacular and mysterious of the deep-sea features are the trenches, steep canyons usually found close to shore lines and continental shelves, which plummet to depths of up to 35,000 feet (Mariana Trench). The deepest North American trench is the Puerto Rican Trench with a depth of 30,246 feet. Trenches are usually deeply filled with layers of sediment a mile or more in thickness. Without these, the trenches would be even deeper. The bottom of the deep sea is usually covered with a monotonous layer of red clay where a few hardy animals make their homes. Hesse, and others ZONES OF THE SEA- WHERE THEY LIVE H (1951) give the following percentages for the distribution of the depths of the seas over the earth. 15.6 per cent of the sea bottom is 0-5,900 feet deep 19.3 " " " " " " " 5,900-11,800 " " 58.4 " " " " " " " 11,800-17,750 " " 6.5 " " " " " " " 17,750-35,600 " " The Movements of Water This is a complex subject which concerns waves, tides, horizontal water currents such as the Gulf Stream and vertical currents such as upwelling. It is very difficult to clearly separate these factors. All of them cause a mixing of ocean water in complex patterns and are thus vital to the life of the sea, particularly in distributing animals and plants and in fertilizing surface waters from below. Waves constitute the least massive of the various types of water movement. They are surface phenomena, rarely having any effect on waters greater than 300 feet (or a maximum of 600 feet) deep. Winds provide the energv necessary to create a wave. The size of a wave depends mainly on four factors of winds: (1) velocity— the stronger the wind the higher the wave, (2) duration— the longer the wind blows, the higher waves will become, (3) fetch— the longer the distance over water that the wind can blow without obstruction, the larger the waves, and (4) direction— the longer the wind blows in an unchanging direction, the bigger the waves will be. These four factors interplay and vary greatly to produce waves of a few inches high to a probable maximum of 40 feet in height. Barnett (1954) states that it would take a 60-mile gale blowing steadily in one direction for a distance of 900 miles to produce a 40-foot wave, and since such conditions are not often met, such waves are rare. The very high velocity of gusty winds does not produce high waves because it literally blows the waves' tops off. The water itself moves very little in the horizontal direction of the wave. It merely rises and falls in a circular fashion as the wave form passes through it. To prove this, all one has to do is to place a floating object on the water's surface and notice that it bobs up and down in a circular pattern with the waves, but does not move forward with them. When a wave reaches shallow water, the circular pattern of the water movement becomes flattened because of the proximity of the bottom. This causes the forward edge of the wave to become steeper and steeper until it falls over on top of itself and breaks Cfig. 2). The force of a breaking wave can be tremendous. It is enough to carve holes in rocks or to lift large stones and even throw them through the air. It has been calculated that some huge waves strike with a force of up to 3 tons per square foot, ample warning that swimming in large breakers near shore can be very dangerous business. Tides are very different from waves, in that they are of cosmic origin. The gravitational pulls of the sun and the moon provide the energy of the tides. When the sun and moon are both lined up with the earth and with each other, 12 UNDERWATER GUIDE TO MARINE LIFE their pulls are complementary and the highest tides, called "spring tides," occur. This happens during the new and full moons. When the sun and moon form a right angle with the earth as the center, their gravitational pulls partly cancel each other and the lowest or "neap tides" occur. This happens during the first and third quarters of the moon. WIND AND WAVC DIRECTION Fig. 2. The breaking of a wave on a sloping shore. Tides are erratic in occurrence and in height. In general they are lowest at the equator and highest toward the poles. In most places high tides occur twice a day, but in some places they occur only once. The highest tides in the world occur in Nova Scotia's Bay of Fundy, followed closely by several spots such as the Turnagain Arm near Anchorage, Alaska. In both of these places, tides must travel up narrow channels in a short period of time. This causes a tidal-wavelike rush of water called a "bore" to form. It is not known exactly why extremely high tides form in some places and not in others. Pre- sumably it has to do with the configuration of the ocean bottom. If the bottom forms a trough, so that water can oscillate back and forth in it like water in a bathtub, and is the right size, so that the period of this rocking vacillation matches the period of the movements of the sun and the moon, water will rock back and forth in the trough to give very high tides of up to 50 feet. Bores will occur when the trough has a constricted end, up which tidal water may rush. The currents in the sea have a more complex origin and are of greater importance than waves and tides, even though they are usually not as spectacular to look at. Horizontal currents traveling over the ocean's surface are motivated primarily by prevailing winds, and their direction is influenced by the rotation of the earth. The prevailing winds in both hemispheres blow from the east along the equator, the trade- winds, and from the west along the horse latitudes (30° north and south), the westerlies. The rotation of the earth causes a deflection of these winds so that their paths become circular— clockwise above the equator and counterclockwise below. In general, ocean currents follow this wind pattern (/ig. 3), but are compficated by the conformation of coasts and the ocean bottom. In the center of these great, rotating bodies of water are eddies where the water just turns around and around and never goes anywhere. One such eddy is the Sargasso Sea. Vertical currents are caused bv density diff^erences of water, that is, by varying temperature and salinity. As water is heated near the equator, it becomes ZONES OF THE SEA-WHERE THEY LIVE 13 less dense, rises to the surface, and flows northward. At the polar regions, it cools, sinks and flows very slowly back to the equator at the rate of a mile a day. In warm seas, water evaporates rapidly and becomes denser due to in- creased salinity. In ihe Sargasso Sea, for instance, water sinks due to increased salinity. Tropical rains have an eflfect in diluting water and making it less dense. These interplaying factors vary so as to produce quite complex results, but temperature has generally the greatest effect in the oceans. Thus, circulation is mostly toward the poles on the surface and toward the equator in deep water. UpwelHng is a special and very important vertical movement caused mosdy by winds. The principal areas of upwelling in North America are off the coast of Newfoundland and widely along the southwest coasts of the United States and northern Mexico. On the lee coasts of prevailing winds, water is blown away from the shore and must be replaced by upwelling water, floating up Fig. 3. Currents of North American seas. The names of the curreyits are indicated on the map. from the depths. Conversely, windward coasts accumulate surface water which sinks. This is reflected biologically by the abundant coral forests in the Caribbean where warm water accumulates, and the comparatively depauperate coral fauna of western Mexico where cool water wells up from below. However, these upwelling seas compensate for lack of coral by possessing a great abundance of plant and animal plankton, and very rich oceanic fisheries, largely com- posed of schooling fishes, become established as a result. In fact, areas of upwelling show the greatest abundance of life to be found anywhere on earth. The champion of all upwelling areas seems to be off the coast of Peru and northern Chile. There are other causes of upwelling besides winds. These are storms, tides, and the passing of two currents of unequal density (when this occurs, one sinks 14 UNDERWATER GUIDE TO MARINE LIFE and the other rises). But these causes are probably minor in comparison to the effect of winds. The net result of all these massive water movements is the production of rather stable, but exceedingly complex water masses in the seas, which may be identified by characteristics of temperature and salinity. The complexities of their movement are introduced largely by the lay of the land, that is, con- formation of the ocean bottom and of the land masses. For instance, the best known of the horizontally moving water masses, the Gulf Stream, is a distinct, blue river of water, 1,200 times the size of the Mississippi, which pours out of the warm Gulf of Mexico through the Florida Strait at a speed of up to 6 mph. It proceeds north to Cape Hatteras, where it is deflected out to sea, runs into the cold Labrador Current near Nova Scotia, and breaks up into four separate currents before reaching England. Water returns to the Gulf of Mexico by the Caribbean Current to the south. Similarlv, the movements of vertical water masses are influenced by the conformation of the bottom. Most of the cold bottom water of the mid-Atlantic is supplied from the antarctic because those of the arctic are blocked by underwater barriers. Local rain, river influx into the sea, melting ice, and sun also have effects on densitv which greatly complicate the picture. In summary, the movements of water have, no doubt, a greater effect on the abundance and distribution of marine life than any other physical factor. The oscillating surface factors of waves and tides are mainly responsible for mixing oxygen and carbon dioxide at the surface. In shallow water, they buffet the living and nonliving alike, either building or eroding shore lines and forcing the sessile, or sedentary, animals and plants that live under the impact of their blows to become either massive and thick like corals and sea shells, or flexible and resilient like seaweeds and gorgonians. The nonoscillating water movements are composed of an interplay between two rotating systems, one which is horizontal and caused by winds and one which is vertical and caused by winds and varying water densities. These produce circulating water masses and serve to mix dissolved salts and gases and to distribute animals, particularly planktonic ones, throughout the seas of the world. ZOOGEOGRAPHY: Communities of Marine Life It is almost too obvious to point out that the sea is not evervvvhere the same, but simple statements are often the most important ones. The studv of the various environmental differences and similarities that exist in the sea or on the land and the precise recognition of their characteristics allows the recognition of definite zones, that is, fairly sharplv delimited areas where a certain set of physical features prevail— certain temperature limits, water movements, depths, light, etc. One such type of zone is the tidal, characterized by movements of waves and tides. Another type is the eu photic or lighted zone, discussed in the section on light. Another is the temperate, delimited bv certain temperature extremes. Any zone will have living in it a group of animals and plants that have become specialized to meet the physical and biological requirements of its zone in order to live more efficiently there. All of these forms of life are grouped ZONES OF THE SEA-WHERE THEY LIVE 15 tooether as a connmmity. It is usually possible to apply the same names to communities as to zones, and so there are, for example, tidal, temperate, and euphotic communities. Provinces are divisions of the sea similar to zones but larger in scope. The whole of the deep sea, excepting the very bottom, is spoken of as the oceanic province, for instance. The communities and zones of land are separated on the basis of plant associations. In the sea, temperature and the nature of the substrate are used. The zones defined bv hght intensitv have already been described. Temperature zonation is simplest and will be considered first. Temperature Zones The temperature zones are three in number: tropical, temperate, and arctic. (The temperature zones of North America are given in figure 4.) Fig. 4. The tetnperature zones of North America. THE TROPICS The temperature of truly tropical waters almost never falls below 68° Fahrenheit. Near shore, most tropical waters are 75° to 80° all year long. Some subtropical waters fall to a minimum of 61° but are not as different from the tropics as are the temperate zones. The tropical seas represent the most ideal 16 UNDERWATER GUIDE TO MARINE LIFE environment for life found on earth so it is not surprising that the greatest diversity of hfe lives there. The tropics are characterized largely by endemism, that is, a great many groups live there and nowhere else. All phyla of the animal kingdom are represented. High temperature is the basic reason for this, but other factors enter in. For instance, there is an abundant supply of food in the tropics so nourishment is not usually a major problem. Not only are the tropics characterized by endemism, but animal groups seem to be more diverse there than anywhere else. The most bizarre shapes and habits and the greatest variation in behavior within any group are usually present in tropical members of that group. A review of the fishes adequately reveals this fact. The tropics are further distinguished by the presence of the lowest tides and the clearest water of all seas. They are the home of the coral reefs and dazzling white coral sands. Tropical seas are divided into these three regions: the Indo-Pacific, the Adantic, and the East Pacific, in order of their richness of species and size. Many animals and plants such as the hammerhead shark, all the sea turtles, and many bony fishes are found in all three, that is, they are circumtropical. Atlantic Tropics. These waters extend north to southern Florida and thence to Bermuda. The whole of the Gulf of Mexico is included. The warm Caribbean and Gulf currents characterize this region. There is good coral growth here, probably richest in the Bahamas, but these are neither generally as lush nor as extensive as those of the Indo-Pacific. The sub-tropics of the Atlantic extend up the eastern coast of Florida to Cape Hatteras. There is very little coral growth here because of low temperatures and the lack of the hard bottom to which coral must anchor (Ekman, 1953). East Pacific Tropics. Tropical waters reach north to the lower tip of Baja California on the west side of that peninsula, and the whole of the Gulf of Cali- fornia on the east is tropical. The subtropics reach north to San Diego. Upwelling of deep bottom water cools almost the whole of this zone rather drastically at times, and the warm waters are rather shallow in extent. Even in the tropics, water of 54° Fahrenheit lies only 600 feet down. The lower limit of the tropical water is a shallow 300 feet. This is a distinct zone which bears a few resemblances to the Atlantic tropics, but which, because of the isolation forced on it by the barrier of the Isthmus of Panama and the depauperate central Pacific, has many endemic groups and lacks several tropical groups completely. Ekman (1953) lists the kelp basses, Paralahrax, as endemic and says that porgies have only one species, butterfly fishes two species, while parrot fishes, common eels, and manatees are completely lacking. TEMPERATE SEAS These are the waters of the greatest temperature variation. Temperatures do not generally rise above 70° Fahrenheit, but they do sometimes reach as much as 80°. However, these zones, being subject to wide seasonal variation, may have temperatures only a little above freezing to the north in winter. The temperate zones show a wide variety of bottom types from rocky to sandy or muddy. The shores are often covered with dense beds of brown algae, which are scarce in the tropics. ZONES OF THE SEA-WHERE THEY LIVE 17 Atlantic Temperate. Atlantic temperate waters lie from Cape Hatteras north to Cape Cod where the Labrador Current ends its southward flow. At Cape Cod the mean annual temperature is 46" Fahrenheit and the extremes are 37° in winter to 66° in summer. Above Cape Cod is a zone reaching to Labrador that may be called cold temperate, boreal, or subarctic. This is the home of the sculpins, halibuts, rock eels, and of huge schools of cod and herring among others. East Pacific Temperate. The temperate Pacific lies from San Diego to the Gulf of Alaska. This is a very uniform zone, both in temperature and in flora and fauna, throughout its whole length. Ekman (1953) lists the following August temperatures for this zone: San Diego 64°, San Francisco 55° to 61°, Seatde 61°, and the Gulf of Alaska 57°. The reason for this uniformity is that upwelling cools the southern part and warm currents warm the northern part. The winter temperatures show much greater variation, however, being much colder to the north. This region has a verv large per cent of endemics. Ekman (1953) lists the surf perches, greenlings, and rockfishes QSehastodes^, and there are many others. Starfishes are more greatly varied in this region than anywhere else in the world. There are 92 species of them, of which 60 per cent are endemic. The sun star, Pycnopodia, is the largest of all starfishes. The boreal or subarctic Pacific is similar to that of the Adantic with its cod, halibuts, rock eels and sculpins. It extends from the Aleutian Islands north to the Bering Strait. Boreal waters in general contain the greatest schools of fishes in the seas. Colder waters have fewer kinds of animals and plants than warmer waters, but this does not prevent those few species from reaching immense numbers in the summer when light is strong, days are long, and surface waters are still cool and viscous enough to support plankton. These constitute ideal conditions for plankton and when they are combined with the upwelling near the Labrador coast, which brings nourishment in the form of dissolved salts to the surface, it is not too hard to see why life is so abundant there. Boreal seas are further characterized by immense beds of algae, rocky shores, and the world's highest tides. ARCTIC SEAS Waters of the arctic zone never get warmer than 41° to 45° in summer and reach these heights only for short periods. Ice is an ever-present characteristic whether in floe or iceberg form. These waters are present north of the Bering Strait and Labrador. In general, the boreal animals and plants are also found here, and among them are sculpins, blennies, sea snails, and dense beds of brown algae. Tides are high and shores are rocky in this zone. Substrate Zones On land, the word "substrate" usually refers to the ground under our feet, since it forms the support of most terrestrial animals and plants. But in the sea, "substrate" may refer either to the ocean bottom or to the water itself since both are dense enough to support life. (In subsequent chapters, "substrate" is 18 UNDERWATER GUIDE TO MARINE LIFE used in general in the former sense.) Therefore, the substrate of the sea, in which anu over which the life of the sea hves, is three-dimensional in contrast to the two-dimensional substrate of the terrestrial world. Flying animals, such as birds, insects, and bats, like fishes, live in a three-dimensional substrate, air. The ocean is divided into two large provinces, the neritic, over the continental shelf, and the oceanic, over the deep sea. Both of these are subdivided into two subprovinces, the henthic, or sea bottom, and the pelagic, or open water. (The divisions of the provinces of the sea are given in figure 1.) Since "pelagic" simply means "living in the open sea," it usually refers to animals of several types, subdivided according to swimming ability. "Planktonic" (from flanktos, meaning "wandering") refers to animals whose swimming powers are not great enough to combat water movements. These usually small or microscopic organisms, about 60 per cent of which are diatoms, are the most important life of the sea. The word "nektonic" is applied to animals that move freely through the water. The highest development of free movement is found in fishes and the re-entrants. The use of these terms is rather simple. For instance, the tuna is largely oceanic-pelagic-nektonic. A snail or a scorpion fish is neritic- benthic. Some animals overlap in zones. A bottle-nose porpoise is pelagic-nektonic but is both oceanic and neritic. Some groups are exclusively pelagic, such as diatoms, radiolarians, jellyfishes, arrow worms, pteropods, and some prochordates. In order of abundance among animal pelagic forms, Hesse and others (1951) list copepod crustaceans, pteropods, siphonophores, arrow worms, octopuses, and squids, then fishes. Diatoms are more numerous than any of these. Exclusively henthic groups include sponges, sea squirts, brachiopods and bryozoans. Most echinoderms, worms, clams and oysters, and snails are also henthic. THE OCEANIC PROVINCE The water beyond the continental shelf is divided into zones according to light. For practical purposes, it is best to consider two major zones to be present rather than three previously set up in the section on light. Thus, there is a lighted (euphotic) zone and a dark or abyssal (aphotic) zone. The dysphotic or dimly lit zone is a transitional zone between these two. The Lighted Oceanic-Pelagic Zone. This zone is, of course, not well-defined from the lightless zone and varies greatly in depth. In general, it is never more than 600 feet deep. All nektonic-pelagic animals, whether they are oceanic or neritic, must be alert or protectively colored or both because of a complete absence of hiding places. In keeping with this, it is here that are found the swiftest swimmers. Coloration is usually countershaded or obliterative (Chapter 2) and is composed of blues or greens above and white or silvers below. The mackerels and tunas are perfect examples of this. The fishes, squids, and mammals are the dominant nektonic-pelagic animals since their powers of swimming are best. There are no large plants and there is no organic debris (detritus) in this zone, so all of the nektonic species depend upon plankton or plankton-eaters either directly or indirectly for food (Chapter 2). The nektonic species are slightly heavier than sea water and must swim slowly but constantly to keep from sinking. The swim bladder of the fishes ZONES OF THE SEA-WHERE THEY LIVE 19 helps keep them near the surface, but oddly enough some pelagic fishes such as mackerels have no swim bladder. Oil and fat help the mammals rise to the surface, but even they almost never cease moving during the whole of their lives. Plankton, on the other hand, is largely dependent upon currents. Most plank- tonic animals and all planktonic plants are microscopic or small, like crustaceans, pelagic tunicates, various larvae, arrow worms, protozoans, pteropods, and diatoms, but some are very large, such as the giant lion's-mane jellyfish, Cyanea, the basking shark, and the ocean sunfish (Hesse, and others, 1951). These large planktonic animals are good swimmers, but they usually just drift with the currents. All of the small plankton depend heavily on flotation mechanisms to keep from sinking. The specific gravity of sea water is 1.02 to 1.03 and that of living matter is 1.02 to 1.06. The flotation mechanisms that compensate for this slight overweight are several. Most protozoa, many larvae, and many crustaceans have extensions of their body or skeletons so that they off^er great resistance to water. These extensions work best in water of rather high viscosity and are found only on very small animals. Other animals and plants reduce their specific gravity by various means, such as reducing the weight of their skeletons (pteropods, protozoa, and crustaceans), taking on extra body water (jellyfish), storing oil droplets (fish eggs, basking shark, penguins, whales, ocean sunfish), or by storing gases (Portuguese man- of-war, bony fishes with an air bladder, chambered nautilus). The Ahyssal-Pelagic Zone. The animals of this zone depend entirely upon the rain of dead plankton falling from above or on other members of their community for food. There are no herbivores, and there is usually a direct relationship between the abundance of plankton at the surface and the abundance of life in this zone. Fishes and cephalopods predominate, and these are usually small although exceptions to this rule occur. For instance, the giant squid is the largest of cephalopods. Some protozoa are larger at depths, but this is related to increased viscosity of deep cold waters. This is a very stable and uniform zone where changes in light and temperature and the movement of water are very slight. Since there is very little water motion, animals with fragile bodies, such as the oarfish (ribbonfish), are common. Many of the fishes are elongate in shape (the chimaera and the frilled shark). Low temperatures in this zone as well as high pressures prevent the efficient formation of lime (calcium carbonate) so skeletons and shells are weak. Pressure also causes protoplasm to take up more water so tissues are gelatinous. Archaic animals are common because of a comparative lack of competition probably. All over the world, the abyssal zones are rather uniform resulting in the wide distribution of the deep-sea animals. The Abyssal-Benthic Zone. This is perhaps the least well-known zone of the sea where cold, lightless, almost motionless waters exist. In a few places, a rock bottom is found, but most of the bottom is covered with clay or ooze, the latter being composed of the countless skeletons of planktonic animals and plants. In shallower places, the protozoan oozes cover huge portions of ocean bottom (Chapter 7). This is particularly true of the North Atlantic where the protozoan Glohigerina, holds forth. A red clay bottom is typical of much of the Pacific and is a very sterile medium for life. 20 UNDERWATER GUIDE TO MARINE LIFE Support and anchorage are big problems in such gooey substrata. Sea spiders (pycnogcnids) creep about on long legs. Crinoids and glass sponges have long stalks on which they are supported, and some crabs have hairy feet. Brittle stars are abundant and support themselves on their long arms. The largest known crustacean is the Japanese spider crab with legs that may span 1 5 feet, lives here. THE NERITIC PROVINCE This is the realm of the underwater swimmer. Whereas oceanic diving has been done and will be important in the future, the shallow waters near shore offer greater variety of life and less harsh conditions; almost anyone is able to enjoy these waters. The boundaries of this zone are the shore line and the outer limit of the continental shelf (if it is present) at a depth of about 600 feet. Therefore, the majority of the neritic province is euphotic or well lighted. This province is variable in almost every respect, including nature of bottom, temperature, animals and plants, and water movements. The Neritic-Pelagic Zone. There is an overlap between the oceanic-pelagic and the neritic-pelagic. Most of what has been said about the oceanic-pelagic also applies to this zone. However, many of the animals of this zone are dependent on the bottom during some time of their life. Pelagic invertebrate larvae usually settle to transform to sessile, benthic adults, and many of these pelagic animals at least feed on or near the bottom. Actuallv, onlv a few habitually neritic animals and plants are free of the bottom throughout the whole of their lives, and these are mostly planktonic ones, small species such as diatoms or some crustaceans. The life of this zone includes planktonic larvae, jellyfish, fish eggs, crustaceans, pteropods, squids, the great majority of sharks and rays, and most bony fishes. Most of these cannot survive if they stray into the oceanic province. The IN eritic-Benthic or Littoral Zone. Of all the zones of the sea, this one shows the greatest diversity of life, especially in a special subdivision of this zone, the coral reef, which will be considered presently. It is rather difficult to decide how to subdivide this area. From the point of view of water movement, there is said to be an eulittoral or tidal zone, which is bounded by the high- and low-water marks of the highest spring tides and which varies in extent with the geographical location, and a sublittoral or non- tidal zone, which extends from low-water to the edge of the continental shelf. Within each of these are several zones which define the all-important communities of the neritic province. Some nektonic species such as small fishes are so intimately connected to the benthic region that they may be spoken of as being nekto-benthic. Among these, the reef fishes are notable examples. But mostly, the zones of these benthic regions are characterized by algae or sessile, plantlike animals or other invertebrates. THE EULITTORAL OR TIDAL SUBZONE This is the harshest environment of the sea, and the only one that can be thought of as having weather as we know it on land. As the tides cover and ZONES OF THE SEA-WHERE THEY LIVE 21 uncover this zone, great changes in wetness, temperature, salinity, light, and friction produced by the harsh buffeting of waves are experienced. On top of all this, food is rather scarce, being present for most animals only when water covers them. This means that the animals and plants of this zone have to be the hardiest in the sea. It also means that from the low-water mark to the high-water mark, conditions become harsher and harsher and more and more like conditions on land, and that life becomes more and more impoverished. Therefore, it is not surprising to find that many eulittoral animals and plants, such as periwinkle snails, blennies, and gobies, show tendencies toward terrestrialism. This zone is subdivided according to the nature of the bottom into rocky, sandy, and muddy shores. Rocky shores are characteristic of eroding coast lines. This solid substratum forms an anchorage for sessile forms of life such as rock- weed (Fmcms), barnacles, mussels, snails of various sorts. Here life piles on life. As soon as one animal or plant anchors itself to the rock, another may anchor next to it or attach itself to the original resident. Great numbers of small animals live almost exclusively attached to seaweeds or shells. Most of the ani- mals either bore into rock for protection or have heavy shells themselves. The plants are tough and resilient so as to be able to vvithstand the beating of the waves. Tidepools are little rock communities in themselves. A great variety of life that can stand wide ranges in salinity, high temperature, and quiet water lives here, including small fishes such as gobies and killifish, bryozoans, hydroids, seaweeds, tunicates, worms, sponges, anemones, barnacles, and mussels. Sandy and muddy tidal shores are typical of depositing coasts. Marsh grasses such as Spartina are invaders from the land and are not bona fide marine plants. The invertebrate fauna is dominated by burrowing forms such as clams, whelks, sand bugs CJ^ippa), and worms. In places where there are no aquatic plants but where marsh grass is found, a great variety of snails, crustaceans, starfishes, crabs, and others are added. Soft bottoms of a mixture of sand and mud are richer than either sand or mud alone. This illustrates the general principle that mix- tures of habitats usually produce richer fauna than either habitat does when pure. THE SUBLITTORAL OR SUBTIDAL SUBZONE The types of animals and plants of the tidal zone are also usually typical of the nontidal zone, except that the species are richer and more varied here. Once again, there is a division into rocky, sandy, and muddy shores. The rocky shores bear extremely rich faunas which have at their base great forests of plants or plantlike animals, such as kelp or corals. As in the tidal zone, life builds on life, but fishes are dominant. Animals here usually have heavy protective shells (abalone), spines (sea urchin), or protective coloration (octopus). Every phylum of the animal kingdom is represented, with the notable exception of the exclusively pelagic arrow worms. The loose substrata of sand and mud are like those of the tidal zone except that they are richer, containing flatfishes, rays, stargazers, anglers, squids, prochordates such as Amfhioxus, and a greater variety of crustaceans, worms 22 UNDERWATER GUIDE TO MARINE LIFE and molluscs. Furthermore, the sublittoral soft bottoms are frequently covered with lush growths of eelgrass or one of the marine algae which, though rootless, can obtain a foothold in sand or mud. This adds another batch of animals such as sea urchins, starfishes, sea horses, pipefishes, eels, killifishes, and even more worms, molluscs, and crustaceans. The presence of plants markedly increases not only the diversity, but also the abundance of life due to the added oxygen and debris which they bring. Here also, a sand-mud mixture supports the greatest amount of life. As might be expected, coarse sand or gravel supports the least. The forms that live in pure mud are very delicate and usually of fairly deep water, since fine mud particles do not come to rest on the bottom until quiet seas are reached near the outer "mud-line" of the continental shelf. Special Zones Three zones, all of them littoral and tropical, must be given special attention because of their unique characteristics. They are (1) a special type of muddy shore, the mangrove zone, (2) a special type of plant community, the Sargasso Sea, and (3) a special type of rocky zone, the coral reef. THE MANGROVE ZONE World-wide in muddy, estuarine flats of the tropics grow mangrove trees, especially the red mangrove, Rhizofhora mangle, whose branching and support- ing aerial and stiltlike roots form a swampy tangle in which the fauna of the sea meets that of the land. Crabs, tree oysters, snails, small tropical fishes, snappers, tarpon, and other brackish-water species meet raccoons, land snails, land crabs, and even some fresh-water fishes. Actually the beginning of man- groves usually means the end of the shallow sea for in the roots of the plants, debris and coral, sand or mud are trapped and a land-building process is begun. In fact, as the debris gets thicker and drier, the mangroves themselves die out. THE SARGASSO SEA In the great eddy of central Atlantic waters from Berm.uda to the south and west toward the Bahamas, Sargassnm, or sargasso weed, the only floating brown alga, is found in fairly large quantities to form a distinctive community of its own. This weed breaks off from its anchorage on rocks in the Caribbean, especially during hurricanes, and is carried to this sea by ocean currents such as the Gulf Stream. There it lives and grows, but it probably does not repro- duce there, until it dies and disintegrates. The total mass of this algae has been greatly overestimated. Clumps of it do become quite large, according to how much gets tangled up together, but it is not a threat to navigation as is some- times stated. Actually, the area of the Sargasso Sea would be quite sterile of life were it not for this weed, for, though this is the area where the world's clearest waters are found, it is also an area of sinking water where nutrients are rather sparse. Sargasso weed is brown and leafy (Chapter 5) so most of the animals that live in it are brown and bear leaflike appendages as concealing coloration and form. The weed forms a substrate, a sort of pseudobenthos, for a rather restricted ZONES OF THE SEA-WHERE THEY LIVE 23 fauna. A comprehensive list of the fauna given by Hesse, and others (1951), and Ekman (1953) includes a few crustaceans, several hydroids, a few worms, brvozoans, two barnacles, a tunicate, some snails, a sea spider, a pipefish, and the extraordinarv sargassum fish. All of these live directly on the weed most or all of their lives. Sargassum also forms a nesting place for flying fishes. The young of many tropical fish use the weed as a place of protection. THE CORAL REEF Most of us, if given free choice of places to dive, would pick the coral reef for several reasons. It possesses by far the gaudiest, most varied, and most luxurious fauna of any place where life is found on the earth. In its never- ending nooks and crannies, innumerable animals make their homes, animals which quite frequently have extraordinary habits. Coral reefs owe their existence to groups of animals and plants which deposit calcium around themselves as a protective skeleton. This building of lime progresses best in tropical waters for two reasons: (1) the chemical reaction which forms calcium carbonate is fastest at temperatures of 68° Fahrenheit and above, and (2) calcium is more soluble, and hence more abundant, in warm waters than in cold. Though the name "coral reef" is given to a limy reef built largely by stony or madreporite coral, these little colonial animals are not the only reef builders. On almost all reefs the stinging, hydroid millepores, called "fire corals," are very common and, to go farther afield, foraminiferans, calcareous algae such as Halhneda, and calcareous bryozoans play a dominant part. In fact, the madre- porite corals sometimes even take third place behind foraminiferans and algae in the building of some reefs. Still, the tropical madrepores are the dominant and most spectacular builders of reefs. Temperature is important; a minimum of 68° Fahrenheit is necessary for the building of coral reefs. The temperature also determines the abundance and variety of corals, for at the marginal temperature of 68°, coral becomes rather depauperate. It takes higher temperatures of about a minimum of 74° to introduce large branching corals to a reef. Borderline waters of 68° such as those off Bermuda, have a noticeably depauperate coral fauna. Hesse, and others (1951), compare Bermuda's 12 coral species to the Red Sea's 125. On the other hand, coral can stand very high temperatures of up to 132° Fahren- heit when uncovered by low tides for short periods. Stony corals do exist in cool temperate waters, incidentally, but reefs are never formed. Temperature is not the only limiting factor for the growth of coral reefs. Moving water must be present, and it must move at the right speed, not so fast as to break the coral or to prevent the coral larvae from gaining a foothold and not so slow as to prevent the distribution of enough planktonic food and oxygen to keep the coral animals alive. Coral must have firm substrate to which to attach itself. It is poor or lacking if hard bottom is scarce. Such is the case on the otherwise adequate southeastern coast of Florida. Coral is usually poor in areas of cold upwelling waters. The eastern Pacific has a poor coral fauna for that reason. 24 UNDERWATER GUIDE TO MARINE LIFE The depth of water influences coral growth. Because of the symbiotic, photosynthetic, oxygen-supplying flagellate protozoans (Chapter 7) that live in coral polyps, growing reefs are not found in waters deeper than about 150 or perhaps 200 feet. Clarity of water has a great deal to do with successful coral growth. In fact, silting can kill coral by cutting down light or by smothering the coral animals. Lowered salinity will eliminate corals. The mouths of rivers never have coral growths and heavy tropical rains are a reef -destroying factor. With all these things taken into consideration, it is almost a wonder that reefs are formed at all. Nevertheless, reefs are one of the most common char- acteristics of all tropical seas, being common between 30° north latitude and 30° south latitude. The Gulf Stream pushes the northern limit up to Bermuda at 32° north latitude. Probably the finest and most extensive coral formations of the North American continent are those of the Bahama Islands. Coral grows at varying rates depending on the local conditions. A usual average of one inch a year is given by Hesse, and others (1951). The growth of branched coral is sometimes marked by yearly ridges. The American Museum of Natural History in New York has a fine specimen of antler coral from the Bahamas showing these ridges a good two to three inches apart. This is adequately rapid to make nautical charts worth very little after a few years. In places like the Red Sea, whole port towns have had to be abandoned because coral growth made their harbors inaccessible. The heavy stone corals and brain corals are the foundation stones of reefs. The branched Acro-pora shows great variety of form, being more delicate in deep or quiet waters. All of these look like they are of a permanent nature, rugged and durable, but the story of a reef is not one of mere construction. It is also one of destruction, as is the whole cycle of life. Coral polyps die eventually. Single coral colonies may live and grow over a period of a couple of hundred years, but this process comes to a halt, and if the dead coral is not quickly built upon by new colonizers, its lime will be etched by the waves and turned to sand or redissolved in the sea. Waves are the greatest of the physical factors which destroy reefs. Two more of these are winds and rains. But probably the greatest overall destroyers are boring animals such as worms, sponges, clams, crustaceans and others. Fishes which gnaw on coral, such as the common parrot fishes, also add their bit in destroying coral formations. All of these forces help reduce coral to sand, which sometimes becomes compacted into coral limestone such as that seen on shores near many coral seas. The best reefs to visit are those in which the forces of construction overbalance those of destruction. The practiced eye soon learns to recognize the difference between dying and flourishing reefs, which are sometimes close together. Three types of reefs are known (/ig. 5). These are (1) the fringing reefs which closely follow coasts near shore, (2) the barrier reef which is separated from shore by a lagoon, and (3) the coral atoll which is a ringlike island formed by the sinking of a volcanic island at sea. The longest construction ever built by living animals, including man, is Australia's Great Barrier Reef, 1,250 miles of coral formations. The deepest constructions of life are also of coral origin. At Eniwetok Atoll in the Pacific, a boring of 4,000 feet was made through limestone before striking a volcanic rock bottom (Barnett, 1955). ZONES OF THE SEA-WHERE THEY LIVE 25 As has been pointed out, the coral reef is the most gaudy and varied realm of life on earth. The animals live so close to each other spatially that they often develop symbiotic dependencies on one another. The pearlfish and the sea cucumber, the conchfish and the conch, and the zooxanthellae and the coral polyps are only a few examples of this. Coral itself "blooms" at night, the polyps mostly keeping to their limestone chambers by day. In fact, the whole reef teems with nocturnal animals, such as big-eyes, squirrel fish, and others that are more active at night than by day. But even by day the coral reef is a bustling community. So ideal is the protection offered among coral branches and caves that many animals never leave the reefs. This protection also seems to allow them to be more brilliantly colored than elsewhere, but it must be admitted that the reason for the exceptionally gaudy colors of reef animals is not known. Possibly, the colors of the animals have evolved to match the brightness of the lisht and the clearness of the waters. Com- o O petition for space on the reef is keen and is probably another reason for the diversity of life, since the animals attempt to adapt to new situations quickly (Chapter 5). Most of the life of the reef is benthic or nekto-benthic since almost all of the species, including the large groupers, make their homes in reefs or feed near the bottom. The great majority of benthic groups are represented on reefs as are the great majority of animal phyla. About the only group which is not well represented on reefs is the large algae, chicflv the brown algae (phylum Phaeophyta). Reefs are largely an animal realm and, indeed, very little matter of nonanimal origin is found there. FRINGING REEF BARRIER REEF ATOLL Fig. 5. The three kinds of reefs Qadapted from Darwin^. The top row of diagrams represents cross-sectional views of reefs; the bottom row, views from, above. Each type of reef may he exposed at low tide and may even form, the foundation for land formation. Since a reef grows fastest at its outer, seaward border, land masses which are sinking will draw the coral downward. The coral will grow to reach the surface only at its seaward edge and a lagoon will be formed. CHAPTER ^ BIOLOGY OF THE SEA— How They Live In this chapter the ways of life of animals of the zones of the sea will be examined under the broad headings of adaptation to habitat, distribution and dispersal, food and food chains, symbiosis, adaptive coloration, animal behavior, and ecology and evolution. Each of these is a huge and complex topic so we can do little more than whet the reader's appetite with general principles here. Furthermore, these few topics by no means exhaust the subject of "how they live." Methods of locomotion, methods of defense, and breeding patterns are three more aspects, for instance (Chapters 7 to 10). It is hoped that the diver will be able to see each animal he encounters under water in the light of the prin- ciples that are given so that he will be better able to appreciate the evolution and adaptation of the system of animate nature and also so that he may be able to increase the depth of his experience in the sea. Most of the principles that are discussed, incidentally, are not peculiar to the sea, but apply to all life, including mankind. ECOLOGY AND EVOLUTION The nature of the sea and its immutability compared to land— its relative homogeneity and lack of sharp environmental contrasts— has been discussed. In this chapter, we shall see how animals adapt to meet this environment. Several billion years ago, probably about 4.5 billion, the earth was formed as an immensely hot conglomeration of condensed gases. Over the next few billion years, this mass cooled to the .temperature of water in its liquid state; below 212° Fahrenheit. By this time, the earth had become enclosed in a hard, wrinkled crust several miles thick, and this crust had trapped water vapor within it which began to be disgorged from volcanoes in great jets of steam. Huge clouds were formed, and a deluge of rain began to fall, which did not let up until the amount of water equivalent to about one-fifth of all the water in the sea had filled the ocean basins. Cloud formation and rains continued with diminish- ing intensity until, about a billion years ago, the waters in the seas had reached approximately their present level. It was in the shallow, warm waters of this sea that life arose by a chemical reaction involving combination of chemicals 26 BIOLOGY OF THE SEA-HOW THEY LIVE 27 drawn from the sea's organic "soup." Before the beginnings of the Cambrian Period, about a half a bilhon years ago, all the major animal phyla had evolved, the history of which still remains largely a matter of mystery. All of these phvla carry remnants of their place of origin in the sea because there is a liquid (plasma) bathing the tissues of all animals which is very much like sea water in its composition. The similarity of sea water and plasma indicates that the seas of a billion years ago, when life arose, were very similar in chemical composition to what they are today. Only one-fifth of all the total number of species of animals (including insects) are found in the sea, whereas four-fifths live on land. This is in spite of the fact that the sea's area is over twice that of land. It is also in spite of the fact that of the sixty classes of animals, only four are not found in the sea, whereas twenty-five are exclusively marine (Hesse, and others, 1951). Therefore the sea has a greater diversity of groups since most groups originated there, but it has a much smaller number of species than the land. This apparent paradox can be explained by the fact that land, where barriers between habitats are much more difficult to cross, which results in isolation, is a much harsher and more variable environment than the sea. Only the most adaptable animals are able to live on land at all, and these are subject to large and constant en\'ironmental changes, with which they must keep step by evolving to meet new conditions quickly. It is isolation, enforced bv barriers, and environmental xariabilitv that lead to increased numbers of species. Land animals have evolved to a higher psychological order in response to a relatively complex and harsh environment. Competition is another factor that affects the rate of evolution. Evolution is fastest in areas of most strenuous competition on land or sea. The greatest competition in the sea occurs in the tropical reef areas, where a great number of species are jammed close together in an optimum environment subject to little variation. Consequently the most advanced and highly evolved species of marine animals are generally found about tropical reefs. Toward the deep sea or toward colder waters, environments become harsher, resulting in elimina- tion of species that cannot adapt to these conditions. Thus, fewer species are found in deep and cold waters than in the tropics, and these are subject to lessened competition. This permits some archaic animals to survive in deep or cold waters, whereas they could not survive in tropical, shallow waters. This is the reason that primitive fish, such as isopondyls (herringlike fishes) are mainly found in temperate or arctic zones, in the deep sea, or in fresh waters. The cold and deep waters are harsh enough to reduce the number of species and to reduce competition but, unlike land environments, are not variable enough and have few barriers to impose isolation. The numbers of individuals is great in northern and deep waters, but the number of species is small. Evolution is not over. Environments constantly change, and all life, including man, must adapt to meet change, or else perish. ADAPTATION TO HABITAT Adaptation refers to adjustment between a living animal and the parts of its environment. This is a phenomenon that is exhibited by every living thing 28 UNDERWATER GUIDE TO MARINE LIFE including man and is a direct result of evolution (Chapter 5). Adaptation is exceedingly complex and can never be measured by a single factor such as adaptive coloration, food-getting mechanisms, or breeding habits for the simple reason that adaptation is an adjustment of all the activities of an animal or plant. Because adaptation is so complex, it is never perfect. It is inconceivable that any living thing could perfecdy adapt in every way to meet the ever- changing requirements of the environment. This is not to say that adaptation is not very successful, however. The adaptations for speed of movement are not absolutely perfect in the mackerels, tunas, and spearfishes, but these fishes are certainly not inadequate in meeting the demands of their predatory ways. Adaptation cannot be studied without reference to the question. Adaptation for what? It is not specific enough to say that an animal is adapted for its general environment. We must define what part of its environment, what specific way of life, that animal has evolved to meet. In other words, where in the community does that animal fit in? We know, for instance, that barbers, bakers, and candlestick makers all have their particular adaptive skills in a human community. It is no less obvious that groupers, parrot fishes, and sponges have their separate places in a coral community as well. The adaptive place of every animal is called its niche, and in order to define it, the community must be re-examined. The various zones of the sea have been defined under the section on zoo- geography in the last chapter. These zones were shown to be different in their general physical characteristics of temperature, substrate, salinity, etc. In other words, zones are defined by a set of physical characteristics. All the animals of a zone were lumped together in a community, each species of which is subject to similar physical characteristics. Different communities may difi^er in many ways, but all communities are alike in one respect: Elton (1935) says that the basic ground plan of all communities is dictated by food chains and that the niche any animal occupies is largely determined by its place in the food chain, that is, what it eats and what eats it. For instance, the herring is a plankton-eater and is eaten by a large number of predatory fishes, such as tunas, swordfishes, etc. Therefore, it is best to define "niche," as Elton points out, by an animal's relation both to its food and to its enemies. "Niche" is an important concept and the discovery of its niche is vital in understanding any animal. But, of course, the description of a niche cannot tell us all there is to know about an animal. The ways it breeds, moves, is colored, and behaves are all vital aspects that do not derive from a study of food patterns alone. One point that must be emphasized is that animals are not continually engrossed in one long struggle for existence throughout the whole of their lives. As Elton (1935) emphasizes, animals are not always doing something. It is true that all animals are adapted to a particular mode of life which revolves around feeding, but it is not true that everything animals do or that every anatomical characteristic they possess are of adaptive significance. Some actions or characteristics may have little or no significance. If the diver is to interpret adaptation correctly, he must learn to distinguish between significant and insig- nificant traits and actions. BIOLOGY OF THE SEA-HOW THEY LIVE 29 Youth, Maturity, Old Age, and Death "The interaction out of which the organism develops ... is between the organism and environment! And the organism is different at each different stage of its development." (Lehrman, 1953.) It is rather obvious to us the way environments change through the years or with the seasons. It is also obvious that animals grow, but it is not very widely appreciated what a drastic effect on the life of an animal growth has. Animals start out life as a helpless egg, which soon hatches or develops into only a slightly less helpless hatchling or newborn. This tiny creature goes through a growth period in which increase in size is rather rapid through juvenile and subadult stages. At the onset of sexual maturity, the growth rate levels off, that is, becomes slower and, though growth continues past the onset of maturity, it tends to be slower and slower as time goes on or even to stop. As old age is reached, vigor is decreased, breeding stops, and some animals (man, for instance) even shrink a little in size. Rarely, in nature, do animals die by "natural causes." Predators, parasites, or disease usually spell the end. There are several fascinating questions that may be raised concerning this cycle of life which every living thing goes through. Some forms of life such as fishes seem never to stop growing. In most animals, aging does not begin until growth ceases and no one seems to know how long fishes could live if it were not for predators, parasites, and disease. Some fishes live one hundred vears or more, but it is not known when they begin to grow old. But these questions are not the main interest here. What is of the greatest significance is that this cycle means that as an animal grows and matures, it must pass through a series of stages, each one of which is different from the others. In general, animals are most vulnerable in the beginning of the cycle when they are small, and therefore the process of elimination of the individuals with the greatest negative selective value is strongest in the young before maturity is reached, a fact of evolutionary significance. Those individuals that do survive do so only because they have been able to find food and elude enemies through- out all stages of their life, and in this rather simple statement, we believe, lies the greatest proof of the great delicacy and complexity of adaptation. For instance, the jewfish is a huge, powerful, carnivorous animal when it is mature. But each jewfish starts life as a tiny egg which hatches into a small hatchling. When the voung fish starts to feed, it is a fraction of an inch long and must have all the adaptive behavior necessary to allow it to live the life of a tiny, secretive, invertebrate-eating carnivore. Soon it grows to a deep-bodied, little reef fish with many of the habits of other small, spiny-rayed fishes that live alert, watchful lives in rocks and reefs. As it grows, its prey and enemies and ways of life continually change, and, at maturity, breeding behavior adds another facet to its life. Therefore, every animal may be thought of as passing through a series of very different stages each one of which must be finely adapted to the environment. The greater the animal's difference in size between youth and maturity, the greater and more significant will be its adaptive changes through life. This is the reason why so many fishes have young that are different in so many respects from their parents and are sometimes mistakenly listed as separate species. 30 UNDERWATER GUIDE TO MARINE LIFE DISTRIBUTION AND DISPERSAL In any community, a number of species is found, each of which has its own distribution Cfig. 6). Some species might be found in only one community— the parrot fish in the coral reef— and some might be found over a wide range of habitats— the common eel, ranging from the Sargasso Sea to fresh-water streams at different stages of its life. U.S. COAST H SPECICS OR 1.3% TROPICAL, SUB-TROPICAL AND TfMPERATE AMERICA 5 SPf CIES OR 0.9 % WCST INDIES AND TROPICAL PACIFIC 8 SPECieS OR 2.5% TROPICAL AND SUB-TROPICAL AMERICA-2IO SPECIES OR 6b% PELAGIC -v3 9 Spec IE S OR (2.3% WORLD WIDE NEAR SHORE IH SPECIES OR 'i.'i% WEST INDIES , EAST ATLANTIC AMD EUROPE 8 SPECIES OR 2.5% SARGASSUM-4SPeCIES OR l-SX Fig. 6. An example of geographical distrihtition— species of Bermuda shore fishes (^adapted from Beehe and Tee-Van, 1933^. The distributions of the species of a community are far from static, and the numbers of individuals of any one species in a community are subject to con- stant variation. Some communities are even compound, that is, there may be several animal communities present in any one zone. For instance, the coral reef has strikingly different day and night communities, and the temperate zones have very different seasonal communities. It even sometimes happens that the fluctuation in numbers of one form of life basic to a community mav change the whole complexion of the environment. This is what has happened in con- nection with the decrease in eelgrass in the 1940's. Fluctuations like these have a great deal to do with factors aff^ecting distribu- tion and dispersal— the questions of why animals are found where they are and how they get there. Such fluctuations complicate the studv of communities greatly, because one is never sure just which species are community members and which are not. The answer to why animals are found in one place and not in another, lies in understanding the principle of ecological valence. Briefly, valence describes a range of tolerance which an animal can stand in relation to any factor of the environment. The valence of coral reef formation with respect to temperature ranges from a low of 68° Fahrenheit to well over 100°. The valence of whales in relation to food depends upon the supply of the planktonic crustaceans on which they feed. Both of these valences are small and limiting in value, but valences can be large. For example, the valence of killifishes with relation to salinity ranges all the way from pure fresh water to pure sea water. The valence BIOLOGY OF THE SEA-HOW THEY LIVE 31 of the sperm whale with relation to temperature allows this species to live in all oceans. Every species of animal has a valence for each of the many factors that de- termine its distribution. There are valence values for food, temperature, salinity, water clarity, light, pressure, enemies, breeding behavior, shelter, other animals of the communitv, and all the other myriad factors that affect that animal at any stage of its life. The distribution of any animal is usually not limited by all of these factors but by only one or two. Figure 7 illustrates the law of the miniviwn; an animal's distribution is limited by the factors to which the animal shows the least tolerance. There are several clues to the discovery of limits to distribution. First, limiting factors are often biotic, that is, have to do with the animals' relation to other animals in the community, and these are sometimes very complex. Second, limiting factors work on all stages of the life of an animal and animals are limited at the stage where they show the greatest sensitivity to environment. For instance, oysters grow in some brackish waters where they are seeded by oyster farmers, but low salinity prevents their breeding there. Third, sometimes the limiting factor is not deadly or even harmful in itself, but affects distribution I I J L ^ LIGHT OXYGEN DEPTH I -|— I TEMPER/^TVBE H SALINITY j I WATETR CLARITY I I -) 1 1 FOOD I I LOWER LIMIT I THE I UPPER LIMIT OF TOLERANCE I f^lNinUM' ^'^ TOLERANCS I I Fig. 7. Ecological valence and the law of the minimum. Each har represents a valence value for one of the important factors in the life of an animal. Distribution in this hypothetical case is limited hy low temperature tolerance and hy the high end of salinity tolerance; this animal coidd not live in waters lower than a certain tem- perature or of above a certain salinity. An animal's distribution is limited hy the factors to which the animal shows the least tolerance. only indirectly. For instance, light in itself is not harmful to most nocturnal animals, but these animals avoid light for protection against other animals (Elton, 1935). Fourth, limits may often be discovered by noticing a decrease in the numbers of animals near limiting conditions. Animals are most common in optimum conditions or away from limiting conditions. Again, coral is a good example. There are few corals in Bermuda where the lower limiting temperature is reached and many in the Red Sea's optimum habitat. Fifth, rarely does any one species of animal approach its physiological limits. Animals find suitable habitats psychologically and travel the path of least resistance away from mar- ginal conditions, unless they are forced there. 32 UNDERWATER GUIDE TO MARINE LIFE The limits of distribution are complicated greatly by the factors of vagility, or the relative power of dispersal of an animal. Sea turdes swim actively and so are found in all tropical seas— they have high vagility. However, many of the groups common to the coral reef communities of the Indo-Pacific are rare or lacking in the east Pacific because their powers of vagility are not great enough to allow them to cross the huge and rather barren central Pacific. Many sessile animals, such as coral and sponges, have pelagic larvae which increase the vagility of the species. Mackerels migrate and have high vagility. Many sedentary fishes or fishes of coral reefs have pelagic eggs or young which serve to spread them to new habitats. In fact, it is relatively common to find young reef fishes far from their parental homes, but rare to find adults of these species far from the tropics. Many Caribbean reef fishes have their distributions listed "north to Cape Cod." This northern extention of range is due mainly to wandering young in the Gulf Stream. Of course, pelagic eggs and wandering young represent wasteful methods of dispersal because many never find suitable habitat and die. For instance, many Caribbean fishes and corals are not found in Bermuda because the wandering young do not live long enough to cross the wide expanse of open sea between these two places. Other methods of dispersal are adventitious (incidentally acquired), for example, sessile animals on ships and driftwood or whale barnacles attached to whales. In summary, the greater the vagility of a species, especially when vagilitv is bv swimming free of the influence of currents (nektonic animals) the more widespread a species is liable to be. An important factor influencing distribution and dispersal concerns barriers. Barriers are like fences which keep animals from moving out of an area. Bar- riers are rather obvious on land and consist of rivers, mountains, plains, or deserts that a species can not cross. But barriers in the sea are harder to see and much more variable. Barriers in the sea are also usually of a less severe nature than on land so that most marine animals have wider distributions than are common for terrestrial animals. The most obvious barrier is a land mass. For instance, the isthmus of Panama prevents most Caribbean species from reaching the east Pacific. Land masses separate the major divisions of the sca- the oceans. Other barriers are salinity, temperature, and oceanic currents. How severe a barrier is depends a great deal on the vagility and tolerance of the animal. For instance, powerful swimmers like tuna can skirt around such bar- riers as land masses, a feat a blenny could not accomplish. Therefore, tunas have wide ranges and blennies small ones. Barriers seem to the human eye to be of a rather permanent nature, but this is not true. Barriers vary widely in geologic time. The earth's temperature is an example. Not too many million years ago Panama lay beneath the ocean's surface. Because of this impermanence of barriers, distribution is influenced by a third factor besides valence and vagility. This is the geologic age of a group. A family of fishes that has existed for 75 million years has obviouslv had more time to spread itself throughout the seas than a family that has existed for 10 million years. This is why the squirrel fishes, derived from relativelv ancient stock, are found in all tropical seas, whereas the grunts of the genus Haemnlon, relative Johnnies-come-latclv, arc found onlv in the New World. Two more factors concerning distribution must be understood. First, when the range for the green turtle is given as "cosmopolitan in tropical seas," this BIOLOGY OF THE SEA-HOW THEY LIVE 33 docs not mean literally that the green turtle is to be found all over the tropics. What it does mean is that this turtle will be found only in the particular habitat of the tropics to which it is adapted, namely shallow beds of marine plants near rocks and coral. Second, the number of individuals of a species to be found in suitable habitats varies greatly with food preference, size, enemies, disease, reproductive capacity, and a number of other factors. Basically, however, density comes down to carrying capacity, a measure of the limit of the numbers of individuals the en\'ironment can support, which is dependent mainly on food supply. There are certain psychological checks that prevent carrying capacity from being exceeded. Territoriality is the chief of these. This means that a breeding pair of a species sets up and defends a certain area where other members of the species are not tolerated. If breeding space is all taken, individuals that do not find adequate territory do not breed. In species that do not set up territories, carrving capacity is limited chiefly by food. Carrving capacitv is not static. For instance, in summer in north boreal seas, long davlight hours and cool viscous waters support huge amounts of plankton so that the carrying capacity for whales and fishes greatly increases at that time. Therefore, the number of a species found in any area depends upon the carrving capacitv for the species in that area at a particular time and is regulated largely by food and the psychological factors. FOOD AND FOOD CHAINS Since the adaptive niche of animals is largely defined by their place in the food chain, it is important that this subject be understood if the life of the sea is to be appreciated. Elton (1935) is mainly responsible for focusing attention on this subject. There would be no food in the sea and no life on earth were it not for chlorophvll and the reaction of photosynthesis. Chlorophyll is present in the great majoritv of plants and in some plantlike animals such as flagellates, and is a catalyst in utilizing the sun's light energy in the following reaction: Light energy + carbon dioxide + water ^ ^ sugars and starches + oxygen Basically, the food chain involves the following relationship which is common to all communities (with the possible exception of the deep sea, to be discussed later in this section) : r ^1 photosynthesis i eaten by i ^ ^- • i l i,- energy of the sun 1 > plants ^ plant-eatmg animals or rierbivores I eaten by flesh-eating animals or carnivores This is an oversimplified diagram, of course. For instance, the omnivores, which eat both plants and flesh, are not included. Neither are the detritus- or debris- eaters which eat dead organic matter from the bottom. Furthermore, there are many grades of carnivores, from the plankton feeders (whales, mackerels) to 34 UNDERWATER GUIDE TO MARINE LIFE the large species which eat large carnivores themselves (killer whale, white shark). Rarely does an animal have a diet that is restricted to only one food species. The dependency of the sperm whale on giant squid, the blue whales on krill, or the sea slug on seaweed are three of the few examples of such dependency. It is much more common to find that an animal will eat a variety of things depending on its food-getting mechanisms and the availability of food species. Probably the animals with the most catholic tastes are the omnivores which eat just about anything they can catch and swallow (some sharks, many crabs, catfishes, etc.). The importance of size on choice of food is easily illustrated. For instance, the killer whale and the blenny are probably equally voracious feeders, but they are forced to eat very dissimilar things because of a vast difference in size. There are exceptions to the size rule that the biggest animals eat the biggest objects. Puffers and killer whales gang up on prey to increase their effective size. Poison, as a food-getting mechanism, increases effective size. On the other hand, the largest of all cartilaginous fishes and mammals (basking sharks, whale sharks, manta rays, and baleen whales) eat tiny planktonic animals. One primitive mechanism of food getting is filter-feeding and is extremely widespread in the animals of the sea. It alwavs involves the straining of plankton- bearing water. The larger plankton-eaters feed by a filtering process. Such fishes as whale sharks, mackerels, and herrings, filter with gill rakers, and whales filter with baleen. Many groups of invertebrates also filter-feed. Among these are clams, oysters, sea squirts, sponges, crustaceans, corals, many worms, and salpas. Some of these, such as crustaceans and salpas, are planktonic and form a vital link between small, single-celled planktonic plants or protozoans and the large fish or mammal plankton-feeders. The other groups are sedentary and depend on small plankton brought to them by water currents. A food chain may be drawn for any animal in the sea. Examples of food chains follow. (The arrow points from the food or energy source to recipient.) Oceanic, plankton-eating, carnivore food chain (whale): sun ^planktonic plant >- krill ^baleen whale Coral reef, omnivore food chain (angelfish) : sun galeae ^anoclfish planktonic pla Small invertebrates — ^ angcU lant / Inshore, carnivore food chain (striped bass): sun ^.planktonic plant ^planktonic animal striped bass -^ small fishes^ BIOLOGY OF THE SEA-HOW THEY LIVE 35 Each of these examples is somewhat simpHfied. The striped bass, for instance, does not eat only small fishes, but may also eat crabs, but in spite of this, food chains are short. The shortest consists of three steps and concerns herbivores as follows: sun >• algae or turtlegrass >■ green turtle The longest are of the large, voracious carnivores, but these are lengthened only because of a step-by-step increase in the size of the carnivores involved: sun ^planktonic plant ►•planktonic animal Killer whale -^ seal -< herring It is often stated that parasitism and mud-eating are examples of simplified food chains. This is not really so. For instance, the mullet is a mud-eater, but its food chain is not the following: mud ^mullet Instead this food chain is quite complex as follows: sun >planktonic plant ^planktonic animal plants (/ ^ plankton-eaters debris and mud planktonic plant 1 X small mud-dwellers illet Therefore, the mullet's diet of mud consists of three separate food chains, all of at least four steps. Similarly, the parasite's food chain is not simple but is exactly like that of a carnivore in that it involves flesh-eating. One food chain deserves to be somewhat set apart from the rest. This is the food chain of the deep sea. It is a popular opinion that life decreases from the surface to the bottom, but several scientists, including Beebe, Barton, and Jacques Cousteau (Cousteau, 1954), have remarked that life decreases to the dimly lit or dysphotic zone and then increases downward. Deep plankton forms the DSL (deep scattering layer) which can be detected on echo-sounding devices. Direct observations made from the bathyscaphe, a remarkable undersea boat which uses the principles of an aerial balloon, indicated an increased abundance of life with depth and that the DSL is the top layer of this abundance. This has led Cousteau and others to suspect that some unknown link in the food chain is responsible. 36 UNDERWATER GUIDE TO MARINE LIFE While it may be true that httle is known of the deep sea, we do not beheve that an unknown hnk in the food chain is necessary to explain the abundance of life in the depths. The sun is still the source of energy, although indirectly, but two food chains, which are linked together, are really involved, one at the surface and one in the depths (fxg. 8). Life in the depths is connected to the fundamental reaction of photosynthesis at the surface and is not self-sustaining, but the basic form of life in the depths is not photosynthetic; it is bacteria which cause decay and nitrification. Bacteria is a sort of "plankton of the depths" with respect to food chains there, and since bacteria is most abundant at the bottom, so are the animals that feed on bacteria. SUN X ^ PLANT PLANKTON ^HERBIVORES ^CARNIVORES TO PLANT PLANKTON DrcAY ^ORGANIC BACTERIA MATTER \ AMMONIA ^-NITRIFYING BACTERIA- -NITRITES AND NITRATES Fig. 8. Deef sea food chains. Animals of the deef scattering layer (DSIS) and deep sea are dependent wpon two sources for their hasic food: I) the constant rain of dead animals from the surface and 2) the deep water plankton consisting of various bacteria and simple plants. Both of these sources are dependent upon the sun, directly or indirectly, for energy. Since surface life is most abundant in the first 250 feet of water and bottom life is most abundant in the deep sea, a graphic representation of the numbers of animals at various depths takes the form of an hourglass as shown on the left. So far, we have dealt only with food chains involving a few animals. When a whole community is considered, the food chains of all the species must be added up to form a food cycle. Figure 9 shows a greatly simplified cycle of the coral reef. Food chains have a great effect on animal numbers. Carnivores, for instance, could never become more numerous than the herbivores they eat. It is estimated that for every planktonic animal there are seven planktonic plants. The numbers of planktonic animals per baleen whale is almost inconceivable. Figure 10 shows a pyramid of numbers in which the most numerous forms of life are shown to be the photosynthetic plankton and the least numerous are the large carnivores. This concept explains the relative rarity of the extremely large, voracious predators like the killer whale and the white shark. BIOLOGY OF THE SEA-HOW THEY LIVE 37 THIRD LtVEl -"' (secondary consumfrs)--^ SECOND LEVEL / ANIMAL PLANKTON (PRIMARY CONSUriERS) / (e ,. protozoa, pferopod. and c«-t*^tttccans) SWIMMING PREDATORS bony (i%h, s«a twr-H^s, 3*A snak« SUN DISSOLVED WUTRIENTS { *o.l1-s and qosea ) SUN Fig. 9. A simplified food cycle of the coral reef arranged in order of energy Qfood') source levels. This is a representation of the total of all the food chains in the com- munity using animals from the coral reef as examples. The first level is photosyn- thetic, the second level herbivorous, and the third level carnivorous. However, any one animal may helong to more than one level— the omnivore which eats both animal and plant matter. Bacteria may he said to constitute a fourth level since they live on decaying matter. In general, the higher the level the less is the dependence upon the level immediately preceding for food and the more efficient the iise of the food supply. Conversely, the higher the level the greater the energy loss through respiration, a fact related to the increased activity in the animals of the higher over those of the lower levels. (^Adapted from Allee and others, 19S0.') LARGE PREDATORS MEDIUM-SIZrO PRfOATORS SMALL PREOATORS I KILLER WHALE i WH\1 I I SEALS » BLUEFISHJI \TE SHARK ILARCC BASS; JACK HERRING ) WRASSE J GOBICS ANIMAL PLANKTON PLANT PLANKTON Vig. 10. The pyramid of numbers; the higher the energy level Qfg. 9) or the farther out on the food chain an animal is, the less numerous it is. The rectangular areas represent relative numbers of animals. Examples are given opposite each rectangle. SYMBIOSIS The word "symbiosis" is derived from syn, meaning "together" and hios, meaning "hfe" and means nothing more or less than "hving together." An animal that lives with another is called a "symbiont." There are various ways, however, in which animals and plants do this, ranging from nothing more than 38 UNDERWATER GUIDE TO MARINE LIFE an occasional meeting in a community to the exceedingly delicate adjustment required by parasitism or mutualism. From the purely chance meetings of species to the most delicately adjusted symbiotic relationship, all shades of gray exist. Therefore, in order to identify the three main categories of symbiosis— commensalism, mutualism, and parasitism —the diver will often find himself in the position of having to make value judgments as to just what the advantages or disadvantages of a relationship might be. The evolution of symbiosis is bound to progress from the disjunctive (no constant bodily contact) to the conjunctive (constant bodily contact), but from there the paths of evolution diverge. Perhaps the most ideal relationship is one in which both symbionts are benefited (mutualism). Commensalism evolves into parasitism, and parasitism often evolves into mutualism. Commensalism This is a relationship in which the symbionts share the benefits unequally, one receiving advantage and the other not affected except, perhaps, in a minor way. Therefore, the symbiont receiving advantage is active and the other is passive. The active member may benefit by obtaining shelter, support, locomotion, or food (exclusive of the tissues of the passive member). Commensalism is the least involved of the three special types of symbiosis discussed here. It is most often disjunctive, but approaches being conjunctive in some cases and is either social or nutritive. If nutritive, however, the relation- ship is not antagonistic. Well-known examples are the pearlfish, which uses the cloaca of the sea cucumber for retreat, the conchfish, which uses the conch for the same reason, and the many bony fishes which live in seaweed or under the bells of jellyfishes. In the case of the man-of-war fish, commensalism is very near to being con- junctive since this fish is rarely seen out of the company of the Portuguese man-of-war and even feeds on some of the food that the jellyfish captures. A truly obligatory conjunctive commensalism is found in the whale barnacles which can only live on whales, the benefit of the association being all theirs. Mutualism In this symbiosis, both symbionts share the benefits of the relationship. Both species need not benefit in the same way; in fact, they usually do not. One may receive shelter or transport and the other food. The fine adjustments necessary for two animals to live and benefit together are sometimes very extraordinary. Usually mutualism is conjunctive, though it may be disjunctive. It is either social or nutritive and is always non-antagonistic. As in commensalism, examples from the sea are common and striking. The zooxanthellae of corals is one. Crabs show mutualistic relations with anemones in which the crab carries the anemone about on its back; the crab is protected by the stinging tentacles of the anemone, and the anemone receives transport and support. Some crabs even hold anemones in their claws and use these as weapons or food-getting devices. Other hermit BIOLOGY OF THE SEA-HOW THEY LIVE 39 crabs become clothed in a sponge, which gives the crab a home and the sponge transport. These are conjunctive examples. Disjunctive examples are found among butterfly fishes, which relieve large predators such as snappers of throat and body parasites with satisfaction to both parties. Parasitism In parasitism, which will not concern us very much in this book because most parasites live within the bodies of other animals and cannot be seen, the relation- ship is either conjunctive or disjunctive and is always antagonistically nutritive. One of the svmbionts is small and benefits by partaking of the tissues of a larger host. The host is always harmed to a greater or lesser degree. Parasitism is derived from a predator-prey relationship, in which the parasite has become adapted to feed on only one host (see lampreys. Chapter 8). In any case, the ideal parasitic relationship is not one in. which the parasite kills its host, for in so doing it destrovs its own home and its source of food. Parasites include various groups of worms, crustaceans, and protozoans among others. Manv of the phyla of the animal kingdom have parasitic members. Certain worms and the remarkable deep-sea angler fish, Photocoryniis, have males which are parasitic on the females. The male angler, for instance, attaches himself at an earlv stage of growth to the body of the female. He is a true parasite, receiving all food from the female, and degenerating to a small animal con- taining reproductive organs only. ADAPTIVE COLORATION Whether the colors of an animal are changeable or not, the colors it exhibits at anv one time are verv often adaptive in nature, that is, are of some benefit to the animal in its environment. Color adaptation can take several forms- concealment or warning, for instance— but in every case it is important to view the colors of animals with careful reference to the animal's environment, its behavior there, and the characteristics of the visual receptors of other members of its community. Otherwise, colors lose their meaning. Furthermore, it is necessary to realize that not all colors and color patterns are adaptive. Some colors have no adaptive significance. Thus, the tasks that are faced when studying colors are, first, to determine if the coloration is adaptive at all and, second, to determine what purpose the adaption serves. Sometimes the adaptions are obscure, and the diver must be very careful in his interpretation. Animals change color with response either to environment or to stimuli associated with courtship or fear. The colors of fighting sailfish become notice- ably more intense; the octopus shows a nearly white color phase when it is frightened or alarmed; blennies and gobies change color with sexual excitement during breeding. Adaptive coloration has been reviewed by Cott (1940) in a brilliant book on the subject. Cott begins his book by pointing out the very important fact that vision rests upon a three-fold basis. First, the physics of light is responsible for the intensity and quality of rays that emit from an object. Second, the physiology of the eye and nervous system gives a picture of that object to the 40 UNDERWATER GUIDE TO MARINE LIFE brain. Third, and most significant, the brain interprets that picture. The eye picks up a picture, but the brain does the interpretive "seeing," and, in general, the brain sees only what it knows. This means that past experience has a great deal to do with what is seen. The whole point of adaptive coloration is a play on the interpretive part of vision. Concealment — Cryptic Coloration In the case of concealment, adaptive coloration serves the purpose of making the adaptively colored animal harder for its enemies to detect or harder for its prey to notice. In other words, the clues by which detection of an object is made are rendered inconspicuous. Methods of concealment are four in number: 1. Color Resemblance. The most common method of concealment involves colors and color patterns that match the background. The flatfishes Qcolor photograph'), round sting rays (fig. 76) and the cowfish (/ig. 12) show this admirably in their resemblance to substrate. The octopus is another notable example of an animal that matches its surroundings. The red colors of moderately deep-water or nocturnal animals might be adaptive, making these species appear black. Several deep-sea animals and the larvae of eels, tarpon, bonefish and ladyfish are almost completely transparent, which is a protective lack of coloration that imitates the transparency of ocean water. Some animals change colors throughout life to match their changing environments. Cott gives the examples of the flying fishes, which are brown when young in sargasso weed and blue when adult in the open sea, and the sea slug, Aplysia Qcolor photograph), which changes from red when it lives on red algae when young to olive-brown when it lives on brown algae as an adult. Great numbers of bonv fishes and a few sharks are able to change color, being, in general, of lighter coloration in open water over a light bottom or near the water's surface and of darker coloration over a dark bottom or near dark places. 2. Obliterative Coloration. This adaption is encountered mostly in open-water species and is explained by figure 11. It involves countershading, in which the animal is darker above than below. This, in combination with illumination from above, makes the animal look flat and inconspicuous. Mackerels are excellent examples of this. The diver who has seen the barracuda swim toward him like a ghost has seen another good example of obliterative coloration. Countershaded fishes appear dark when seen from above, and when seen from below appear light against the surface. Most fishes are countershaded in addition to their other coloration adaptations. 3. Disruptive Coloration. In most cases, matching colors alone are not enough to conceal an animal. Methods which break form and outline, the greatest of all clues to identification, are very often employed. Stripes or bars are effective in breaking outline as shown by figure 12. The jaw outline of many bottom-living predators is concealed by the presence of fleshy, protectively colored tabs. Such is the case with toadfish, anglers, stargazers, sculpins, and scorpion fishes. BIOLOGY OF THE SEA-HOW THEY LIVE 41 TOP -LIGHTING & MO COUMTERSHADING COUNTERSHADING & UNIFORM ILLUniMATION TOP- LIGHTING & COUNTERSHADING Fig. 1 1 . An example of ohliterative coloration— the mako shark. The top figure shows the shark as it would appear if it were uniformly colored and lifted from above— a shadow is thrown on the lower side and the animal is conspicuous. The center figure shows the shark uniformly illuminated, hut colored darker on its hack (as the mako shark actually is')— still conspicuous. The bottom figure shows the shark as it actually appears in the water with a combination of top-lighting and counter- shading— flat and inconspicuous. (^Adapted from Cott, 1940.) The eye of many animals, being an especially noticeable part of the body, is frequently concealed by a stripe or disguised as those of butterfly fishes Ccolor photograph) and the porkfish QColor Plate 5). The eye also may be included in the pattern of the rest of the body, as seen in anglers, some groupers, and the scorpion fish, by extending body pattern and color onto the eyeball. Shadow Elimination. Animals are very often not betrayed by their own bodies but by the shadows that they cast. Shadows frame animals conspicuously, and the elimination of shadows is a valuable aid in avoiding detection. This method of concealment is common in bottom fishes. Depressed species such as rays, flatfishes, and batfishes cast little or no shadow. Rarely is only one of these four methods of concealment used by an animal. The mackerel shows ohliterative coloration, but its blue to blue-green ground 42 UNDERWATER GUIDE TO MARINE LIFE -a-a!^ Fig. 12. Four examples of disruptive coloration in tropical reef fishes. Upper left: the rihhonfish, Eques species, is vertically striped on its head and horizontally striped on its hody, which serves to break the fish into two major parts. Upper right: a midget bass, the harlequin serranid, Prionodes tigrinus, breaks its body into seg- ments with vertical bars and puts a stripe through the eye to mask it. Lower left: the Nassaii grouper, Epinephelus striatus, breaks the outline of its massive jaws by con- tinuing the pattern of stripes from the snout onto the lower jaw (note the con- spicuous tuning fork pattern on the forehead, a good identifying characteristic^. Lower right: the cowfish, Lactophrys tricornis, shows a disruptive coloration as well as color resemblance and protective habits, all of which cause the fish to look like a rock Qa large stone coral is at the right in this picture^. ri ■ ■ ■ < m i'^^V 1 SflB^^vjBHI j^ H tj p^ , vl^^fl H i |4 ^w^ l^^ii p r^^^HJl'^^X ' W "/ ^ m- '■■m,^ ^*4i ^\l I \. 4 J ^^^^^ \. BIOLOGY OF THE SEA-HOW THEY LIVE 43 color matches the open water, showing color resemblance as well. The sergeant major has bars for disruptive coloration, color phases which match environment, and countershading. The octopus is one of the few animals that shows only one method, color resemblance, though even in this case, its sometimes noticeable blotchings could perhaps be interpreted as disruptive coloration. The round ray is both protectively colored and flat to eliminate shadow. These examples should serve to illustrate the point. Do not look for one simple answer to concealment; it is most often a combination of interplaying factors. Conceal- ment of one part of the body may even be combined with advertisement of another part. For example, the four-eyed butterfly Hsh conceals its real eye and advertises a posterior false eye. Advertisement — A posemotic Coloration Diametrically opposed to concealment is advertisement. Some animals are conspicuously colored either to serve as a warning or as an aid for food-getting. Bright colors and striking patterns by themselves do not reveal to us whether an animal will be conspicuous or not. Rather, how the animal looks to the other members of its community is again the important criterion. Animals that advertise are frequently sluggish and stand their ground, especially when they are protected by having poisonous spines or flesh. They are sometimes gregarious, so that their conspicuousness is reinforced by their numbers, and they usually display themselves freely. There are two aspects to advertisement, action and color pattern. Sometimes these work alone, but they usually go hand in hand. Animals that are conspic- uous-looking in their environment also act in a conspicuous way. 1. Warning Colors. Conspicuous colors are sometimes taken on by animals to advertise the fact of their ill-tasting flesh, poisonous flesh, poison spines, or anything else about them that is definitely distasteful or dangerous. These animals travel more or less with impunity and with little fear of attack. This, of course, involves learning on the part of the predators which avoid attacking these animals. By advertising themselves, recognition is made easier, and there is reinforcement of dangerous qualities by striking appearance. Many sea slugs (nudibranchs) are very brightlv colored to advertise their distastefulness iColor Plate 10^. Warning display goes hand in hand with warning colors. The porcupine fish inflates not only in order to appear larger, but also as a warning of its dangerous spines and poisonous flesh. The sculpins erect their preopercular spines when molested. The squids and octopuses cause waves of color to pass over their bodies, causing hesitation on the part of predators. Toadfishes utter a grunt probably as a warning to advertise the power of their jaws. Some warning colors are adventitious and the result of symbiosis. Thus, the man-of-war fish achieves protection due to the warning colors or form of the Portuguese man-of-war's tentacles. Some Indo-Pacific demoiselles live in the protection of stinging anemones. One crab uses a live anemone, held in its claws, as a warning to predators. 44 UNDERWATER GUIDE TO MARINE LIFE 2. Conspicuous Localized Characters. In some cases, it is not the whole animal that is advertised but some specific part of it. Lumped in this category are a wide variety of fascinating examples, most of which involve some sort of misrepresentation or deception and which can almost be thought of as being a sort of concealment-advertisement combination, for in making one part of a body conspicuous, the other parts are made much less so. The one sort of exception to this general rule is exhibited by animals which conspicuously advertise a particularly dangerous part of their bodies. No deception is involved here. The sharp, preopercular spine of angelfishes is usually a bright yellow. The poisonous spiny dorsal fin of the European weaver, Trachinus, is a conspicuous black. The lancetlike white spine on the caudal peduncle of the blue tang contrasts with its otherwise blue body. Electric rays frequently have ocellar spots on their backs as warning marks. The other purposes of conspicuous localized characters are several and involve misrepresentation as stated above. First, conspicuous contrasting colors may serve to startle predators. This is a protective device involving bluff. The batfishes spread their pectoral fins to show a startling black and white banded pattern when threatened. Sailfishes are known to erect their huge dorsal fins when fighting or excited, and they may thus alarm aggressors. Second, conspicuous colors may serve to lure prey to the most dangerous part of an animal. The angler fishes use their first dorsal spine as a lure to attract prey, and the stargazers have a similar lure inside their mouths. In both of these cases, the lures are wiggled to make them even more conspicuous. Anemones have attractively colored stinging tentacles which lure small fishes to their deaths. Third, conspicuous characters may serve to distract attention awav from the most dangerous part of an animal. The cuttlefish, Sepia, and probably other squids as well, exhibits rapid waves of color on the posterior part of its body, which draw attention away from the tentacles. When the cuttlefish draws close to its prey, these relatively inconspicuous tentacles are shot out, and the capture is made. Fourth, colors and conspicuous marks are often used to misrepresent the posture or position of an animal or to direct attack to less vulnerable parts of the body. The combination of an eye stripe and an eyelike spot near the tail of the four-eyed butterfly fish creates the impression that this fish is swimming backwards. Longitudinal stripes through the bodies of such fishes as shark-suckers, cobia, and the rainbow runner make these animals appear to stand still while they are actually moving speedily forward. Squids, octopuses, and some sea slugs QAflysia^ emit clouds of inky fluid when annoyed, which confuse an enemy and mask their retreat. Some animals leave parts of their own bodies behind to confuse enemies. This habit is called "autotomy" and is well known in the cases of many lizards, including the glass "snake," which can discard its tail at a moment's notice, then grow a new one later. In the sea, the crustaceans are the best-known practitioners of autotomy. Many of them have a special autotomous joint where a limb may be broken off. This limb then wiggles BIOLOGY OF THE SEA-HOW THEY LIVE 45 furiously for a minute or two, distracting the enemy, and allowing the crustacean a chance to slip away unnoticed. The discarded limb may be eaten, but this doesn't bother the crustacean which simply grows another as good as the one lost. Disguise Disguise is like concealment, but in the case of disguise the resemblance to objects is real and not abstract. That is, animals are adapted not just to become inconspicuous by destroying parts of their own identity, but by resembling objects about them. Cott (1940) calls these animals "posers." Concealingly colored animals are "self-effacers." Disguise is put on either for protection or ambush as is the case with concealment. The sargasso weed community shows several good examples of disguise. The sargassum fish QColor Plate 10^ and some crustaceans that live in this weed are not only colored like the weed, but they bear fleshy, weed-resembling protruberances on their bodies. Se\'eral sea horses and pipefishes bear extensions of their bodies to make them look weedy. The angler, Lophins, is colored a'nd roughened to look like a flat rock. Scorpion fishes look like weedy pieces of coral rock. Filefishes imitate floating seaweed. The decorator crab, Stenorhynchiis, clothes itself with seaweed as an adventitious disguise. Several other crabs become clothed with sponges or anemones in a symbiotic relationship. Disguise is particularly interesting because it demands not only that an animal look like some foreign object, but also that it act like that object. The trumpet fish is brown and slender like a eoreonian stalk, but this eff^ect would be lost if this fish remained horizontal. Therefore, trumpet fishes assume a vertical position (usually near gorgonians) when alarmed and let the water currents sway them so that the illusion is complete. Similarlv, the angler fish stays motionless like a rock and the sargassum fish moves very little in sargasso weed in order to accentuate their disouises. Mimicry Mimicrv involves a resemblance even stronger than in disguise for it involves strong imitation of one living organism by another in both habits and appearance. The purpose of mimicry is restricted to warning and false warning. Two types of mimicry are known. First, if a relatively scarce, palatable, or nondangerous species mimics a relatively abundant, distasteful, or dangerous species, Batesian mimicry is said to exist. This mimicry involves learning by predators of the pattern of color and form of the distasteful species which is not eaten. Animals which imitate this distasteful species then are presumably also not eaten. However, the mimic can never become abundant or the learning of the warning pattern is weakened. For instance, the tasty viceroy butterfly mimics the distasteful monarch, but viceroy butterflies are much less plentiful than monarchs and are generally shunned by predators. Examples of Batesian couples in the sea are rare. Cott states that the English sole, Solea inilgaris, has a dark spot on its side which mimics the black, poisonous dorsal spines of the European weaver fish, Trachinns. 46 UNDERWATER GUIDE TO MARINE LIFE Second, if a number of species, all of which possess the same distasteful or dangerous attributes, come to resemble each other, the association of dangerous qualities with their appearance is made simpler for the predator, that is, learning is reinforced. Perhaps this is why the distasteful sea slugs (nudibranchs) look so much alike and are so brightly colored. ANIMAL BEHAVIOR The purpose of this section is to give some understanding of the principles involved in activities of marine animals as they adjust to their environments. The stories of fish behavior told by fishermen and others who come in contact with the sea are often interesting and fascinating, but for the most part they have been greatly misinterpreted. Often human qualities have been attributed to fishes and superficial conclusions drawn from only bits of behavior. To the diver who has to deal with the aggressive fishes in their environment this can be dangerous. A more rewarding approach can be taken by careful scrutiny of the evidence available. This involves looking into the physical and mental equipment possessed by marine animals, into the kind of behavior one can expect with a particular type of nervous system possessed by an animal, and into the part that sense receptors (visual, olfactory, etc.) play in their total behavioral adjustment to their environment. It may become apparent to the underwater naturalist as he studies the marine animals and their environment that these animals do not perceive their environment as the diver does. Many of these animals are deaf (the starfish, worms and snails, and probably some fishes). Many are blind, others have poor vision (sharks and rays), and some have excellent vision (some bony fishes, squids, octopuses). It is suspected that some fishes can hear sounds of higher frequencies than humans. Most fishes, such as the bottom feeders, have a highly developed sense of smell and are able to detect very weak chemical concentration. It becomes clear that the underwater world as seen by the diver appears different to each species, and what sometimes appears to be bizarre coloration, shape, or behavior in a fish to the diver can be explained as a method of survival adapted to the sensory and motor abilities of the animal that feeds on it. The diver perceives the underwater world mainly through vision. Man's vision underwater is far superior to that of fishes. He is very sensitive to color, he can focus over a wide range of distances with excellent visual acuity, and he is a good judge of distance because of binocular vision. But he is limited when compared to most fishes, for their behavior depends upon smell and lateral line receptors for contact with the environrnent. Man cannot smell underwater nor can he detect vibration as fishes can over great distances. This makes man primarily a day animal in the sea, for at night, unless he brings a light source with him, he loses his visual superiority and becomes as helpless as a fish out of water. Marine Invertebrates The behavior of marine invertebrates is for the most part dependent upon the type of structural equipment (action equipment, sensory receptors, nervous BIOLOGY OF THE SEA-HOW THEY LIVE 47 system) that a given species has and upon the environment in which it is able to live. It is able to live in a particular environment only if the environment continues to provide the stimulus (light, food, temperature, etc.) to which its receptors are capable of responding. The higher vertebrate animals (mammals) have advanced structural equipment (well-developed nervous system, sensory receptors, and action equipment) which, in contrast to invertebrates, is well integrated, making possible complex behavior patterns and modifications of behavior by learning from previous experience. Behavior of invertebrates varies, depending upon the efficiency of their sense organs, the type of nervous system, the kind of action equipment possessed, and it is also dependent upon the animal's past experience and inherited instincts. The intelligence of a species is determined by the degree to which sensory receptors cooperate with one another in governing the nature of behavior. The marine sponge, Sycon (phylum Porifera), is one of the lowest of invertebrates. After its free-swimming larva stage, it attaches itself to the bottom and becomes immobile, filtering its food from the surrounding water. There is no nervous system. There is only protoplasmic transmission of impulses from one cell to a neighboring cell, each cell serving the combined functions of action equipment and receptor. Pore cells are stimulated by water currents and chemicals, which start the beating of flagellated cells found lining the body walls. This animal has no other sensory receptor and its stationary existence makes it completely dependent on its environment. The sea anemone (phylum Coelenterata), more highly organized, shows an advance in sensitivity to its environment far greater than that of the sponge. It is able to move if the food supply becomes scarce, and it grasps its food by its tentacles and passes it to the mouth. These tentacles have contact sense and chemical sensitivity. The anemone also shows a reaction to light. The actions of coelenterates are integrated by the most primitive tvpe of nervous system known, the nerve net (/rg. 13^. The brittle starfish is part of a group called "echinoderms," which include the starfishes and sea urchins (phvlum Echinodermata). They are slightly higher in psychological position than the sea anemone because of their more efficient type of nervous system involving a nerve ring Qfig. 13^. The nerve ring is not a controlling center like a true brain but is a more efficient transmitter of impulses than the nerve net. This nerve ring is used mostly for controlling locomotion, and cooperation between parts is slightly better than in the coelen- terates. These advances, although the echinoderms have radial symmetry, usually allow whatever area is more strongly stimulated to take the lead in locomotion. Brittle stars are sensitive to light, moving about at night capturing their food. The light-sensitive cells are found at the ends of the arms. The sea urchin has light-sensitive pigmented spots on the body. Chemical-sensitive buds are found on the tentacles, tube feet, and the mouth area of brittle stars. The whole body is somewhat chemically sensitive with the ventral side slightly more so. Contact sensitivity is well developed and is important for locomotion. Contact receptor cells have hairlike projections found on the tube feet and tentacles. The sea urchin relies on the spines for its contact responses. Most starfish move about with the action of the tube feet. The britde star moves by the action of its arms, which are powered by large muscles. The sea urchin, when excited, moves by the spines. Otherwise it uses its tube feet. 48 UNDERWATER GUIDE TO MARINE LIFE The worms (phylum AnneHda) have a definite advantage over the starfishes and sea urchins in that the head end of the body is well supplied with sense organs, there being progressively less sensitivity toward the posterior end. This permits the head to take the lead in behavior and makes possible a more complex SAN OUORM, Nereis, NCRV0U5 SXSTEM CRUSTACEAN NERVOUS SYSTEMS STARFISH MCRVC RING pr.» SQUID NERVOUS S'ySTEM FISH HtRVOOS 5YSTCM OUFAtTORY 3ENSE Fig. 1 3 . Nervous systems and sense organs. adjustment to environment. The sandworm, Nereis, is an example of this group. It burrows into sand and has its chemoreceptors (smell, taste) mainly on the head end of the body, allowing a more directed food-locating behavior than is present in echinoderms. One other factor important in this group for adaptive orientation is the development of a centralized nervous system involving a brain Cfig- 13). These improvements of nervous structure with concentration on the head end integrate the body processes in a better way than the lower animals previously discussed. The marine worms have paired ocelli (simple eyes) which are sensitive to light, and the worms react by hiding by day. Chemical sensitivity is mainly located on the tentacles on the head end although there is some sensitivity on the tail and body appendages. There is strong sensitivity to vibration from the substrate. Some members of the phylum Mollusca (snails, octopuses, and squids, but not clams and oysters) have a much better-developed head end, and this anterior part of the body exercises greater control over other parts of the body than that found in the worms. This superiority is also evident in improved sense receptors and nerve conduction. The groups of this phylum have a great variety of behavior, which ranges from the permanently attached oyster and the slow- moving clam to the fast-moving octopus and the squid. Chemical sensitivity is present in the tentacles of the snail and scallop and in the sucking discs of the BIOLOGY OF THE SEA-HOW THEY LIVE 49 octopus and the squid. The snail has hght-sensitive receptors at the end of the tentacles. The octopus and the squid have eyes that are almost as highly developed as those of the mammals. This has been one of the chief reasons for attributing a high intelligence to these animals. These highly developed receptors and a well-developed nervous system (fig. 13^ place the octopus and the squid among the highest in intelligence among invertebrates, but their apparent lack of sensory and motor integration does not warrant placing them higher than fishes in psychological standing. The scallops have a row of light-sensitive receptors on the mantle edge near the base of the tentacles. Clams have light- sensitive cells on the inner surface of the siphon. An elongated muscular foot in clams provides locomotion. Sea slugs and snails have a gliding movement caused by waves of muscular contractions on the foot. The octopuses move on the ground by using their tentacles and in open water by jet action of expelled water. The phylum Arthropoda is a very diverse group composed of almost a half-million species, of which crabs, shrimps, and lobsters are part of a group called "crustaceans." This group lies above the Mollusca in psychological standing because of the existence of a brain and the ladder-type nervous system, which has a pair of ganglia in each body segment. This results in better integra- tion of the parts of the body than is found in molluscs. The head with the specialized sense receptors plays an important part in the animal's orientation. The crustaceans have chemoreceptors in the antennules, or small antennae. These receptors are used to locate food by picking up juices diffusing in water from some distance away. They are also used to detect food by contact. Substrate vibrations are picked up with legs and body. Most of the crustaceans have compound eyes, which are especially well adapted for picking out moving objects. Fishes Much of the behavior of the invertebrates is closely related to the extent of the development of their nervous system. In the vertebrate fishes, as well as in the invertebrates, the nervous system is principally a bridge between the sense organs and action equipment, but it is distinguished from the invertebrate system by having a spinal cord, which runs along the back, associated with a much better brain (/ig. i3). This tubular conductor permits a unification of the action equipment not possible in the invertebrate ladder-type system. Although fishes have this advanced tubular system, they still retain a form of segmental system, which acts like a ladder-type system (not, however, evolutionarily related to the ladder system) and which accounts for some of the stereotyped behavior found in fishes. This segmental system is retained in part by all higher vertebrate animals. SENSE RECEPTORS OF FISHES IN GENERAL VISUAL The eyes have a flattened cornea with a fixed-focus lens. Vision is usually 20-30 per cent binocular because of the position of the eyes on the sides of the 50 UNDERWATER GUIDE TO MARINE LIFE head and each eye is able to move independently. The eyes are able to focus only on near objects. Distance focusing is limited. Fishes have a wide-angle field of vision. Careful experiments have shown fishes to have color vision, but it seems weak. Verrier (1928) found that the structure of the retina does not permit good visual acuity. Acuity seems to be similar to the peripheral regions of the human eye. It allows good vision of movements of objects, but perception of form seems defective. The eyes are used by all fishes to some degree except for a few nocturnal species. OLFACTORY Ciliated cells in sacs, which have an opening to the exterior nostrils are able to detect the presence of weak concentrations of chemicals in the water Cfig- 13). These receptors are used for locating food at a distance as well as for detecting chemical gradients in water. Smell is used to some degree bv about half of all fishes. TASTE There are many grouped cells with hairlike projections in and around the mouth, on barbels (when present), on growths which hang from the lips, or often covering the body surface. These receptors detect chemicals emanating from food close to the body. The contact taste receptors in the mouth cavitv are used for the selection of food and rejection of foreign objects. Some fishes, mainly those that feed by sight, swallow objects without much attention to selection, and often strange objects are found in their stomachs. Taste seems to function in all fishes. THE LATERAL LINE Lateral line receptors are composed of ciliated cells clustered at intervals along a lateral line on the sides of the body from head to tail and on some fishes branching on the head (fig. 13^. These receptors are sensitive to the intermediate vibrations between those perceived by the ear and those perceived by the body surface. While the auditory sense can detect vibrations of high frequency, lateral line receptors can detect only low frequency water movements. Skin contact receptors are sensitive only to solids, water pressure, and very slow vibrations. Lateral line receptors are used to locate prey, to avoid predators, and to localize obstructions by detecting direct and reflected water movements. In some fishes at night, these organs become the dominant sensory receptors. They are also sensitive to relative water pressure as well as to direction and rate of flow of water. ALIDITORY The auditory sense in fishes is poorly developed. However, there is evidence that some fishes can hear sound frequencies of up to 8,000 cycles per second or more. Some fishes are not able to hear at all. The ears of fishes are not visible on the body surface and are used mainly for equilibrium and orientation. It is possible that some fishes use vibrations reflected from the surface, the shore, and the bottom as aids to orientation. COMMON EEL- SPOTTED MORAV HOUN DFI SH ^jjyeyiMHHH»GWiiU»Mi«-: PIPEFISH MORTHERNJ SEAHOFtSE SHORT-WIN&ED FL.YIfslO FISH COLOR PLATE i COLOR FLAl E 1 BIOLOGY OF THE SEA-HOW THEY LIVE 51 THE ROLE OF SENSE RECEPTORS IN THE BEHAVIOR OF FISHES When the environment of fishes is compared with that of land animals it is found that the sea is much less variable than land. The chemical content and temperature of the sea vary little, food is easier to get, and there is no weather as we know it on land (except near shores). This stability of environ- ment requires a smaller repertory of adjustable activities, which accounts for the lack in sea animals (except for the re-entrants) of the advanced psychological organization that is found in the higher land animals. In spite of the fact that fishes live in a comparatively constant environment, there are a great many types of habitats in the sea— coral reefs, seashores, open sea, and the great depths. Physiological factors such as weak vision, a good olfactory sense, possession of a swim bladder, and well-developed action equip- ment determine in which of these environments a species will live and a species' environment determines to a great extent its type of behavior. In the environment of a particular species, the pattern of behavior is to a certain degree determined by the sense receptors it possesses. This close relation- ship between sense receptors and characteristic behavior is clearly evident in the feeding habits of the rays. These animals have weak eyesight and a well- developed sense of smell and touch; they are for the most part bottom-feeding animals (except the devil fish and the manta), which use their smell and touch organs to locate crustaceans and molluscs in the sand. The jacks are an example of fishes with excellent eyesight, which obtain their food by darting at or hunting down small schooling fishes at the surface. The important place of sensory equipment in the activities of fishes is pointed out by Jarmer (1928), whose studies show that when a fish is exposed to a visual or olfactory stimulus (depending upon which is its main operating sense), it automatically bites. This is characteristic behavior of fishes in general, resulting from the possession of a primitive type of vertebrate nervous system. In the higher vertebrates, as in the mammals, when one sensory system makes contact with a prey, the animal becomes alerted and other sensory systems are brought into play for a better orientation before striking. These animals have the ability to delay their responses to a stimulus because of their advanced nervous system, whereas in fishes the stimulus brings on a direct and automatic response. An example of this type of behavior is given in the case of the diver who dove into the water within a few feet of a barracuda, a visual feeder. The fish turned instantly and struck the swimmer at the moment of the splash. Approached underwater, the same barracuda would probably do no more than show curiosity. The diver's splash and movement resembled the splash and movement of the prey of this fish, that is, he resembled the characteristic visual feeding stimulus, the splashing at the surface of a wounded fish or of schooling fish. The stereotyped visual feeding behavior of the barracuda was demonstrated when the authors placed freshly killed fish on the bottom near a barracuda, resulting in no feeding response. But moments later when a moving, wounded fish was presented, the barracuda swiftly struck. A shark with weak eyes and a strong sense of smell, on the other hand, would have reacted readily to the smell of the dead fish on the bottom and also to the wounded, moving fish. Conse- quently, an excellent method for understanding fishes' behavior is to group them according to the way they secure their food. They generally fall into two 52 UNDERWATER GUIDE TO MARINE LIFE groups— the visual feeders and the chemical feeders (mainly through sense of smell)— with one system dominating consistently. 1. Chemical Feeders. The chemical feeders, such as the mullets, rays, and most sharks, among others, usually have poor eyesight, but they have a well-developed olfactory sense supported by touch and taste organs, and these are used to locate bottom life and carrion, and to sift organic matter from the bottom. The olfactory system is a general "exciter," which increases searching activity around the area where the scent is detected, and as the fish gets nearer to the source of the scent, the paired olfactory organs direct the fish to the food. Vision and the contact sense probably come into play in the immediate vicinity of the food. 2. Visual Feeders. Many species of fishes respond more to the visual stimula- tion of a moving object than they do to chemical stimuli (barracuda, bluefish, tuna, cod, blennies, etc.). These are the fishes that pursue their prey, and vision is the dominant sense used in making contact with their environment. These fishes, upon immediate sighting of a moving prey or preylike object across their fields of vision, respond with a compulsive and direct strike. This response appears only when the prey lies within the field of visual acuity of a given species, an acuity which in fishes is generally limited to short distances. The eyes and body move simultaneously, as if sighting and striking were one movement, so powerful is the role of vision in these species. The voracious bluefish is a good example and is well known among fishermen for striking at any object that falls within its visual field. The eyes have a rather powerful, direct and stereotyped influence on behavior in the visual group, almost always producing the biting response, whereas in the chemically dominated group, olfaction plays the part of a general exciter and accounts for less stereotyped behavior. The other sensory systems generally support the dominant systems, but they may take the place of chemical or visual senses as the dominant sense under some conditions; for instance, at night or in murky waters the lateral line receptors often become the dominant sense. These exteroceptive systems in fishes generally work independently with little intersensory integration. In the behavior of the higher vertebrates, such as some amphibians, reptiles, birds, and mammals, no one sensory system dominates consistently in the way that fishes' behavior is dominated by vision or olfaction. One sensory system, when stimulated, brings another system into action with overall integration of activities. Bringing other sensory systems into play requires more time, gives a better orientation to the stimulus, and allows greater flexibility in responding, enabling the animal to make a better and wider adjustment to its environment. CHAPTER *| MAN AND THE SEA, I— Photography and Equipment UNDERWATER PHOTOGRAPHY This section has been written to help the underwater swimmer to explore, document, and enjoy underwater life. With photography as a tool, the swimmer can better identify the different species that swim about him as well as make possible contributions to science. The aesthetic aspect of the underwater world has been one of its greatest attractions, an attraction that is hard to describe. Only through photography can it be brought to the surface. One has only to see coral, which takes endlessly exciting formations, and the angelfishes, which are the colorful ballerinas of the underwater world, to realize the possibilities for taking artistic photographs. The graceful movements of schools of fishes as they swim about the reefs, the birdlike flying of the rays, and the sleek beauty of sharks as they move through the water— all make fascinating pictures. There are startling nature photographs to be taken: the gigantic whale shark standing on its tail feeding on plankton, the manta ray leaping out of the water as it drops a newborn in midair and heralds the birth with a thundering slap as it hits the water, the parrot fish standing on its tail with its mouth open while little wrasse clean its teeth and face by eating the food particles left there. There are hundreds of similarly interesting instances in the lives of sea animals yet to be recorded on film. Effects of Sea Water on Light Underwater photography is mainly for clear waters. Ideal locations are tropical waters such as those surrounding the Bahamas, where visibility is sometimes better than 100 feet. Excellent photographs can be taken, however, in waters with visibility of 15 feet or better. Water conditions which allow visibility less than 15 feet limit picture taking to close-ups. Penetration of sunlight into the sea is limited by the roughness of the surface, the density of water, and the turbidity caused by the presence of organic (planktonic) and inorganic (silt) particles. These particles cause an effect called "scattering," which tends to make pictures taken under water at 53 54 UNDERWATER GUIDE TO MARINE LIFE a distance lack contrast and three-dimensional effect. This is similar to shooting pictures in a fog. Because of scattering, the camera should be as close to the subject as possible in order to obtain sharp contrast. Sea water also has the effect of absorbing colors differentially, the red end of the spectrum suffering the greatest loss. The deeper sunlight penetrates, the greater the loss of colors. The deterioration of color from the surface to the depths in clear water is as follows (for photographic purposes): At about 3 to 4 feet from the surface, red begins to disappear noticeably; at 15 feet so little remains that photographs taken without filters show no red. Orange goes out at about 30 feet, yellow at about 65 feet. Below 65 feet only green and blue remain, until at approximately 1,800 feet all light visible to the human eye disappears. These figures are approximate and dependent upon water conditions. (Chapter 1.) Exposure Under Water Exposures under water depend upon surface conditions, the type of bottom, and the turbidity of the water. Because of turbidity, light-meter readings are more accurate the closer the subject is to the meter; the greater the distance, the less reliable the reading. In shallow water with only one kind of bottom (e.g., sand) generally one reading is sufficient for all exposures as long as the surface conditions remain the same. Clouds and water currents that stir up particles are factors that change lighting conditions. In calm water only about 3 to 4 per cent of the incident light is lost because of reflection, but choppy water results in a 20 to 30 per cent loss in intensity at 2 or 3 feet below surface. On bright calm days in clear waters, exposures at about 5 feet below surface are almost the same as above the surface. To 30 feet, use a meter or increase the lens opening one more stop. A direct-reading light meter (Weston) can be mounted in an underwater housing in line with the lens for constant readings. Shutter speeds under water are generally the same as above water. A general working shutter speed is 1/50 to 1/100; for static subjects (e.g., coral) 1/25 may be used for increased depth of field, provided the camera is not in motion. The success of increased shutter speeds for taking fast-moving objects is somewhat limited by the low light-level conditions. Camera Under certain conditions any type of camera can be used to take underwater pictures, but the most successful underwater pictures are taken with the camera arrangement which reduces mechanical handling to a minimum. This allows the photographer to take advantage of opportunities and to concentrate on safety and subject. These are important considerations when one remembers the equipment the swimmer has to carry, the constant motion, the attention that must be paid to safety, and the time element involved in use of the aqua-lung. Also, if an exciting specimen swims by, the swimmer with an ordinary camera could not take more than one picture if he had to struggle with setting the shutter speed and lens opening, cocking the shutter, and advancing the film. Another factor operating to complicate picture taking is that the reaction time MAN AND THE SEA, I 55 Fig. 14. Underwater 'photographic equipment, (a) Plastic housing for Dejur Spectator 8nim. inovie camera and ^Aleston Direct Reading exposure m.eter. (b) In- expensive Voit plastic camera hag for beginners, (c) Praktina 35mm. single-lens reflex camera with rapid sequence spring motor, (d) Plastic housing for a 35mm. single-lens reflex camera vAth a Weston Master exposure meter, (e) Cast metal housing for Robot 35m,m,. rapid-sequence camera. The plastic cases were assembled by plastics dealer and the controls added by the authors at a total cost of less than twenty-five dollars for each case. 56 UNDERWATER GUIDE TO MARINE LIFE of fishes and most marine animals (about one twenty-fifth of a second) is faster than that of humans. This means that if a photographer and a fish sight each other at the same time, the fish could be at least halfway turned around and headed away before the photographer raised his camera. The 35mm. camera is widely accepted as the camera best suited to underwater work. Both camera and case are small and easy to handle. It takes a thirty-six- exposure roll of film, enabling the photographer to take fewer trips to the surface to reload. Some 35mm. cameras have bulk loading attachments which take up to four hundred frames. There are 35mm. cameras available which have a spring mechanism that transports the film and cocks the shutter, leaving only the trigger release to be pressed for rapid sequence exposures. The Praktina Cfig. 14} and Robot meet all these requirements. Movie Cameras — For Underwater Use When selecting an 8mm. movie camera, one should: 1. Select a camera with an f/1.9 normal and f/2.5 wide-angle lens. 2. Select a camera which has the longest film run possible on one winding (e.g., 10 feet). Long scenes are effective, short scenes are meaningless. A long film run cuts control manipulations and allows concentration on the subject. A good example of such a camera is De Jur (/ig. J4). 3. Select a camera you know won't jam in the middle of an important scene necessitating a trip back to the boat to clear a jam. Jamming is time- consuming, troublesome, expensive, and results in the loss of previously exposed film. The factors which determine the selection of a good 8mm. camera for underwater work also apply to the 16mm. camera. The 16mm. camera should load the largest footage possible; short footage cameras mean many trips to the surface and loss of opportunities. Many of the professional photographers use motors attached to their 1 6mm. cameras to eliminate hand winding. The Use of Wide-Angle Lenses Under Water With both motion-picture and still cameras, a wide-angle lens is necessary when one wishes to encompass larger subjects without having to shoot at great distances. Because of the refraction of light through glass, air, and water, objects under water appear one-third larger or underwater distances appear one-fourth less. This affects the goggled eye as well as the camera, making it necessary for the photographer to back off farther to fill the picture frame and to set the focusing scale at 75 per cent of the actual distance. The wide-angle lens helps overcome this by permitting the camera to be taken closer to the subject. The lens offers another advantage in that it has a larger depth of field, which means one can shoot almost without chanoing focus. This is highly desirable, consider- ing that most underwater pictures are taken at a low light level which calls for MAN AND THE SEA, I 57 wide apertures— resulting in lower depth of field. It is important when selecting a wide-angle lens for scientific work that there be no distortion at the edges and corners— the type of distortion found with some extreme wide-angle lenses. This distortion is acceptable for amateur work. The Normal Lens and Underwater Close-up Photography There are all forms of interesting life— coral, crabs, small reef fishes, plandike animals, etc.— that require close-up photography. Most camera range finders are useless under water, but single-lens reflex viewers for critical focus give a large enough image to be useful. The depth of field at short camera-to-subject distances is slight (even with small apertures) and a miscalculation of only an inch or two is enough to cause out-of -focus pictures. (A workable rule for determining depth of field for any aperture is one third the distance in front of the subject facing the camera and two thirds behind.) The wide-angle lens is desirable for photographing large marine subjects, but it necessitates that one get close to the subject in order to fill the negative. This lens is not effective when working close to small moving subjects; small reef fishes are generally -skitterish when approached closely, within a few feet, and swim off or become hyperactive. The use of a reflex camera equipped with a normal lens set for distances of 18 inches to 3 feet gives a slight telephoto effect allowing the photographer to back off so as not to disturb the subject yet fill the negative. This is contrary to the general belief that only wide-angle lenses are suitable for underwater work. With nonreflex cameras a measured stick is useful to determine distances when photographing static life, but drives off the active life. Since most small fishes are constandy moving about a small area around the reef, a ruled stick can be used to determine probable distances where the subjects have a good possibility of appearing. This involves studying subject behavior before shooting, but it can be very rewarding. Keep in mind that the yardstick appears one third longer under water. Color-Correcting Filters Color correction filters will function to a depth of approximately 50 feet. Beyond this depth in most waters not enough of the remaining red, orange, and yellow light is left (because of the effect of selective absorption, scattering of light, and lack of intensity) to make filtering practical. Some source of arti- ficial light is necessary to restore color below this depth. There are shallow-water photographers who feel that filters are unnecessary and that the blue-green effect is desirable and truly representative. This is a matter of individual preference. The authors feel that the complete elimination of the blue-green effect results in the loss of the unique color tones of the underwater world and prefer the use of filters which do not completely eliminate this effect. For waters that tend toward blue with litde green use a CCR series (color- correcting red) filter. If the water tends toward yellow, that is, has a definite green tinge, use a CCM series (color- correcting magenta) filter. For a clear tropical 58 UNDERWATER GUIDE TO MARINE LIFE waters like the Bahamas the authors prefer CCM :^20 or #30. These filters will give an overall balance of color with a slight blue-green effect without introducing a large filter factor. Be sure to include the filter factor along with the exposure meter reading to get your camera exposure setting. For instance, the filter factor for a CCR #30 filter is 1.8. Black and white pictures show increased contrast if a filter of the series CCY (color-correcting yellow) is used. A CCY #10 or #20 is adequate for most waters. All these filters are available in gelatin sheets at low cost, and they may be cut and mounted on the back or front of still or movie lenses, or pasted on the inside of the camera case in front of the lens. When using filters and a meter it is possible to put the same filter over the meter to eliminate exposure computation involving filter factors. Film Films for underwater photography are the same as used above water. Consider- ing that most underwater lighting conditions are of a low light level, the best color films are the faster speed Anscochrome and Ektachrome. Ektachrome gives good results, but Anscochrome gives better color rendition under water. The exposure latitude is one stop under and over ideal exposure for good color rendition. Black and white films have a wide exposure latitude with three stops above and below ideal exposure giving a workable negative. The best films are Panchromatics with an ASA 50 emulsion speed or better. For caves and poor lighting conditions found near reefs, take advantage of the faster speed films. However, increased speed of film usually results in increased graininess which cuts usability when enlarging. Any color film like Anscochrome 16mm. reversal movie film that has an ASA 32 rating and gives excellent color rendition at low light levels is well adapted for underwater use. This film allows the underwater photographer to use smaller lens openings and to shoot under conditions which formerly did not allow color. Anscochrome reversal film allows a latitude of one stop over and one stop under ideal exposures. Thin emulsion black and white films have extremely high resolving power, very fine grain, and fair contrast which make them good for underwater work. The exposure latitude of thin emulsion films is about the same as ASA 32 color films. Film manufacturing is undergoing great changes to which the underwater photographer should be alert. Artificial Lighting There are tremendous photographic opportunities in waters shallower than about 60 feet for the amateur and the professional. Below 60 feet, in caves, and where poor lighting is found, artificial lighting units such as flash attachment and strobe light units that can be converted to underwater should be used. These units can bring light and color to the blue-green depths. It would MAN AND THE SEA, I 59 take many pages of technical information to go into the techniques of artificial lighting in deep waters. There are excellent books devoted to the subject. (See Bibliography.) Underwater Camera Techniques A good technique to ensure not missing any opportunities when exploring is to take along two cameras, one in hand and the other hanging from a neck strap. Each camera might be set for special work: one set at IV2 feet to 3 feet for close-up work and the other set for distance shots; one with color film and the other black and white; one a still camera and the other a motion- picture camera. It is a frustrating experience to have a camera loaded with black and white film when a good color situation presents itself Whenever possible, avoid shooting into open water, but try to frame sub- jects with a reef, coral, or a school of fish. Pictures of fishes with people in the background are also interesting and show relative size. Special effects are obtained when subjects are shot against the surface or with back lighting. Generally the best results are obtained with side lighting. Get as close to the subject as possible to avoid pictures without conttast. Underexposing slightly and overdeveloping negatives will increase negative contrast and permit the use of a smaller lens opening for increased depth of field. Sunshiny days are best for color, and then around middav. Avoid fogging and light streaks in the film by reloading in the shade or, if in an open boat, under a canvas or towel. The lack of gravity experienced under water permits all sorts of special camera effects equal to the most expensive studio setups. For example, by using the feet for propulsion and handholding a movie camera, one can get a dolly effect as well as all sorts of angle shots. For an exciting experience as well as a method that affords wonderful photo- graphic opportunities, put on an aqua-lung with extra weight, and sit on the bottom in an area where there is much life. As soon as the fishes come to see you as an unthreatening part of their environment, they will swarm around you with curiosity or completely ignore you, allowing you to study and photo- graph them in their natural state. Underwater photographic technique is by no means standardized. There is much that the photographer can still do toward evolving new techniques. Underwater Housings In order to take pictures under water, it is necessary to have a watertight case for your camera. If you own one of the more popular cameras, such as the Robot, Ditto, Leica, Argus, Rolleiflex, Contax, Nikkon, or Canon, there are manufactured cases available. (Write the manufacturer of your particular camera for information.) Inexpensive camera arrangements for the beginner are shown in figure 14. Many photographers build their own cases because of the expense of manufactured models and because many cameras do not have cases designed for them. 60 UNDERWATER GUIDE TO MARINE LIFE Fig. ]5. Some diving gear for snorkeling and self-contained diving. Snorkel tubes, mask, emergency air pack, and flippers are in the foreground. Aqtia-lungs and air coinpressor for filling tanks are shown in the rear. DIVING GEAR Basic Equipment The basic equipment necessary to start underwater exploring includes swim fins, mask, and snorkel. The mask or goggles is a must, for the eyes see every- thing out of focus without it. One must constantly keep in mind that the mask makes objects appear one third larger or distances one fourth smaller. The mask should fit comfortably, allow maximum vision, and not leak water. The flippers, or swim fins, will greatly increase swimming ability and speed as well as free the hands for photography and safety. They wall also give better balance and maneuverability. The fins should feel comfortable— if too tight they will MAN AND THE SEA, I 61 impede circulation, leading to cramps. An allowance of an extra size should be made if one plans to use a full rubber suit that covers the feet. The snorkel tube Cfg- ^5) is usually made of plastic, rubber, or metal. It allows the swimmer to breathe surface air with the head under water and the eyes out of the sun. One of its important features is that it permits the bodv to float just a few inches below the surface at the point the body would normally be in a dead man's float. Consequently, little energy is expended in keeping afloat as compared to treading water without a snorkel. Thus, the swimmer can swim great distances without tiring. The snorkel is a very simple device to use. The valveless tube is used by the experienced swimmer and the ball-valve type by beginners. A good technique for almost doubling your breath-holding time under water is to take many quick, deep breaths before taking the last one for the plunge; this rapid breathing before diving builds up oxvgen in the blood. Safety under water, as in all other activities, works best when it is an attitude rather than a set of rules, for trouble can come from unexpected directions. Keep on the alert for the unexpected. The diver should carry at all times a very sharp, double-edged, pointed knife for emergencies. He should also carry an emergency air pack (underwater parachute), which, responding to a squeeze, will inflate, take the diver to the surface, and keep him afloat. The Aqua-Lung or Scuba This self-contained underwater breathing apparatus gives the swimmer almost all the freedom that a fish experiences. It allows the diver to go into deeper water with litde effort, without the need to surface for air. It is useful to the naturalist and especially useful to the photographer, for he can sit and wait for opportunities. (Figure 131, the "kissing" grunts, was taken after one of the authors remained still on the bottom at 35 feet for almost three quarters of an hour.) The experience of using the aqua-lung in clear waters, abundant with marine life, can be one of the most memorable in one's life. It can also be the last. It is such a simple device to start using that the novice can easily get himself into trouble. The authors recommend strongly that no one attempt to use any self-contained underwater breathing apparatus without at least six hours' training under competent instruction. Besides instruction on how to use the lung, a portion of the six hours should be devoted to the part water pressure plays in the aqua-lung's operation and to the eff^ect of pressure on the body. A beginner would be wise to start with the snorkel, which will make him conscious of his breathing, then move to the aqua-lung. BASIC SAFETY RULES 1. Never dive alone; use the "buddy" system, and keep within sight of each other. 2. Plan in advance what is to be done in an emergency. Be sure all members of a diving party know the technique for executing the back pressure-arm lift method of artificial respiration. CHAPTER 4 MAN AND THE SEA, II— Dangers and Chumming DANGERS OF THE SEA When one thinks of dangers of the sea, sharks, barracuda, moray eels, and other large, predatory animals immediately come to mind. These animals can be dangerous, but they are not usually every man's greatest danger in the sea. Dangers come from two sources primarily, from within the diver and from the sea. By far the most potent of these dangers are those that derive from within, for the diver can carry within himself the attitudes and knowledge that make the sea, like an automobile, either a safe or dangerous place. The diver must develop an attitude of safety and respect for the sea which involves thorough knowledge of equipment (Chapter 3) and a resolve never to dive alone or out of sight of other divers. The diver should take every precaution that his equipment gives him every bit of advantage possible. The diver must have a knowledge of his own physical limitations which involves the general principles of diving physiology (bends, nitrogen intoxica- tion, squeeze, lung collapse) and the peculiarities of his own body. This knowl- edge will keep him from the overconfidence which would lead him into situations beyond his ability to handle. The diver must equip himself with a knowledge of the sea's inhabitants, their identification, habits, and adaptations to their environment. This, of course, implies to a certain extent that the diver is often free from possible injury from the life of the sea to the extent that he is skilled in biology. One of the diver's most common errors is attaching such anthropomorphic qualities as cowardice to animals of the sea. This is a fundamental and even dangerous mistake in trying to understand behavior patterns and mentalities very different from our own. One of the important dangers that comes from within is fear, which leads to panic or even immobilization of the diver. This fear usually arises from misinformation and lack of understanding of the sea and its life. Consequently, the way to eliminate fear is to know and understand the sea, in which environ- ment the diver is a misfit, able to exist only in a highly organized state. Even with his technology, the diver moves under water with disadvantages and fears similar to those which primitive man experienced in the Stone Age, being subject to similar dangers from the elements and from wildlife. Man's real 62 MAN AND THE SEA, II 63 superiority in the sea, an environment which for him is foreign, is not physical, but is psychological and intellectual, for the physical elements of this environ- ment prohibit man from being as effective in the sea as he is on land. It is not the authors' intention to review the life of the sea minutely for dangerous species. This would require a book in itself (see Bibliography). Instead, the following is a review of the ways in which animals and plants are dangerous, with examples of each. The usefulness of this review depends a great deal on the common sense of the diver. For example, a complete list -of all fishes with sharp spines, sharp gill rakers or heavy teeth would include most of the fishes in the sea. The diver must use his own judgment when handling these fishes. Details of the dangers of the species mentioned are described in Chapters 7 to 10. Passive Dangers STINGING, IRRITATING, BITING, AND SPINY ANIMALS Most dangers in the sea are passive ones, such as fire coral and the poisonous spines of some fishes, which become dangerous Only through the actions of the diver. It is a good rule never to touch any 'plant or animal which cannot he properly identified. Wear gloves. The dangers from stingers, poisonous spines, etc., exist in a very great number of species. Usually, such mechanisms are used as modes of defense by these creatures, though in some cases they are also food-getting mechanisms, such as the tentacles of the Portuguese man-of-war or those of anemones. But in all cases, man plays the aggressor, and therefore he can almost completely control this aspect of danger merelv bv knowing where the dangers lie. A good example of this is the case of the diver who, having caught an octopus, slung it over his shoulder and was bitten on the back as he carried the octopus up the beach. He died from the poisonous bite in a short while. He was the victim of the now popular belief that octopuses are harmless. There is a huge array of species spread throughout most of the phvla of the animal kingdom that can sting or injure with spines and claws. Some of these species advertise their harmful parts, but some do not. On the whole, spines and stings are defensive in purpose and do not cause serious injury, but a few are very dangerous. For this reason the very dangerous species are listed separately. Dangerous examples: Portuguese Man-of-War; Lion's-mane jellyfish; Sting ravs; Eagle rays; Chimaera; Scorpion fishes; Sea snakes; Octopuses; Squids. Less dangerous examples: Red tide protozoan; Fire sponge; Poison-bun sponge; Manv sponges with sharp spicules; Many coelenterates with large nematocysts, including some hydroids, anemones, and fire corals; Parasitic flukes which cause "swimmer's itch"; Bristle worms of several species; Biting worms such as Nereis; Cone snails; Sea urchins; Crabs and lobsters with strong spines or claws; Spiny 64 UNDERWATER GUIDE TO MARINE LIFE dogfish; Bony fishes with strong teeth such as the toadfishes; Bony fishes with fin spines such as catfishes; Bony fishes with opercular spines such as the squirrel fishes and the angelfishes. This list covers almost all the species that can be dangerous to the diver. However, judgment and common sense are the best guides for handling most animals with spines, claws, or teeth. Many fishes have spiny gill rakers inside their gills which can inflict cuts when handled. ENTANGLEMENT AND ABRASION In extensive beds of algae, particularly kelps, it is fairly easy to get dangerously entangled in these long, tough plants. Several corals are very sharp and can inflict cuts that are painful, liable to infection, and slow to heal. The reason for the slow healing and extra pain of coral cuts is not known. Any cut that bleeds freely should be attended to at once because of the danger of infection. A diver, simply for safety's sake, should not remain in the water if bleeding because of the danger of sharks. The diver must use his own judgment in this area, staying away from kelp and branching coral or rocks in rough water. POISONOUS FLESH Almost all animals of the sea are good to eat, but the flesh of some can produce results varying from an upset stomach to death. The maki-maki or "deadly death" is a Polynesian species of puffer fish with extremely virulent flesh. Fortunately, North America is relatively free of species with poisonous flesh. The poisonous parts may either be the muscles or some other organ of the viscera such as the liver, gall bladder, or reproductive organs. It is sometimes rather difficult to trace down what organs of an animal are poisonous or under what conditions a normally nonpoisonous species may become poisonous. For instance, some species, such as the yellow-fin grouper, barracuda, and various jacks among others, have poisoned people, but very little is known of the causes responsible for this. It is suspected that these fish sometimes eat the flesh or eggs of poisonous fishes. This is a subject about which very little is known. Because more and more animals of the sea are being used for food, this subject is becoming very important. Some well-known examples of poisonous animals: Many filter-feeding clams and oysters (they become poisonous in the warm months due to accumulation of toxic plankton in their digestive tracts); Six-oilled shark; Chimaera; Escolar; Some triggerfishes; Some filefishes; Most puffers (Chapter 9— Plectognaths); All porcupine fishes and boxfishes. Active Dangers THE AGGRESSIVE SPECIES These are few in number, and each of these is considered in detail individually in later chapters. All predators are aggressive for food and will attack only if the diver falls into their feeding patterns, which usually he does not. Many of these animals can be driven off by taking the psychological advantage and swimming toward them (with the exception of the killer whale). MAN AND THE SEA, II 65 Anything the diver does that resembles the normal prey oF these predators might cause them to bite. For example, blood in the water stimulates sharks into a nervous frenzy called the "feeding mood" so that they will bite at any- thing in and around the area scented with blood. The smell of blood leads them to the prev, and when the victim or any object comes into range of their poor evesight thev will almost always bite. This is a normal feeding pattern of most sharks, which have a highly developed olfactory sense. For that reason, carrying dead or wounded fish on a belt while swimming can also be dangerous. Many fishes feed mainly by sight, and those dangerous to man are usually the faster-moving fishes such as the barracuda. They usually hunt down their prey with quick, darting strikes. Any splashing about the surface, especially in water with poor visibility, may resemble a school of frightened fish or a wounded fish, causing barracudas or other fishes to strike blindly. This is almost like a reflex with them (Chapter 2). Such fishes as wahoo or tarpon might possibly become dangerous in the same fashion. Bright, shiny objects such as rings or belt buckles look like the bright side of some fish and attract predators. When fishes are hurt they often show their light undersides. Having objects or clothing that flutter as the diver moves through the water can be dangerous for the same reason. Diving at night or in water with poor visibility can be extremely dangerous because many fishes such as sharks and fishes that would not normally attack man do so under these conditions (barracuda, Chapter 9, and sharks. Chapter 8). Here again, the diver enters the feeding pattern of fishes, for many fishes locate prey at night by picking up vibrations of swimming movements with their lateral line receptors (Chapter 2). These fishes pick up the vibrations of prey moving through the water and strike in the direction from which they are emitted. Some fishes have chemoreceptors on the skin of the body, and it is possible a touch by a diver will emit a biting response in the direction of the touch. Of course, almost all animals on land or sea when cornered or wounded will fight with every ounce of their strength. When a diver reaches into a hole for a lobster there is a chance that he will be bitten by a moray eel. The eel sees this move as an aggressive action and bites to defend itself. Toadfishes and wolf eels can also be provoked into biting in this way. The killer whale, being an intelligent mammal, must be viewed apart. It is particularly dangerous because of its liking for larger warm-blooded prey. Luckily, it is rarely seen out of cold northern waters. Almost nothing is known of its reaction to man under water. Aggressive animals dangerous to man: Some sharks; Large moray eels (when molested); Barracuda; Killer whale. Other large predatory animals which might be dangerous if man enters their feeding pattern: Sawfish; Tarpon; Spearfishes; Giant jewfishes; Wahoo; Some other large predators. 66 UNDERWATER GUIDE TO MARINE LIFE LARGE SPECIES The size and power of some marine animals should be a deterrent to handling them or being careless in their presence. Since the sea is an environment to which they, not the diver, are adapted, all animals of large size must be respected. The best defense upon meeting large animals under water is to let these animals see you as part of the environment and to move slowly and without alarm. Here, as in aggressive species, taking the psychological advantage and swimming toward them will usually drive them off. Possibly dangerous large animals: Large squids; Large octopuses; Whale shark; Basking shark; Manta ray; Other large sharks and rays; Tarpon; Tuna; Spearfishes; Large groupers; Jewfishes; Sea turtles; Seals; Sea lions; Porpoises; Whales. CHUMMING— ATTRACTING ANIMALS In most places, the majority of the animals of the sea are remarkably unafraid of the diver because, until recently, he has left them comparatively unmolested. In areas where spearing is done, fishes have become wary and difficult to approach. For purposes of photography and observation, chumming is extremely useful in attracting the wary and tame alike. Eventually, with periodic feeding, quite quickly in fact, fishes and other more intelligent animals learn to recognize their benefactors. It is a great thrill to be able to establish harmony with the animals of the sea by chumming them. Very little is known about chumming. New techniques, methods, and subjects should constantly be tried. It would be just as well not to try to chum sharks, manta rays, huge jewfishes, or barracuda by feeding methods, but almost all others, including sting rays, which have been known to be able to recognize their feeders in captivity, might be tried. Chumming of very small fishes such as wrasse, blennies, and gobies is very rewarding. Three general methods of chumming may be used: 1. Feeding. Good chum may be prepared from a mixture of sand, bread, and shrimp. In place of shrimp, various other sea foods may be used, but red meat is not successful because it bleaches out in water. Natural foods such as broken sea urchins, shellfish, and large crustaceans are also successful. The first step is to get fishes to eat in the presence of the diver— chum- ming the area. If the diver is to be associated with the food, it is important that the time between presentation of food and eating be as short as possible and also that presentation of food be made often. Eventually, the subject will be following the diver about, waiting for more offerings. Groupers are particularly good subjects of this method. Morays are some- what less "friendly" subjects, food placed near holes in reefs usually pro- ducing one or more of them. Great masses of smaller fishes such as wrasse never fail to accept food vigorously. 2. Sounds. This is a subject about which very little is known and about which critical experimentation is yet to be done. Mullets are known to be attracted by bells. Croakers and grunts might respond to an imitation MAN AND THE SEA, II 67 of the noises they make. Porpoises communicate by sounds under water and could probably be attracted by imitating them. Movements. Smooth, regular movements will attract very little attention under water, but jerky, splashy movements are known to scare most fishes away and to attract such large predators as barracudas and sharks. The clever use of body movements can serve to attract some fishes. The authors used retreating movements to attract barracudas for photographic purposes. The same have been known to elicit the curiosity of groupers. Small fishes such as blennies and demoiselles are often attracted by wiggling fingers. Movement is another area about which little is known. CHAPTER /^ EVOLUTION, NAMES, AND CLASSIFICATION As the swimmer explores the underwater world, he soon becomes aware of the diversity of shapes, forms, and habits of marine life. The bizarre shape of the sargassum fish, the sleek beauty of the tuna, and the plantlike form of gorgonians lead him to wonder what their relationships are, both to the environment and to each other. The adaptation of each of these to their environment depends upon the many small hereditary changes which, if advantageous, allow them to find food more easily, breed more successfully, or live longer. This adaptation involves evolution. EVOLUTION Figure 16 gives in "tree" form an outline of evolution of fishes for the last 400 million years, the period of time that their evolution is known through paleontological studies. No more than the barest summary of the evolutionary process from which the tree took root and grew can be given here. (The excellent books noted in the Bibliography, especially Simpson's Meaning of Evohition, should be consulted for further information.) The basic unit in evolution is a 'population, all members of which have free access to all other members without spatial, ecologic, social, or any other barriers. Take, for example, a hypothetical isolated coral reef, over and around which swim several individuals of a certain species of butterfly fish. These individuals compose a single population; all of them could breed freely together if they so chose. This population may be thought of as possessing a number of char- acters, hereditarily controlled by genes, in common. That is, the population represents a common gene pool. Because all members of the population breed tooether and share genetic features in common, thev would not be likelv to diff"er from each other significantlv, perhaps only varying slightly in size or intensity of coloration, etc.— differences due to the possession of slightly different assortments of genes from individual to individual. Butterfly fishes, for example, show variations which are due to different gene assortments. For instance, members of a single population may have eye stripes of varying widths and 68 EVOLUTION, NAMES, AND CLASSIFICATION 69 Fig. 16. The evolution of pshes. The stippled areas are approximately proportional in size to the numbers of species in each group. Notice that the more advanced groups tend to replace more primitive groups with time. Adapted from Romer (1945) and Young O950'). 70 UNDERWATER GUIDE TO MARINE LIFE lengths. However, no single individual can have all widths or all lengths of eye stripe. Instead, there are individuals with long, wide stripes, short, wide stripes, long, narrow stripes, and short, narrow stripes. A species may consist of one population, but it is usually composed of several populations which are more or less independent, but do have some gene flow between them. That is, some members of neighboring populations do breed, so that these populations also have genes in common, though the species gene pool is less restricted in scope than is the population gene pool. Within the species, gene flow may be restricted enough between some populations to allow them to differ recognizably from others. In such cases, these are called suhs-pecies. In humans the four races, Caucasoid, Negroid, Mongoloid, and Australoid, could be biologically classified as recognizable subspecies. Each has its own charac- teristics and, until recently, had its own geographical location. With increased gene flow between these races, the result in time will probably be a breakdown of race distinctions. Evolution would probably not progress very far, however, if it depended solely on differing assortments of genes that are already present. Somehow new build- ing blocks must be found, and these are produced by mutations. The nature of a mutation is only partially understood. Presumably, it is a structural change in a gene or even a rearrangement of genes on a chromosome which causes that gene to have a different effect than formerly. Most mutations are small changes. Drastic changes are rarely tolerated in nature, and in the great majority of cases of large changes due to mutation, the result is early death. Mutations are usually deleterious in effect, since mutations are purely random changes, not dictated by the environment or the needs of the organism. Purely random changes do not usually benefit an or- ganism which has evolved over a period of thousands of years in response to its environment. But, of course, beneficial mutations do occur occasionally. Mutations are rather common. Any one gene may mutate once in several thousand individuals. Evolution is the process by which the fate of these rather common, mostly small, sometimes beneficial mutations is determined. The assortment of genes also plays an important role since certain assortments of genes are more or less beneficial than others. The environment determines the fate of genes in evolution. Every gene or combination of genes can be assigned a selective value in its environment. If the gene is beneficial, the selective value is plus; if deleterious, the value is minus; if it has no effect, the value is zero. The environment determines this value since a gene, and an animal possessing genes, does not exist in a vacuum but in a definite place at a definite time. What may be beneficial in one place may not be so beneficial in another. By the process of natural selection, the beneficial genes are selected for and the deleterious ones selected against. Since the genetic change is not usuallv drastic, neither is the selection drastic. It usually expresses itself in a differential breeding rate between the individuals possessing beneficial and deleterious genes. Those individuals possessing beneficial genes find adjustment to their environment easier and produce on an average more offspring than those possessing deleterious genes. This results in time in a marked shift in the whole population toward the beneficial change within an environment which is presumed to be stable. Environmental change usually results in a change in direction of evolution. EVOLUTION, NAMES, AND CLASSIFICATION 71 Let us return to the butterfly fish population on the isolated reef which was previously mentioned. Suppose that these fish were at one time plainly colored, small yellow fish without a vertical stripe through the large black eye. Suppose that one individual was the result of a mutation which caused a narrow stripe to appear through its eye. This stripe would be an advantage in camouflaging the otherwise very noticeable eye (Chapter 9), and the gene controlling the stripe could be said to have a plus selective value. It would aid this fish to avoid its predators a little better, give it a little more time to eat, and allow it a slightly longer life in which it could produce more off^spring, some or most of which would also have this camouflaging eye stripe. Eventually, stripeless in- dividuals would be rare or lacking. More mutations might also occur to strengthen this trend toward a stripe through the eye in butterfly fishes. Such a trend is, in fact, very evident in these fishes. But evolution cannot be concerned merely with the change within one population. Somehow there must arise. a situation whereby two types are pro- duced from one, a splitting by which new species, families, and higher categories may arise out of the old. This can only occur if somehow the population is broken into two parts which cannot interbreed with one another. Suppose one segment of the butterfly fish population was swept away or wandered away to another distant reef. There would now be two populations on two different reefs. These populations would have slightly different mutations, and the reefs would represent slightly different environments. The results of the evolution in these populations would thus be different, and, given time, two new species would evolve which, even though rejoined, might have become so different that breeding would not occur between them. With expansion of this process of splitting, new families, orders, and higher groups may come into being. It is not too difficult to imagine how, by this process, certain fishes in shallow waters became amphibians adapted to spend part of their lives on land by selection in favor of supporting limbs, how certain amphibians became reptiles adapted to spend all their lives on land by selection in favor of an impervious dry skin, how certain reptiles became mammals adapted for active life by selection in favor of warm-bloodedness, and how certain apes became men adapted for con- templative life by selection in favor of high-mindedness. In each case, some of the descendants of the groups which gave rise to higher categories remained behind, so we see that evolution is a process that creates new forms and builds on itself. The nineteenth-century biologist Thomas Huxley likened evolution in filling the available living space on earth to the filling of a barrel. First, the barrel may be filled with apples. Then pebbles may be put into the spaces between the apples. Sand fills the spaces between the pebbles and finally water may be poured in to fill the barrel. Simpson (1949) adds that the barrel itself expands as the new forms of life create new ways of life bv their verv presence. For instance, without birds, there would be no bird lice, without insects no insect-pollinated flowers, and without whales, no whale barnacles. Evolution is studied from three aspects chiefly. First, the history of evolution is given in fossil form— paleontology. Second, the mechanics of evolution are revealed by genetics. Third, the all-important environmental effect is examined by ecology, commonly known under the name of natural history. We have briefly explained the first two. This book mainly concerns ecology, and three 72 UNDERWATER GUIDE TO MARINE LIFE phenomena observed by ecologists and paleoecologists studying evolution are so instructive that they must be mentioned. First, there is the phenomenon of convergence. Unrelated animals living in similar environments or having similar habits are under similar environmental stress and frequendy come to look alike. Porpoises, for instance, are adapted for a swift-moving, fish-eating existence on the open sea. Before mammals became dominant, when the reptiles reigned, a group of reptiles called "ichthyosaurs" with porpoiselike habits were remarkably like porpoises in body form. The whales and whale sharks are remarkably alike and have similar habits. There are many eel-shaped fishes all adapted for secretive predacious lives, yet derived from widely different groups of fishes. These are all examples of convergence of body form in response to similar habits and habitat. The second phenomenon is faralleUsni, which differs from convergence but little. Again it concerns evolution of animals in response to similar conditions, but, unlike convergence, the groups were not widely divergent to start with and have merely followed parallel courses in evolution. The demoiselles and the butterfly fishes are very similar in habits, size, and form and were derived from fairly closely related suborders of fishes. There are several predacious species of bottom-dwelling fishes such as toadfish, scorpion fish, stargazers, etc., which have developed from closely related stock. The third is adaptive radiation. All groups of animals will tend to evolve in all ways that are open to them within the limits of their own anatomies. Thus the reptiles and mammals have both had swimming, running, flying species. The fishes have radiated into almost every conceivable niche from the surface to the deep sea, from plankton and plant-eaters to voracious carnivores, and from life on rocks and reefs to sandy beaches. Even smaller groups of fishes show their own radiation patterns. One has only to examine the varied mackerel- like fishes— swift predators, plankton feeders, eellike forms, etc.— to see radiation at work. Evolution is a dynamic, continuing process. It is going on today even in the human species. Rarely does it come to a halt in any group. When such a halt occurs, the result is usually extinction— especially when environmental change occurs— but occasionally a seemingly changeless "living fossil" is left behind: the oyster, the horseshoe crab, the sturgeon, Lingula. Evolution is a peculiar combi- nation of the randomness of mutation and the orientation of natural selection. It is natural selection imposed by the environment that keeps species within bounds. Without it, deleterious genes could accumulate, and the result would almost certainly be genetic pandemonium, which could only result in environ- mental maladjustment and extinction. Evolution is highly opportunistic. New situations are quickly met by organisms that seize new opportunities, but such meeting is largely due to the chance that the animals are present to meet these opportunities and the chance that mutations of plus selective value occur. Islands or isolated coral reefs, for in- stance, have notably depauperate faunas simply because many groups of animals have not found their way there. Evolution seems slow to humans who measure time in minutes, hours, and years. Geologic time is not so reckoned. To the human eye, the world is a static place, but this eye sees only one frame of a moving picture. We should EVOLUTION, NAMES, AND CLASSIFICATION 73 not forget that the human Hfe is too short to see evolution in action and that the human animal has been on earth only one thousandth of the time that life has existed here. The age we live in is, in fact, an extraordinarily dramatic one typified by the harsh climatic conditions of an ice age, by fluctuating currents and water levels of the sea, and by the geologic upheaval of whole great mountain ranges such as the Pacific Coast range. But our lives are too short for us to notice most of these changes. Evolution has no goal, but it does have method and a reason. The necessity of life to adapt to an ever-changing and complex environment is its reason. The sensitive man will contemplate and study these things as he swims among the wonders of the sea that lie about him. NAMES It is of utmost practical necessity that names be given to the species of marine life about us. For common everyday purposes, common everyday names are used. These are sometimes adequate. Mostly they are not. When you eat a sardine, do you spread a small herring or a pilchard or a Spanish sardine on your bread? These are three different fishes, all -called "sardine." Different languages have different words for the same fish; for example, "ronco" (Spanish) and "grunt." "Langouste" (French), "crayfish," and "spiny lobster" are all names referring to the same animal. Examples are virtually endless of the confusion, multiplicity, and overlap of common names that are given to marine animals and plants. Common names are given by just about every method imaginable. The margate fish was named for the sailors from the English port of Margate. The yellow grunt was named for its color. The pompano was named for its shape. Gill's mojarra was named after a man. No one knows how or why the ridley turtle got its name. Obviously, some way had to be found out of this imbroglio of terms. In the mid-eighteenth century, a Swedish scientist named Linnaeus founded the system of scientific nomenclature which we use today. There is no need to be con- founded or overimpressed with scientific names. They are only names, after all, which may be a little foreign to most of us but which offer us some stability in the confusion of names, especially when we are not familiar with the local terminology. CLASSIFICATION Linnaeus's system for naming animals is built around binomial nomenclature. That is, every species is given two names, a first, capitalized, generic name, and a second, uncapitalized, species name. The genus of the spotted moray is Gyninothorax; the species is moringa. The complete name is Gymnothorax moringa. The subspecies name, if any, adds a third name. The Linnaean system was originated in a time before the acceptance of the validity of evolution. Linnaeus himself was a special creationist. Nowadays his system has been adapted into an evolutionary scheme and supposedly reflects the tree of evolution as it is revealed to us by the study of fossils. But since the 74 UNDERWATER GUIDE TO MARINE LIFE fossil record is far from complete and since the interpretation of it is subject to human frailty and differences of opinion, there are several systems of classifi- cation which involve duplications and overlaps of the same type that occur in common names. Are the triggerfishes, trunkfishes, and puffers three separate sub- orders, or are they three parts of one suborder? Opinions differ and names are invented to express the differences. Luckily, there is little difference of opinion regarding families, and they can be given the strongest reliance. This is due to the fact that the families of animals and plants form fairly distinct groups. For some reason, the transitional species that unite families are short-lived and usually do not stay around long enough to give scientists a chance to examine them or their fossil remains. This is probably because families share the char- acteristics of an adaptive type which is established quickly in evolution, so quickly that there are few intermediates. Basically, the classificatory system of living things is built as follows: Kingdom: only two in number— animal and plant. Phylum: the largest subdivision, e.g., Chordata, including protovertebrates and vertebrates. Class: a major phylum subdivision, e.g., Osteichthyes, including all bony fishes. Order: a prominent assemblage of families, e.g., Apodes, including all true eels. Family: the basic "adaptive type" grouping, e.g., Muraenidae, including all moray eels. Genus: a distinct morphological tvpe; e.g., Gymnothorax, including an as- semblage of very similar species. S'pecies: the unit of interbreeding individuals, e.g., Gymnothorax moringa, the spotted moray eel. It will be noticed that there are seven subdivisions in this svstem. As we progress down from kingdom to species, the evolutionary relationship between comparable groups becomes closer and closer Cfig. 17^. Most often, however, seven subdivisions are not enough to express the numerous branchings of the evolutionary tree, so the prefixes "sub," "super," or "infra" mav be added to the seven primary divisions or new categories such as cohort or tribe may be added. The basic system between class and genus may thus be expanded: Cohort Superorder Order Suborder Infraorder Superfamily Family Subfamily Tribe Just as evolution is complex and increases in complexity as we learn more about it, so the classificatory system increases in complexity in an attempt to reflect evolutionary patterns. There is some effort to stabilize this system. This EVOLUTION, NAMES, AND CLASSIFICATION 75 takes the form of rules set up by an international committee on nomenclature (Schenk and McMasters, 1948). Family names end in "idae," subfamilies in "inae"; modern taxonomists end orders in "iformes," suborders in "oidei," and superfamilies in "oidea." In the following pages an attempt is made to use the simplest classification possible. Classification is no idle business. The whole point of it is to simplify, and the underwater swimmer will find it to his benefit to try to think in terms .of a classificatory system in which each species has its position in a family which, in turn, has its place in an evolutionary system. An effort should be made to have in mind a hypothetical family image which gives the species of the family their major characteristics. This book is built around the convenience of having such family images, and a "type" system has been used here (Introduction). Fig. 17. The relationship between the evolutionary tree and classification. SECTION TWO CHAPTER 6 PLANTS OF THE SEA With the notable exception of eelgrass, 7:ostera, and turtle grass, Thalassia, flowering plants, most of the marine flora is made up of algae. The word "algae" is a catchall for several very distinct groups. Their variation in size is immense, ranging from microscopic to 200 feet in length. The variation in color is no less striking, ranging from blue-greens and greens to reds, violets, and browns. All of them, however, possess the pigment chlorophyll, which is without doubt the single most important plant compound known. Without it, plants would not be able to utilize the sun's light energy to manufacture carbohydrates (and, by addition of minerals, proteins) from carbon dioxide and water. This process is known as "photosynthesis" and is responsible for all life on earth since it is at the very base of the food chain. Chlorophyll absorbs light at the blue and red ends of the spectrum and reflects green, the color it appears to our eyes. The blue and especially the red light that is absorbed provides the energy used in photosynthesis. Since sea water absorbs light difl^erentially, eliminating the reds and then the yellows until, at over 65 feet, very little but green and blue are left, the quality of light available at various depths influences the distribution of the variously colored algae. This subject will be considered as each group is discussed. The smallest algae as well as the most important in the sea's economv are the planktonic ones, which are single-celled, usually enclosed in a skeleton, and which float or swim near the ocean's surface in immense numbers. There they form the grass of the sea, a vast and nourishing pastureland. The larger algae are mostly confined to shore, shoal, rocky, and reef waters, where they are anchored to rocks, shells, wharves, or even other algae (in which case they are said to be "epiphytes"— epi, meaning "on," fhyte, meaning "plant." These tvpes are most dense on shores sheltered from beating surf and, for that reason, Long Island Sound and Monterey Bay have particularly dense and varied floras. The large algae are easily separated from all other plants bv their lack of true roots, stems, or leaves, although thev have basal holdfasts which look superficially like roots and an upright, usually branched structure that looks like a stem and bears flat, leaflike extensions. 76 PLANTS OF THE SEA 77 The one place where no photosynthetic plant can exist, of course, is in very dim light or darkness. Because photosynthesis is impossible in such places, one would not expect to find photosynthetic algae below a depth of about 200 feet. Algae become rare below 100 feet. These figures vary according to the nature of the water. One group of single-celled animals (Protozoa) is commonly referred to as being more closely related to plants than to animals. This is the zoologists' Flagellata or Mastigophora or the botanists' Euglenophyta. Flagellates are con- sidered by many scientists to be a perfect link between animals and plants since they achieve mobility by means of flagella as do many animals, yet are able to manufacture food by photosynthesis as are most plants. Obviously, according to evolutionary theory, before there were true animals or plants, there must have been some group that gave rise to both and thus possessed the potential attributes of both. Such a group is Flagellata, a group which is considered in the chapter on invertebrates simply because its members look and act more like animals than plants. Flagellates, like small one-celled true plants, are plank- tonic and form a basic food of the sea. Many algae have sexual stages, notablv the male sex cells, that closely resemble protozoan cells. Bacteria are very minute and simple plants. They are not true algae. Their very simple order of construction sets them aside from all other plants. They are well known as the causal agents of many diseases and are probably the most common of all living things. They will not be included here, but it must be remembered that they form a basic food for many of the small one-celled animals. Some of them are unique among living things in their ability to use inorganic compounds (nitrates, nitrites, and sulphur compounds) as a source of energy. This method involves no photosynthetic process and indicates the primitive nature of bacteria (deep sea food chains, Chapter 2). Many plants of the sea, like garden plants, are known only by their scientific names. Most of the genera described and figured here have several species widely distributed on both coasts. DIATOMS: Phylum Ghrysophyta— F/^wr^ 18 Diatoms are microscopic one-celled algae that live in immense numbers near the surface of the sea, being the most common planktonic plant. They are enclosed in a siliceous shell of great variation and beauty. The shells may be ,^3«Wj%_ Fig. IS. Diatoms. The diatoms are shown much enlarged, the circular one being only about one-fiftieth of an inch in diameter natural size. Drawn from microphoto- graphs hy the authors. 78 UNDERWATER GUIDE TO MARINE LIFE rectangular, round, ovoid, or oddly club-shaped but are almost always marked by regular lines or pores, which may be, in some instances, so fine and regular that they are used to test the resolving power of the finest microscopes. The accumulated shells are deposited as diatomaceous earth, which has a number of extremely important commercial uses, such as insulating furnaces, preventing afterglow in matches, filtering oil and beer, acting as an abrasive in silver and car polishes, purifying water, and clarifying antibiotics. The accessory pigment fucoxanthin is present as in the brown algae, and its role will be discussed under that group. BLUE-GREEN ALGAE: Phylum Cyanophyta The dark, blue-green color of most of these most primitive algae is a com- bination of the blue of phycocyanin and the green of chlorophyll. Some of them also have a red pigment. Trichodesniiiini is one of these algae and is sometimes present in such quantities that it gives the sea a reddish color. The Red Sea is said to have been named because of the occasional presence of this alga. Blue- green algae are small, inconspicuous, slimy algae found most typically in fresh-water ponds as scum and in the sea adhering to mud, rocks, or wharves. Their small size frequently requires microscopic examination for identification. Mermaid's hair is found adhering to rocks, mud, and wharves of both coasts. Like other blue-green algae, it forms a dark, fuzzy, somewhat slimy mat on the surface to which it is attached. The pigment phycocyanin absorbs orange and red very efficiently and passes on the energy so acquired to chlorophyll, which uses it for photosvnthesis. Almost no blue light is absorbed by these algae so they must live in very shallow water, although Marshall (1954) reports the presence of blue-green algae in the deep sea. They must live on decaying organic matter, somewhat like fungi, there. GREEN ALGAE: Phylum Ghlorophyta— F/^wr^ 19 The vivid green color of these plants is due to chlorophyll, which identifies them. They are very numerous and are the most diversified and complex of algal groups, having 5,700 species the world over. They are especially familiar as pond scum, on rocks in streams, or on the bark of trees CProtococciis^ . In the sea, the green algae are mostly small and of temperate to tropical distribu- tion, being more varied to the south. Thev are not as conspicuous as red and brown algae. The marine green algae varv from flagellated, planktonic, micro- scopic, colonial, or single-celled forms, Dunaliella, to the small, branched, filamentous forms found attached to rocks in the intertidal zone, Clado-phora, and the showy, large sea lettuce, Lllva, which is usuallv found attached to rocks, other algae, or wharves just below the low tide mark. Sea lettuce reaches 3 to 4 feet in length and has a very wide distribution from the subarctic to the sub- tropics. It is fragile and is frequently broken by wave action and thrown up on beaches. Enteroiuorpha, sometimes called "grass," is a genus of manv species, most of which are very small. Some, however, reach 2 feet or more in length. The plant is either single or branched, and the blades are usuallv round and tubular and are always hollow. Most of the species are of the temperate zone. Sea moss, Bryopsis, is a lovelv, plumose plant which grows to 8 inches high. PLANTS OF THE SEA 79 merman's shaving BRUSH- P«-»»OS5 Ave. 'I i IRISH MOSS Chondrvs AqoLftlhteiUK CALCAREOUS ALCA Ccr&flino. +0 b in. Fig. 22. Re^ algae. Adapted from Smith C1944') and Taylor Cl937^. PLANTS OF THE SEA 85 Most red algae are small and filamentous or branched. For that reason they are frequently called "sea mosses." Many genera are much alike and frequently hard to identify. Red algae are most common in temperate to tropical seas. There are 2,500 species the world over. Laver, Porphyra, looks like sea lettuce and is frequently epiphytic— found growing on other plants. Like sea lettuce, it is fragile and washes up on beaches or into shallow water. It is found from the subarctic to the tropics and is orange or red to purple in color. It reaches 3 feet in length. Gymnogongrus is a beautiful and striking dark purple alga which looks like a small rockweed. It frequently grows among tropical coral but is also found in the temperate zone. It has a firm or even horny texture. Gigartina has a wide range of habitats from quiet sandy shores to rugged rocks where it must withstand the pounding of a heavy surf. It may be olive to purple or rose-red in color, and the surface is roughened by little outgrowths. The texture is leathery, and the plant may be branched, single, or spatulate in shape. Polysiphonia is very mossy in texture and finely branched. It is commonly only a couple of inches in length but may reach 3 feet. The color is a dark purple to brown or black, or even olivaceous. The many species are found from the arctic to the tropics of both coasts, and they grow anchored to rock, wharves, or other plants from the tidal zone to moderately deep water. The Irish moss, Chondrns, is very common, growing on rocks in shallow water. The various species are quite variable, but all are bushy and not over a foot tall, most being only a few inches. The color is a rose- to purple-red, or even with a green tinge, and the distribution temperate. It is a source of industrial chemicals. The dulse, Rhodymenia, gets into fairly deep water and is also found in the tidal zone. It looks like rockweed or Gymnogongrus in its branching and is a pretty, bright purplish red in color. It is of temperate distribution, keeping to deeper waters to the south of its range. Very often species of this genus have little bladelets at the tips of the main blades. Several species reach a length of a foot, but most are smaller. Agardhiella is a treelike, small, branched red alga which is very common on temperate coasts. It is frequently heavily burdened with crustaceans and other animals that live in its branches. Several red algae deposit calcium carbonate or lime around themselves. Some of these aid in the building of reefs. CoralUna ranges from the tropical to the temperate zones of both coasts. It is a dull purple or red in color and reaches a height of 6 inches. Bossea is a West Coast coralline alga, which grows on rocks in shallow water. It reaches a little over 6 inches but is mostly much smaller. SEED PLANTS: Phylum Tracheophyta The eelgrass, Zostera, and turtle grass, Thalassia (^g. 43^, and their close relatives are alone among the truly marine seed plants. They live in sandy, or muddy, quiet shallow waters. The leaves are slender and reach up to 4 feet in length. These plants send out long runners to propagate and bear small flowers and seeds. Many animals such as sea horses and green turtles depend upon these grasses for homes or food. It is unfortunate that because of pollution and parasitic infestation eelgrass has disappeared from much of its former territory. These plants range from the arctic to the tropics of both coasts. CHAPTER 7 THE INVERTEBRATE LEGIONS The diverse series of phyla that compose the invertebrates is so complex that the subject will be treated here on the level of the largest groups— classes and orders. This task in itself can be more complex than it would first appear to be, since 95 per cent of all of the species of animals are invertebrates. So huge is the invertebrate group, that no living man can say that he thoroughly comprehends even a majority of their vast array. Thus, the study of them is divided into a great number of subsciences— entomology for insects, protozoology for protozoans, conchology for molluscs, and so on. Nevertheless, comprehension of the types of invertebrates on at least a phylum and class level is possible, and we believe that such a treatment is vital in understanding the life of the sea. (More thorough treatments are available as listed in the Bibliography.) The study of invertebrates can be a fascinating thing. Because their ways of life are so far removed from the realm of man, one cannot help but be constantly struck by a sense of the bizarre. Most invertebrates possess very limited brain power and do not vary their habits as much as do vertebrates, but within the limited behavioral pattern of each lie constant surprises. The colors and the beauty of form and motion of many of them are often breath-taking, and their odd habits are constant sources of wonder. The invertebrates are united by the possession of no single characteristic, but rather by the lack of one. None of them has vertebrae or a backbone. Formerly, Aristotle separated invertebrates and vertebrates on the basis that vertebrates possessed blood and the invertebrates supposedlv lacked it. His word on this matter was accepted until only 150 years ago when invertebrates were shown to possess blood, though it is frequently colorless. In the following pages, the major habitat and ways of life of each group are given, together with brief descriptions of a few representative species. NONGELLULAR ANIMALS: Phylum Protozoa {first animal) A noncellular animal is one whose body is not di\ided into cells. Therefore, within the single cell of most protozoans, all the functions of digestion, 86 THE INVERTEBRATE LEGIONS 87 respiration, locomotion, and so on are carried out. It is true that some protozoans are colonial— several living together as a single group— but none of these has a division of labor among cells in which the cells are united into organs for special purposes. This phylum is the most variable one in either the animal or plant kingdom. No one body plan, form, or type of symmetry typifies all protozoans. Some species, like Amoeba, are even able to change their form as they move about, but most species have one form that typifies them throughout their lives. Protozoans are among the largest of phyla, existing practically everywhere— in sea water at all depths, in fresh water, on land, and broadcast through air as spores. Some are parasitic, causing such diseases as amoebic dysentery, malaria, and African sleeping sickness. Protozoans are all small. A microscope is required for seeing their details of structure, though many can be seen with the naked eye. They range in size from a few thousandths of a millimeter (1/25 of an inch is equivalent to 1 mm.) to an extreme of over ¥i of an inch for some fossil, amoebalike protozoans with skeletons. Protozoans reproduce by fission, a simple splitting of a cell in two, a method which allows them to increase their numbers rapidly when the right conditions prevail. Just when and where these conditions prevail is the key to understanding protozoans, for they are all extremely sensitive to environmental modifications. Under the proper conditions of salinitv, temperature, viscositv, pressure, light, food, etc., the numbers of a certain species may become incomprehensibly immense. If these conditions change only a little, the species will often die off and produce dormant stages, such as resistant spores which will become active again only when conditions once more become adequate. But so great are the numbers of species of protozoans that almost any conditions are adequate for at least one or two species. Rarely, if ever, are any waters free of them. Protozoan behavior never varies much within a species. Some species are light-sensitive, moving toward or away from it. All react to mechanical shock and temperature change, and most can sense food, moving toward it. Most of the protozoans with which we shall be concerned are planktonic and occupy a vital place at the very base of the food chain, either deriving their body-building energy directly from the sun or from bacteria or other minute particles which they eat. Flagellates: Class Flagellata or Mastigophora — Figure 23 This group, which forms a link between true animals and true plants (algae), occupies an extremely important place in evolution. Some of them contain chlorophyll and are photosynthetic, looking very much like some single-celled green algae (Dunaliella, fig. 19'). Others eat food such as bacteria or decayed substances as do animals. The photosynthetic species have a light-sensitive eve and move toward light. They are extremely important in the sea as basic converters of light energy to carbohydrates. All of the flagellates have one or several flagella, or whiplike filaments, which move the animal. Some flagellates are colonial with several cells living together as a unit, and these bear a remarkable similarity to the sponges in that there is a UNDERWATER GUIDE TO MARINE LIFE subordination of the single cell to the life of the whole colony, a prerequisite for a many-celled animal. This group forms the greatest of all linking groups in evolution. Not only does it unite animals with plants and protozoans with sponges, but the other groups of protozoans, as well as higher animals, were probably derived from flagellates. NootiltA.CQ. .. / - 5jllCeou.s CKry^omonacl Cryp+on^onad "x V Radiolaricxrx Fig. 23. Protozoans. Adapted from several authors. The chrysomonads form a group of predominately photosynthetic planktonic flagellates. They very often have a siliceous or calcareous skeleton and almost always have two large, conspicuous yellow or brown bodies (chloroplasts) within them which contain chlorophyll. The cryptomonads are oval-shaped and have a large gullet in which two flagella arise. Some are photosynthetic and others ingest organic matter through the gullet. None of these has a hard skeleton. The dinoflagellates are the largest and most numerous of the marine, plank- tonic, photosynthetic protozoans and are of enormous importance in the economy of the sea. The body is most often encased in plates made of cellulose, and there are two flagella, one of which rests in a groove which circles the body. Ceratinni is a common form in which long spines are developed as a flotation mechanism. The red tide organism, Gyimiodiniuvi hrevis, occurs sporadically in tremendous numbers along the coast of the Gulf of Mexico and in other warm seas. When it appears, sea water is turned into a muddv, reddish color and svrupv consistencv, causing a high mortality of marine life. If the temperature is high, it produces a vapor that is very irritating to eyes, nose, and throat. No one knows what causes sudden outbreaks of red tide. Gonyaulax causes similar damaging out- breaks of "red water" in Southern California. Noctiltica is luminescent and non- photosynthetic. It is large, to 1/25 of an inch in diameter, and when it is disturbed by a wave, a boat, or a swimmer, will emit a sudden light. The com- bined light of countless millions of them causes a beautiful, sparkling glow on the water's surface. The natural color of NoctiJiica is pale pink and bv day a pinkish tint may be given to the sea by them. All of the three groups of flagellates mentioned have some species which THE INVERTEBRATE LEGIONS 89 live in symbiotic relationship in the tissues of a wide variety of invertebrates, especially in corals, jellyfishes, and radiolarians. These species are known collectively as "zooxanthellae" and are small, degenerate, photosynthetic animals which live and multiply in the tissues of their host. They are able to live free of the host, but rarely do so. Zooxanthellae use the carbon dioxide, phosphorus, and nitrogen wastes of the hosts in order to synthesize food. They yield oxygen to the host and may even be digested by the host when the latter is hungry (Hyman, 1940). The brownish or yellowish color of corals and radiolarians is due to the presence of zooxanthellae. Some coral animals are greenish, and this is due to the presence of zoochlorellae, small, single-celled green algae that live in the hosts' tissues in the same way as do zooxanthellae. Amoebalike Protozoans: Class Sarcodina or Rhizopoda — Figure 23 Among the most delicate and beautiful objects of the sea as well as the largest protozoans belong to this group. All of the species have extensions of their bodies called "pseudopodia " ("false feet") by which they move and capture food. Pseudopodia are neither constant in form nor number, but are best thought of as being tentaclelike extensions which may appear or disappear at any place on the body. How pseudopodia are formed by the protozoan is not known. The foraminiferans ("hole bearers") are a group of amoebalike protozoans that build calcareous skeletons which are perforated by many minute holes through which the slender pseudopodia extend. The animal starts life with a small shell of one chamber. New chambers of ever-increasing size are added, usually in a spiral fashion (like the growth of snails) until the time of death. Foraminiferans are found in immense numbers from surface waters to the abyssal zone. Their skeletons have accumulated in huge quantities in oozes on the sea bottom. One form, Glohigerina, has covered 30 per cent of the forty million square miles of ocean floor with its shells (Buchsbaum, 1948). The chalk of England's Dover cliffs, a thousand feet thick, is composed largely of foraminiferan shells. If one considers that upwards of fifty million shells may occur in one pound of bottom ooze, the number of shells on the sea bottom becomes inconceivably great. Radiolarians are like foraminiferans, but have siliceous skeletons. They have yellow-brown zooxanthellae in them, which probably contribute to their nutrition. The delicate and frequently very beautiful skeletons, as well as oil or fat droplets in the body, and the fine pseudopodia aid these animals in floating at the surface of the ocean. In the very deep parts of the ocean (over 15,000 feet), radiolarian oozes are formed of the skeletons of these animals. Somewhat less than 5 per cent of the ocean floor is carpeted by this ooze. At these depths, the calcareous skeletons of foraminiferans dissolve, leaving onlv the relatively insoluble siliceous skeletons of radiolarians. Giliates: Class Ciliata — Figure 23 Ciliates are named and distinguished from all other protozoans by the possession of little hairlike processes called "cilia" which cover their bodies. Most are free-swimming, but some live attached to substrate or are colonial. 90 UNDERWATER GUIDE TO MARINE LIFE There are a "reat number of marine species, none of which has a skeleton. They are not common as oceanic plankton, but they are very numerous inshore in shallow water or as parasites. The species of ciliates live wholly by hunting down prey or by filtering sea water by means of currents set up around their "mouths" by the cilia. Because of these active modes of feeding, ciliates are the most active of protozoans and show more response to environment than do other protozoans. One group of ciliate derivatives called "suctorians" has lost cilia and has acquired long tentacles by which prey is captured and sucked dry. Prey usually consists of other protozoans or small larvae of invertebrates and may be several times the size of its captor. Suctorians are very common attached to substrate or to other animals in shallow waters. SPONGES: Phylum Porifera {"pore-bearer") Colonial animals are those in which several individuals have banded together to form one functional unit. Multicellular animals are those in which the different cells have become associated into different organs for the performance of different jobs. Sponges sit on the fence between being colonial and multi- cellular. They probably arose from flagellate protozoans because they are composed of aggregates of flagellated cells, but these cells are not gathered together to form organs. There are several types of cells, some wandering and amoebalike, and there is a supporting skeleton. Therefore, sponges, though they are not on the direct evolutionary line toward multicellular animals, do show how multicellular animals might have arisen through associations or colonies of single cells. The partial independence of the cells of sponges is demonstrated by the sponges' powers of regeneration. Sponges are a bit like the magical broom of "The Sorcerer's Apprentice." If a sponge is cut into many pieces, each piece is capable of producing a new sponge. It is even possible to separate the cells of a sponge by squeezing it through a fine cloth. These cells, if they are still living, will group together again and grow into a new sponge. Sponges are very plantlike and, in fact, were not known to be true animals until the nineteenth century. Basically, a sponge is a hollow vase perforated by holes (hence the name "pore-bearer"). There are many small holes by which water enters the vase and one larger hole, or a few larger holes, bv which water leaves. The latter are large enough to be easily seen by the naked eye. Currents of water through the vase are caused by the beating of the flagella of the cells on the inside and bring microscopic, planktonic food to the cells of the sponge. In most cases, sponges are not merely simple vases, however, but have become much infolded and complicated in form, so that the internal surface area of the sponge is greatly increased without involving much of an increase in size. Sponges have skeletons composed of calcium carbonate, silica compounds, or spongin fibers. The first two are brittle and in the form of delicate, ncedlelikc, branching spicules which are enmeshed together to form the sponge's supporting structure. Spongin is resilient and is the substance known to us all as the fibers of the bath sponge, Eiispongia. It is largely on the basis of their skeletons that sponges are classified. THE INVERTEBRATE LEGIONS 91 Sponges exist in a wide variety of shapes. Some are small and even parasitic. Others, such as the Pacific Neptune's cup, Poterion, are big enough to sit in. In general, sponges are like plants in growth pattern. For instance, a tree has a pattern of growth by which it may be recognized, but there is no set number of branches it must have. The same is true of sponges in that there are branching, erect, encrusting, or vaselike patterns of form, but each species shows consider- able variation within that form. The great majority of sponges are marine, there being only one small fresh-water group. Sponges are largest and most varied in shallow, tropical waters. Almost everywhere that they are found, a host of other animals, mostly worms and crustaceans, are associated with them, living in the cavities for protection. The most notable of the sponge symbiotic relationships exist between several species of hermit crabs and a few species of a family of horny sponges. The sponge grows on the shell occupied by a crab and eventually grows to cover the entire shell. Then the sponge dissolves the shell away and itself forms the home of the hermit crab. This is a case of mutualism in which the crab is protected from predators by the bad taste of the sponge and the sponge benefits by receiving transportation. Although most sponges taste and smell badly and are protected by their sharp spicules, two groups of animals, the nudibranch molluscs or sea slugs and the sea spiders, have some species that eat sponges. The sea slugs frequently imitate sponges in shape and color. The sea spiders suck the juices of sponges. Calcareous Sponges : Class Calcarea — Figure 24 These are the simplest sponges, small and vaselike, either single or grouped in colonies, yellowish to whitish or muddy in color, and with spicules of lime. VENUS FLOWER BASKET Euplec+ella-lOin. BREAD-CRUMB SPOMGE Halichror,dr,o.-(nficih+4in- BATH SPONGE Eu-sponc^ia- 8in. Fig. 24. Sfonges. Drawn from life. 92 UNDERWATER GUIDE TO MARINE LIFE Almost all of these sponges are found in shallow temperate waters on pilings or rocks or in tidepools. Leucoselenia is one of the most common of the colonial or branched forms, and Grantia is an "urn sponge." Both of these rarely reach over an inch or two in height. The distribution of these sponges is world-wide. Fig. 25. Top. This colonial vase sponge, three inches high, has large holes called excurrent openings through which water, taken in through many minute holes in the sides, leaves the animal. Bottotn. The basket sponge, Hircina, anchored on a small coral outcrop and almost large enough to sit in, is the largest species of sponge in North American waters. THE INVERTEBRATE LEGIONS 93 Glass Sponges: Class Hexactinellida — Figure 24 These are mostly vaselike sponges of deep water. The skeleton is formed entirely of six-rayed spicules of silicon compounds. Since most of them must anchor in the mud of deep water, there is commonly a rooting tuft of elongated spicules at the base of the sponge. The skeletons are frequently extremely beautiful glassy objects like that of the Venus flower basket, Eii-plectella. This group is found in deep water of all seas but is most common in the tropics. Horny Sponges: Class Demospongiae — Figure 24 The great majority of sponges belong here, and most sponges encountered by the diver will be of this class. The skeleton is largely formed of spongin fibers, which gives a breadlike texture to most of them. Some of them, however, have siliceous spicules (never six-rayed as- in glass sponges), and a few have no spongin but only have siliceous spicules. Some of the species have echinating spicules, that is, the spicules are sharp and break off in whatever object they touch. The fire sponge, Tedania, and the poison-bun sponge, FihiiUa, are two of these. The fire sponge is shaped like the red sponge Ccolor photograph'), but it is usually a much more brilliant reddish orange in color. The poison-bun sponge is breadlike in texture, brownish in color, and grows in rounded clumps like many common sponges. Both of these are West Indian and capable of producing more or less severe burning stings. Because of the irregular shape of many sponges, it is hard to recognize some of the species, so it is a good precaution to wear heavy gloves when handling them. The colors of horny sponges vary widely from the yellow bread-crumb sponge to the brilliant red of the fire sponge and the red-beard sponge, the purplish red of the red sponge, the green of some boring sponges and vase sponges, and the drab brown or black of bath sponges and basket sponges. The form of horny sponges is even more variable. The bread-crumb sponge, Halichondria, and boring sponges, Cliona, which can bore into the shells of molluscs or into rocks, are encrusting; the red-beard, Microciona, and the finger sponge, Chalina, are erect and branching; many are like vases and are colonial (/ig. 25 and color ■photograph); the bath sponges, Euspongia, are rounded; the basket sponge, Hircina (/xg. 25) is cup-shaped and almost large enough to sit in. Almost all of these types have extremely wide distribution, but horny sponges are most varied and are largest in the tropics. The area of the Bahama Islands is especially rich with them. POLYPS AND MEDUSAS: Phylum Coelenterata These are the jelly animals— jellyfishes, hydrozoans, gorgonians, corals, anemones— so named because of the presence of a jellvlike substance, consisting of about 98 per cent water, which is found between their outer (ectoderm) and inner (endoderm) layer of cells. These are the lowest members of the animal kingdom which have cells organized into organs— digestive, muscular, nervous, reproductive— and they also show, for the first time, a nervous system. This is a very primitive system with no brain and consists only of a network of nerve 94 UNDERWATER GUIDE TO MARINE LIFE fibers, but it does allow coordinated muscular action so that these animals can perform simple coordinated movement, such as contraction of the animal as a whole or movement of tentacles. Basically, there are two types of body form in this group, the sessile polyp and the free-swimming medusa. Actually, these two are merelv two forms of the same thing, the polyp being long of body with the mouth up and the medusa being short and turned over with the mouth down (fig. 26). The primitive species have both of these forms present in an alternation of generations, a name given for the extraordinary breeding method in which the polyp reproduces asexually by budding to produce medusas and the male and female medusas reproduce sexually by union of egg and sperm to produce more polyps. The whole phylum is divided into classes on the basis of which one of these generations, the polyp or the medusa stage, has become dominant. Because medusas are active and free-swimming, they show greater responsiveness to environment than do polyps. All of the species of coelenterates are united by the possession of two charac- teristic features. First, they are radially symmetrical; in other words, the top of the body differs from the bottom, but all the sides are the same. Second, most have unique stinging structures called "nematocysts" (_fig. 26), which, in response to a touch on a trigger (cnidocil), are able to shoot out a piercing barbed thread, injecting poison into the object with which it comes into contact. The poisonous thread acts like a fish line, holding the prey to the tentacle. One nematocyst by itself would not be very potent in either poisoning or holding prey, but as the prey struggles, it becomes more and more enmeshed in nematocysts and is subdued shortly. Some nematocysts are large and powerful enough to cause serious poisoning to man, particularly those of the Portuguese man-of-war and the lion's-mane jellvfish. These cells are the organs of offense and defense of coelenterates. Coelenterates are found in great numbers all over the world in shallow to deep waters. Some eat fishes, some feed by filtering plankton, and some have food manufactured for them by other organisms living in them (symbiotic zooxanthellae or zoochlorellae). Some of them are verv striking because of the beauty of their skeletons, as in corals, or the beauty of the iridescent blues, yellows, and reds of their bodies, as in anemones and jellyfishes. As unattractive as jellyfishes may be lying dead on a beach, in the water they are very beautiful as they move by gentle, rhythmic pulsation of their translucent bells, their tentacles streaming gracefully behind. But the diver should not be too carried away by the beauties he finds in this phylum. While most are harmless, many can produce painful stings. Hydrozoans: Class Hydrozoa — Figure 27 This is a group of great variation, having species that show alternation of generations between polyp and medusa stages and those that are exclusively polyps or medusas with no alternation of generations. There are twenty-seven hundred species of hydrozoans in all waters of the world. The hydroids are the most typical of the hydrozoans, showing typical alternation of generations. Most of them arc small, colonial, plantlike, branching, and have a skeleton of a stiff, horny material (though some are without THE INVERTEBRATE LEGIONS 95 budd barbed, poisonous +Kr£Q.d eicloderm POLYP (asexual) MEDUSA (sexual) AFTER DISCHARGE Fig. 26. The alternation of generations of a typical hydrozoayi coelenterate is shown on the left. The asextial polyp and sexual medusa exhibit the same general hody plan except that the medusa is greatly shortened, inverted, and has much more jelly than the polyp. Coelenterate nematocysts, on the right, fire a poisonoiis thread into prey or enemies on contact. skeleton). These are among the most common of seashore animals, but they usually go unnoticed because of their small size. Among the many genera of small hydroids is Ohelia, which shows, among its branches, two kinds of polyp animals, a feeding one with tentacles and a reproductive one which produces the sexual medusas by budding. At some times of year these little hydroid medusas are very plentiful. There are many species of small hydroids like Ohelia. One of these is Hydractinia, which is commonly found on the shells of hermit crabs (fig. 38^, a case of mutualism in which the crab receives protection from enemies because of the presence of the stinging hydroid tentacles and Hydractinia gets free transportation. Most hydroids are too small to sting man, but a few have very large nematocysts which are painful to touch. Phillips and Brady (1953) list Lytocarpus of the eastern coast of Florida and southward as one of these. This is a small colonial species like Ohelia and grows on rocks and pilings among seaweeds. Treatment for the sting consists of thorough washing with soap, followed by application of ammonia or calamine lotion to soothe the burn. Tuhularia, of temperate waters, is a beautiful reddish-pink hydroid that grows unbranched out of a mat and reaches a height of almost 6 inches. The most common small hydroid of the West Indies is Pennaria Cfig. 29), which looks like a small Christmas tree. Not all hydroids are small, however. One gigantic, noncolonial polyp, Branchiocerianthiis, reaches 6 feet in length and lives in the ocean depths. The stinging coral, or fire coral, Millepora Qfig. 28 and fig. 144^, is not a true coral, but is a hydrozoan related to hydroids. It grows in a great variety of forms— branching, erect and bladelike, massive, or encrusting (often on sea fans). They have dense, limy skeletons with very small pores through which the polyps project. Stinging corals are major contributors to the formation of reefs in all tropical seas. The sting of the West Indian species is only moderately painful and the effects usually wear off in a few hours. Indo-Pacific fire coral can produce more painful stings. 96 UNDERWATER GUIDE TO MARINE LIFE HYDROZOAN JELLYFISH GonionemiJ-S - ■§■ in. STAR CORAL As4ran<^.a-3,n. SETA FETATMLR liPennatula- fo i. Fig. 27. Polyps and medusas. Adapted from Mayer (7970) and drawn from life. See Color Plate 10. THE INVERTEBRATE LEGIONS 97 Trachyline hydrozoans are all medusas and look like true jellyfish. (One species, Geryonia, is shown in Color Plate 10.) They have no polyp stage, contrary to most hydrozoans, and are small to moderate in size, not reaching more than 4 inches across the bell. They may be distinguished from true jellyfish by the presence of a shelf called the "velum," projecting inward from the lower bell margin Cfigs- 26 and 27). Fig. 28. Stinging coral, Millepora, takes on a number of forms including that of these foot-high hlades, hut it is most easily recognized hy its mustard-yellow color and knohhy textiire. In the background is a large hush gorgonian. 98 UNDERWATER GUIDE TO MARINE LIFE Fig. 29. Top. Pennaria, a form of colonial hydrozoan, is only an inch high and looks like a Christmas tree. Each of the white knobs is the head of a single animal. The tentacles are too slender to he seen. Bottom. Sea whip gorgonians, unlike most anthozoans, "bloom" by day to capture their planktonic food. Each animal is barely one-fourth of an inch in diameter. THE INVERTEBRATE LEGIONS 99 The last of the hydrozoan groups is that of the Portuguese man-of-war, order Siphonophora. These are not single animals as is commonly supposed, but are floating colonies consisting of several kinds of medusas and polyps living together. One medusa forms the bell or float of the colony. Other medusas form swimming bells, which move the colony through the water. The polyp members of the community contribute the feeding individuals, tasting or feeling individuals, stinging individuals with nematocysts, and reproductive individuals. Thus, a siphonophore is actually a complex colony of manv kinds of individuals all specialized for diff^erent functions. Not all kinds of individuals may be present in a colony. For instance, the Portuguese man-of-war, Physalia QColor Plate 10^, lacks medusoid swimming bells and depends solely on the wind for distribution. All of these colonial siphonophores are pelagic. Most are small and delicate, but the purple sail, Velella, and the Portuguese man-of-war are rather large. Of these, the latter is famous for the power of its large nematocysts. Bad stings can cause cramps, nausea, difficulty in breathing, or, very rarely, death. One of the authors received the stings of only two nematocysts in his heel. In very short time, the poison had traveled to the region of the groin, making walking painful. Bad stings should receive immediate medical attention. The man-of-war reaches a size of a foot across the float and with tentacles over 30 feet long. It is thus able to catch a fish the size of a mackerel. True Jellyfishes: Class Scyphozoa — Figure 27 This is a rather small, compact group of only two hundred species. No true jellyfish has a velum as do the hydrozoan jellyfishes. Most have polyp generations of small size, which produce medusas by a type of budding (strobilation) during the warm months. 1 hese are the common, large medusas and are found in all seas, particularly those of temperate and cold zones. They are common objects cast up on beaches. (A typical species, Pelagia, is shown in Color Plate 10.) The true jellyfish common in warm or tropical seas are the cubomedusans, which are named for the high, squarish or cuboidal shape of the bell. At each corner of the bell is a single tentacle or group of tentacles. Caryhdea is a common example. It is an inshore jellyfish, a strong swimmer, a fish-eater mainlv, and has a strong sting. Most cubomedusans are only 1 to 2 inches across, but some reach 10 inches. Their sting has given them the name of "sea wasps." The discomedusans are the typical jellyfishes which are most common in temperate and cold waters. They have round, hemispherical bells and are the largest of all coelenterates. One species, the lion's-mane jellyfish, Cyanea, reaches a diameter of 8 feet or more with tentacles 200 feet long. Huge shoals of this animal are common in arctic or boreal seas, but, luckily, large ones do not range far south. They possess a dangerous sting because of their large size. The blue jellyfish, Aiirelia, is the most common jellyfish of the Atlantic and Pacific. This genus reaches a foot across the bell and can produce an irritating sting. Pelagia and Dactylometra, the speckled jellyfish, are two very beautiful and delicate discomedusans of temperate to tropical waters. Both can produce painful stings. The former is only a couple of inches across, but the latter reaches up to a foot in width. The rhizostomes are jellyfishes without tentacles and are common in shallow 100 UNDERWATER GUIDE TO MARINE LIFE tropical waters. One in particular, Cassiopeia, is common in the West Indies. It has no true single mouth and cannot sting, but it possesses a much-divided and lobed digestive system which contains symbiotic algae. This jellyfish is often seen lying lazily upside down in shallow water, exposing the algae to sunlight. The algae, therefore, produce some of the food for this jellyfish by photo- synthesis. All rhizostomes have the many-lobed digestive system like Cassiofeia, but most have no algae in them and therefore must live by catching plankton or small fishes. The young of many fishes use jellyfishes for protection, escaping the tentacles by alertness. Sometimes, however, they fall prey to their jellyfish protectors. Gorgonians, Anemones, and Corals: Class Anthozoa — Figure 27 There is no medusa stage at all in this group. Instead, the polyp has become complex and either secretes a home around itself and is colonial or is large in size and solitary. There are over six thousand species of anthozoans found in all seas, but they are far more common and larger in tropical seas than elsewhere. The colors found in this group are often very beautiful and caused partially by pigment and partially by the presence of organisms, zooxanthellae and zoo- chlorellae, that live in the tissues of the polyp. Hard corals are most often brownish, but they may be green, blue, or, in the case of the Mediterranean precious coral. Cor allium, a gorgeous red. The sea pen is red and phosphorescent. Gorgonians are brownish. The browns, yellows, and greens are the colors caused by symbiotic animals and plants. Most coral skeletons are white like coral sand. All of the members of this group are carnivorous, but the size of the prey varies greatly, and in many, the oxygen supply is augmented by the presence of zoochlorellae or zooxanthellae. The small, colonial anthozoans eat planktonic food, and the largest polyps, the anemones, catch fish or whatever else they can trap in their stinging tentacles. All are sedentary, though anemones can creep very slowly on their bases, and must wait for food to come their way. They either feed by passively filtering sea water or by attracting fishes with their flowerlike body form and colors. Some anemones even have little fishes living symbiotically in the protection of their tentacles, and these may serve to attract other fishes near enough to be caught. In any case, food is not acquired actively by the tentacles as in the case of the octopus. A fish must bump into a tentacle through its own carelessness to be captured. Many of the anthozoans, notably the corals, are nocturnal. With the coming of dusk, the coral colony blooms as thousands of tiny polyps emerge from their calcareous homes. This seems to be somewhat a contradiction of the fact that light is needed by the zooxanthellae that produce oxygen for the coral, but enough light is probably able to reach the retracted polyp to allow photosynthesis to occur. Very few members of this group possess nematocysts large enough to do damage to humans. Only one large anemone of the Indo-Pacific is really dangerous. Others may produce a barely perceptible burning when touched. Corals, however, may inflict cuts that are slow to heal and become easily infected. This is known as "coral poisoning" and is not very prevalent in North America. Ergophine ointment is said to be of use in alleviating the irritation of coral cuts. THE INVERTEBRATE LEGIONS 101 Fig. 30. Top. Sea fans, Gorgonia, over a foot in diameter, often anchor to feden- cidate coral in fairly calm water. Several small corals may he seen to the left, includ- ing a hliie coral Qtof left^. Bottom. Eusmilia is one of the most heautiful corals. The colony is not large, hut each coral polyp is of good size, up to three-fourths of an inch in diameter. 102 UNDERWATER GUIDE TO MARINE LIFE Among the several groups of non-stony corals, the gorgonians are the most prominent. These are the bushy sea whips C/igs. 28, 29, 97 , 144^ and sea plumes C/xgs. 97, J48) and sea fans Gorgonia Qfig. 30~). Sea whips and sea plumes have a resilient skeleton composed almost entirely of a horny material called "gorgonin" and because of their toughness often form dense beds in places where wave action would break stiff, stony corals. The beautiful iridescent, purplish sea fans add a considerable amount of lime to the skeleton. Being less .resihent, they more often grow attached to stony corals in calmer waters. The precious red coral, Coralliiim, of the Mediterranean is a member of this group whose skeleton is composed almost entirely of lime of a bright red color. Fig. 3]. Top. Brain coral, Meandra. Female hlue-head wrasse often cluster around the tops of large corals like this very large and almost perfectly symmetrical specimen. Bottom. Staghorn coral, Acropora cervicornis, forms extensive thickets where m.any fishes make their homes. THE INVERTEBRATE LEGIONS 103 Gorgonians are secondary reef builders and are largely tropical in distribution. Several animals live in their bushy branches. Among these are the basket star, Gorgoncephahis, hydroids, oysters, and some others. Dead man's fingers, Alcyonium, is representative of a group of anthozoans called "soft corals." It is usuallv found in rather deep water, but it reaches up to the low-tide mark. It has a flabby, mushroomlike texture supported by calcareous spicules. Soft corals vary in shape from squat, spongy masses to erect, treelike, branching forms and are found in all seas, but they tend to keep to fairly deep water. The pennatulaceans include the very beautiful sea pansies, ReniUa, and sea feathers, Pennatula. Both of these have a long single polyp that forms the axis of the colony and which serves to anchor the colony in soft substratum. This axis can move the colony slowly or withdraw the colony into the bottom when it is disturbed. These are the most active of the anthozoans. On both sides of the axis, there are extensions that carry the small feeding polyps. Most species are beautiful shades of yellow, red, or purple and inhabit shallow, tropical, and subtropical waters of all seas. Many of them are luminescent and will give off light when disturbed. The anemones form a fairly compact group of anthozoans. They are devoid of a skeleton and are noncolonial and flowerlike. Their distribution is world-wide in all seas, and they are almost always found attached to hard, rocky, or coral reef bottoms, where they sit waiting for prey to blunder into their tentacles. The size range is from less than an inch to 2 feet in diameter. Anemones usually avoid bright light and can move slowly on their bases more or less like a snail. They like to attach themselves in crevices, with only the tentacles exposed. Their colors are often extremely beautiful, ranging from bright green to white or dark brown with delicate blues and reds. Anemones are able to contract greatly in size, and when they do so the tentacles are drawn inside, which makes them difficult to spot. (The common West Indian anemone is shown in a color photogra'ph.^ Metridiuin is the most common temperate Atlantic species. The west coast green anemone, Crihina, owes its color to the presence of zoochlorel- lae. Adainsia (/tg. 38) is a remarkable anemone that lives in a mutualistic relationship on the shells of hermit crabs, giving the crab protection by means of its nematocysts and even patching holes in the shell the crab occupies (Hyman, 1940). The anemone benefits by receiving food and transportation. Several other anemones live in similar mutualistic svmbiosis with crabs. The true or madreporite corals are like small anemones but are colonial and secrete calcium carbonate (lime) around their bodies for protection. (Their environmental requirements are discussed in Ghapter 2 under coral reefs.) Gorals are found north to temperate waters, but form reefs only in the tropics. Star coral, Astrangia, is one that is found north to Cape God. Gorals show a great variety in form. Some are small like star coral, some are massive like several kinds of stone coral Qcolor photograph'), and the brain coral, Meandra Qfig. 31 and color photograph), some are stump-shaped like pedunculate corals Cf^g. 30), and some are branching like the staghorn coral, Acropora cervicornis Cfig- 31), or the antler coral, Acropora palmata Ccolor photograph). There are many species of small decorative corals. One is Eusmilia (fig. 30), an extremely beautiful coral, which has large polyps even though the colonies are quite small. 104 UNDERWATER GUIDE TO MARINE LIFE Fig. 32. Comb jellies. Adapted from Mayer (I9i2). On the West Coast is found the sohtary coral Balenofhylla, which reaches a half-inch in diameter. COMB JELLIES: Phylum Gtenophora {"comb-bearer")— Figure 32 There are only a few species of these exceedingly beautiful animals, but these often occur in tremendous numbers and are common in all seas. Their transparent bodies shimmer with delicate colors of glassy, iridescent beauty and bear eight comblike rows of cilia which move the animal through the water and have given these animals their name. Ctenophores are exclusively pelagic, predatory animals. They catch planktonic prey or small fish by means of two long retractile tentacles that extend from the sides of the body and which bear sticky "lasso cells." Some have no tentacles and filter plankton from the sea. Almost all the species have very wide distribution. The little sea gooseberry, Pleurohranchia, is found in all the seas of the world. The sea walnut, Mnemiop- sis, is like the sea gooseberry, but it is larger and is luminescent. If a boat is driven through a swarm of them, the disturbance causes each one to emit a bright, sudden flash of light. The Venus girdle, CesUis, is very different from these, being elongate and like a ribbon. It reaches a length of 3 feet occasionally but is usually seen much smaller. FLATWORMS: Phylum Platyhelminthes— F/^wr^ 33 Flatworms are not common or prominent in the sea, but are very well known as parasites— the flukes and tapeworms. One group of small flukes, the schisto- somes or blood flukes, has a larva that normally infects sea birds. It is also able to burrow partially into human skin, where it forms an extremely itchy swelling known as "swimmer's itch" or "seabather's eruption." These larvae die soon after entering the swimmer's skin, so do not produce parasitic infection in man. Phillips and Brady (1953) suggest immediate fresh-water bathing plus applica- tion of dilute ammonia or antihistamines for relief of the itching. Not all flatworms are parasitic. The turbellarians are flattened, frequently leaflike worms that creep about among plants or over the bodies of marine animals such as crustaceans. They possess amazing powers of regeneration. If the head of a turbellarian is cut off, the headless body grows a new head, and THE INVERTEBRATE LEGIONS 105 the bodyless head grows a new body— result: two worms. If the head is cut longitudinally in two the result is the regeneration of two heads. Almost all free-living flatworms have two eyes on the head which are turned in, making the animal appear cross-eyed. These eyes do not form an image, but inform the worm of the presence of light or dark. In general, flatworms avoid light. They feed on organic matter of all sorts, which they locate by means of chemical reception. Flatworms either glide on their bellies by means of cilia or swim by body undulations. The acoel flatworms have no digestive system and range up to a half-inch in length. The red Polychoerus is found in the tidal zones of both coasts. Convohita is a green acoel that derives its color from zoochlorellae in its body. This animal may feed when young, but when it becomes adult, it lives entirely by digesting the zoochlorellae, which results in the death of the worm, an example of poorly balanced mutualism (Buchsbaum, 1948). Rhabdocoel flatworms have a straight intestine and are very common on muddy and sandy shores or seaweeds. The triclads have three-branched intestines, are of dark color, and are most common on stony bottoms of northern seas. Both rhabdocoels and triclads rarely reach a half-inch in length. Bdelloura is a triclad common on the gills of the king crab, Lhnidus. The polvclads have many-branched intestines and reach a length of a half-foot or more. The tropical ones are frequently very beautiful, matching their sur- roundings in color. Some polyclads are pelagic. They swim by means of wavelike motions of the sides of the body. All of the flatworm groups mentioned show tendencies toward commensalism, suggesting that the parasitism so common in other flatworm groups, such as flukes and tapeworms, evolved first from commensalism. The end product of parasitism is often mutualism which, however, is not present in flukes and tapeworms. proboscis wi+k s+yle+ V SCHISTOSOME LARVA POLYCLAD Pie ucloceros-tii. Rhabdocoel triclad Plo-qios-fortiarn-rrii Bdelloui-a --t in ARROW WORM PHORONID ^a<)i + U- Jj-.n Phoroni5-^i-i Fig. 33. Unsegmented wonns. Adapted from several authors and drawn from life. 106 UNDERWATER GUIDE TO MARINE LIFE RIBBON WORMS: Phylum Nemertea— F/^wr^ 33 These are the lowest members of the animal kingdom to have a circulatory system through which blood flows and a digestive system with a separate mouth and anus. Both of these characters are of immense importance, allowing animals to be more independent in their environment. The presence of blood, by which food and oxygen are distributed throughout the body, and an anus, which frees the mouth of the task of egesting undigested food, gives greater freedom of action to an animal by increasing general body efficiency. The seas contain 550 species of nemerteans, which range from an inch to 100 feet long. The long ones, however, are not much thicker than a piece of ordinary string. On the head end is a sheath bearing a long noselike proboscis. At the tip of the proboscis a sharp stylet which resembles the nematocysts of jellyfishes is usually present. The proboscis is shot out at prey hke an entangling lasso, and those nemerteans which have a stylet use it to poison the prey. Worms, fishes, or crustaceans may be captured and are swallowed whole. The proboscis may also be used for defense and for burrowing. Nemerteans are found all over the world from the arctic to tropic seas. The largest in North America is the flesh-colored Cerehratulns of the temperate east coast. It reaches a length of 20 feet and burrows in sandy or muddy shores by day, but it may swim about at night near the bottom. It is very common and is often dug up by clam diggers. Other nemerteans are as numerous as Cerehra- ttihis, and some are very beautifully colored with bright red or green stripes and bars. A few species are pelagic. ARROW WORMS: Phylum Chaetognatha ("bristle- jawed") — Figure 33 These are swift-swimming, predatory, planktonic worms which reach the small size of only about % of an inch and are extremely common in cold temperate and arctic waters. They capture small planktonic animals in their bristly, hooked jaws. There are about thirty-five species. Sagitta is the most common genus. Their evolutionary position is not known. ROTIFERS: Phylum Rotifera ("wheel-bearer")— Figure 34 These little animals are extremely interesting to watch, but unfortunately a microscope is needed since their size is about that of a large protozoan. Some rotifers have the most extraordinary shapes and are very common attached to objects in shallow water. Some are pelagic. MOSS ANIMALS: Phylum Bryozoa ("moss animal")— Figure 34 Some of these little plantlike animals are so similar to the hydroid coelen- terates that the two groups can only be distinguished by an expert with a microscope. As their name implies, they look mossy. They are colonial. Some are erect and branching, and others are flat and encrusting in form. All are enclosed in an external skeleton, which gix'es the colony support and may be THE INVERTEBRATE LEGIONS 107 ROT ITER Qrea+ly enlarged Bird's hcad bkyozoam 8u.<^u.la.- 2. in ENCRUSTING BRYOZOflN £Q.ch OLniroal - li^ in. Ffg. 34. Rotifer and hryozoans. Adapted from several authors. either hornv and flexible or heavily calcified and stiff. Bryozoans feed by filter- ing plankton out of sea water by means of a crown of tentacles called a "lophophore." The bird's head bryozoan, Biigula, is a common and very widespread form found in most waters of the world. It looks like a small brownish tuft of moss, is 2 to 3 inches high, and grows anchored to almost any hard bottom in shallow water. Some individuals of the colony are reduced in size and look very much like the head and beak of a bird. These are called "avicularia" and constantly move from side to side snapping at any object that lands on the colony, keeping the colony free of foreign matter. Living BuguJa seen under a microscope show these amusing little individuals very well. An amazing habit of manv moss animals is the formation of the brown bodv. Periodically, the tentacles and anterior digestive system of a single animal degenerate into a brown clump which is ejected through the anus. Then a new set of internal organs is regenerated. No reason for this is known. The brvozoa lack a circulatorv svstem, and their evolutionary position in the animal kingdom is not known. PHORONID WORMS: Phylum Phoronida— F/^wr^ 33 The genus Phoronis with a very few species comprises almost all of this phylum. These worms are small, usually about a half-inch in length, and are like moss animals in habits and appearance except that they are adapted to live in straight tubes in shallow mud or sand flats rather than attached to hard substrate. Also, they are not colonial, although they are frequently found grouped together. They have a tentacle tuft (lophophore) and filter plankton from sea water as do moss animals. While they are not common on the east coast, thev are fairly abundant on the west coast. SEGMENTED WORMS: Phylum Annelida The worms whose bodies are divided into segments are called "annelida" which means "ringed." In some of these worms, like the sandworm. Nereis, the rings, or segments, are obvious, but in others they are less so. This is a huge group of several thousands of species. Segmented worms are evolutionarilv 108 UNDERWATER GUIDE TO MARINE LIFE very important as the group which gave rise to the arthropods, the largest of all phyla, which include insects and crustaceans. This phylum is divided into three major classes— earthworms, leeches, and marine worms. Only the last of these will concern us, the marine worms or Polychaeta. Two other small classes, the echiurids and sipunculids, are tacked onto this phylum because they seem to have been derived from segmented worms. In habits, this group varies greatly, from errant (wandering), predaceous worms to tube-dwelling, sessile plankton-feeding types, but the important thing about segmented worms is that they have a well-developed nervous system and a segmented muscular system so that rather refined movements are possible. This is an important advance over the nonsegmented animals that we have dealt with so far. Sexaal +a.il PALOLO WORM Eu„,ce- 10 in. lft+rover+ «&, BUDDING, SEKUAL WORM Syllis-ea.ch PARCHMEMT WORM BRISTLE WORM Hermoc(ice-U+. Fig. 35. Segmented worms. Adapted from the Reports of H.M.S. Challenger Cl 873-76^ and drawn from. life. Marine Worms: Class Polychaeta — Figure 35 The marine worms are obviously segmented, and most of them are provided with appendages on each segment called "parapodia," which bear bristles. Parapodia represent the initial effort made by invertebrates to develop limbs. They function as limbs, as gills, or in food-getting. THE INVERTEBRATE LEGIONS 109 The species of this class are extremely common in all seas. They may be rather neatly divided into three groups on the basis of habits— errant or wander- ing species, burrowing species, and tube-dwelling species. 1. Errant Polvchaetes. These marine worms are the most common. They are the most active worms and are armed with a jaw-bearing pharynx which can extend and retract for the capture of living prey. Some of them can swim, but most are found secreted under stones, among weeds, and other- wise hidden from their many enemies which relish their soft bodies for food. The breeding habits of some errant polychaetes are amazing. Most famous is the palolo worm, Eunice, one species of which is found in the South Pacific and another in the West Indies. During the last quarter of the October-November moon in the South Pacific or the third quarter of the June-July moon in the West Indies, the bottom-living palolo breaks off its tail (Borradaile and Potts, 1935). The tail, ripe with either eggs or sperm, then swims to the surface of the water, where thick swarms of these reproductive tail ends occur. When eggs or sperm are released, the tail, now an empty bag, dies and sinks to the bottom. Meanwhile, the original head end of the worm on the bottom regenerates a new tail which becomes reproductively ripe the following year. Swarming of breeding individuals at the water's surface is common in errant polychaetes. It insures fertilization of a maximum number of eggs since many worms are grouped closely together. Some swarming worms emit a light so that the females and males can more easily find each other. Others have the remarkable ability to bud off new individuals from the posterior end of the bodv- Each of these individuals can breed but cannot feed since it has no mouth parts. Sometimes a long chain of budding worms may be found at the water's surface during the breeding season. Two of these budding types are Syllis and Autolytus, both of wide distribution and growing about an inch in length. The sandworm. Nereis, is perhaps the most familiar of errant polvchaetes. There are great numbers of worms found the world over that look like this worm. Some are well over a foot in length, and the larger ones can give a painful bite with their strong jaws. The sea mouse. Aphrodite, is a mud-dwelling, stout-bodied, very beautiful species with long, silky, iridescerit bristles. It reaches 4 inches in length. The scale worm, Lefidonotus, reaches 2 inches and is common under stones near shore. These are both stout worms of wide distribution which stay near the bottom. One family of errant polychaetes deserves special mention because of the stinging powers of its bristles. This is the Amphinomidae, or the silky bristle worms. There are several species, all of which have long, detachable bristles that easily penetrate skin and cause a burning sensation. These bristles are hard to remove. Phillips and Brady (1953) suggest removing the bristles with adhesive tape. The most common bristle worm is Henno- dice of tropical waters. It often reaches a foot in length and is warningly colored with red marks on the sides. The white bristles tipped with 110 UNDERWATER GUIDE TO MARINE LIFE Fig. 36. Tof. The arrow crab, Stenorhynchus, three inches long and named for its long nose, moves slowly and depends on its concealing shape for protection. Bottom. These strange mounds, over a foot in diameter, failed to reveal their builders when the authors dug into them. They are probably biiilt by burrowing polychaete worms. red are erected when the worm is annoyed. Eiirythoe is a west coast bristle worm of pink color and snow-white bristles. 2. Burrowing Polychaetes. These worms have reduced parapodia and head and frequently look very much like earthworms. Most of these worms THE INVERTEBRATE LEGIONS 111 travel in soft or loose bottom. The lugworm, Arenicola, is common in temperate seas and reaches about a half-foot or more in length. 3. Tube-dwelling Polychaetes. The beautiful fan worms of the family Sabellidae, plankton-feeding worms with a delicate crown of plumes at the head end CcoJor photooraph^, belong to this group. They are found in all seas but are largest in the tropics, where their flowerlike tentacles are often seen protruding from tubes in massive corals. When disturbed, they quicklv retract into their tubes. The tubes, which are secreted by these worms, may be hard and calcified, as they usuallv are when in exposed places, but some of these worms live in U-shaped, parchmentlike tubes in soft bottoms. One of these is the parchment worm, Chaetopterns, of temperate seas. It is among the most luminescent of animals and is extremelv delicate. It circulates water through its tube bv flapping delicate appendages in order to filter-feed on plankton. Sipunculids: Class Sipunciilida — Figure 35 These are large, sand- or mud-dwelling, burrowing worms of most seas. There is no segmentation to the muscular body. Thev live- in burrows and have at the head end a retractile, tentacle-bearing, muscular "introvert" with which they grasp and swallow mud and sand. Thev are, therefore, much like earth- worms in habits. Echiurids: Class Echiurida — Figure 35 These worms are very much like sipunculids, but instead of an introvert, they have a trunklike extension for feeding. Echmrius is a stout-bodied species which lives in a U-shaped tube in mud. The larvae of the European echiurid, BoiielUa, have an unusual method of establishing which sex they will be. The larvae are free-swimming. If they land on the trunk of an adult female, they develop into small, parasitic males on that female. If they do not land on a female, they grow into solitary females. The trunk of echiurids serves both for locomotion and for food-getting. Some species live in soft bottoms and some in clefts in rock, and they frequently move from place to place, not staying in one cleft or tube. They do not eat sand or mud, as do sipunculids, but eat animal and plant matter, probably mostly debris. ARTHROPODS: Phylum Arthropoda ("joint-legged") Four-fifths of all species of animals on earth are arthropods (insects, crusta- ceans, spiders, etc.). They literally abound everywhere, in every imaginable habitat, occupying more different kinds of habitats than any other phylum. For these reasons, arthropods are undoubtedly the most successful animals on earth. Arthropods are named for their many-jointed appendages. In the primitive species, there is a pair of appendages present on every segment of the body, but in the advanced species, these appendages look very different in different parts of the body, being modified into antennas, various mouth parts such as 112 UNDERWATER GUIDE TO MARINE LIFE mandibles, maxillae, or maxillipeds, claw-bearing "arms," walking legs, swim- merets, and tail "fins" on the abdomen. It is the character and number of the appendages that form the basis of classification of arthropods. The skeleton of arthropods is well developed and typically formed of a complex substance called "chitin," which sometimes is strongly calcified (crabs and lobsters). Unlike the skeletons of vertebrates, however, the skeleton of arthropods covers the body rather than forming an internal structure. Also unlike vertebrate skeletons, the external skeleton does not grow but must be periodically shed as the animal grows inside, the new skeleton always being a little larger than the old. The nervous system and the eyes of arthropods are well developed, allowing interesting and complex behavior. The eyes are compound, that is, are composed of several individual small eyes grouped together yet acting as a unit. They are especially well suited for detecting movement since an object produces an image first on one facet of the eye, then another in succession as it moves across the field of vision. For the first time in the review of invertebrate phyla, a separate respiratory system is present. This may take several forms— either gills, book "lungs," or tubelike tracheids— but whatever the form, it allows a more efficient respiration than that of previous phyla, which picked up their oxygen through absorption through the surface of the animal. This advance, in turn, allows larger size, higher metabolism, and increased activity. For all these reasons the arthropods are well-protected, active animals which have complex behavior. The crustaceans form the most numerous group of arthropods in the sea. Crustaceans: Class Crustacea— Figures 37 and 38 All crustaceans have three rather definite parts of the body— head, thorax or trunk, and abdomen. Sometimes the head and thorax are united into a single unit, as they are in crabs and lobsters, and covered by a single "shell" called the "carapace." Crustaceans have a larval stage called a "nauplius" Qfig. 3i7^, which is common in plankton and is often an extremely bizarre little animal. It is by means of larvae that many crustaceans are distributed through the seas. The primitive crustaceans are filter-feeders which sift plankton from sea water. More advanced species are carnivores or scavengers. In fact, the crabs and lobsters are the vultures of the sea, eating literally any organic material, living or dead, that they can lay a claw on. Some crustaceans have good powers of defense in their very strong claws. Others are protectively colored or shaped to mimic rocks or seaweeds. Still others can change color so as to match the lightness or darkness of the bottom. Many of the highly developed crustaceans have a remarkable defense involving intentional loss of a limb or autotomy. If a limb is seized bv an enemv, the crustacean snaps it off, then retreats, later growing another limb. This class occupies a vital place in the economy of the sea. Many of its species form the major food source of m.any animals. Notable among the many examples is the dependence of the baleen whales on the Euphausian decapods THE INVERTEBRATE LEGIONS IB called "krill." Man, when he speaks of possible future food from the sea in terms of "plankton cakes," usually means the small planktonic crustaceans which are extremely abundant in polar and boreal seas in the warm months. The several groups of crustaceans are briefly reviewed below. WATER FLEAS: Order Cladocera Water fleas are enclosed in a large, round, bivalve carapace, which does not, however, cover the head, and have two large antennas which are organs of locomotion. Daphnia is a fresh-water form commonly sold in pet shops as a living fish food. Some water fleas are verv oddly shaped, and all are small, scarcely the size of the head of a pin. Their distribution is world-wide. They are planktonic, very numerous, and an important food for plankton-eating fishes. OSTRACODS: Order Ostracoda Like water fleas, ostracods are planktonic, small, swim with the antennas, and are enclosed in a bivalve shell. But ostracods look just like little swimming clams. The study of ostracods is especially important to oil prospectors because certain fossil ostracods are found associated with oil deposits. NAUPLIUS LJ\RVA MENTIS SHRIMP 1 Scjuilla-l-ol ft. SHORe SLATER u,,,.- 1 ,„. Fig. 37. Lower crustaceans. Adapted from various authors including the Reports of H.M.S. Challenger Ql 873-7 6') and drawn from life. 114 UNDERWATER GUIDE TO MARINE LIFE SNAPPING SHRIMP Cro-nqon- i m PELAGIC SHRIMP Lencifer - 4- ■". PHYLLOJQIvifl LARVft . SPINY LOBSTER Pig. 38. Advanced crustaceans. Adapted from the Reports of H.M.S. Challenger Ql 873-76") and drawn from life. REV SNAPPER MARGATE SRUMT \. 1 "^^-^^^ COLOR PLATE % YELUOWTAIL DEMOISELLE REEF FISH BAN O EP BUTTERFLY FISH FOUR- EYED BUTTERFLY FISH COLOR PLATE 6 THE INVERTEBRATE LEGIONS 115 COPEPODS: Order Copepoda This is the third and last of the small, planktonic crustacean groups. Again, movement is accomplished by means of two large antennas. Usually there is a single large eye in the center of the head, which has given one fresh-water form the name of Cyclops, after the Homeric giant. The body is usuallv pear- shaped, and the females are often seen carrving two little sacs of eggs attached to their abdomens. Calaniis is one especiallv abundant, cosmopolitan copepod which is eaten by fishes and baleen whales. Some copepods are parasitic (fish lice). BARNACLES: Order Cirripedia Barnacles have been described as crustaceans that lie on their backs surrounded by a shell, kicking food into their mouths with their feet. These "feet" are actually long appendages that form a sievelike net for the trapping of plankton as thev are rapidly thrust in and out of the shell in a grasping motion. Barnacles are the only sessile crustaceans and are hermaphroditic (both sexes in one animal). Hermaphroditism is present in many sessile animals, doubling the num- ber of voung produced as the result of one copulation. Sessile animals must produce high numbers of voung at a breeding because they can not move from place to place and are thus restricted in acquiring mates. In the case of barnacles the usefulness of hermaphroditism is particularly well illustrated. Unlike many sessile animals which merely shed their eggs and sperm into the sea, barnacles have internal fertilization in which the very long penis of one barnacle deposits sperm directly in the body of a neighboring barnacle. But the barnacle is limited in its mating by the length of the penis, and isolated barnacles out of reach of other barnacles can not mate, although it is possible that these indi- viduals fertilize themselves. The barnacle thus compensates for being sessile by doubling up, each individual being both fertilizer and fertilized, with twice the number of larvae being produced. The goose barnacles, Lepas, attach themselves to substratum by means of a long, muscular stalk. Goose barnacles most often attach to floating objects in the sea, the bottom of ships or to seaweeds. One species attaches to jellyfishes. The acorn barnacle, Balanus, is an extremely common animal of the tidal zones of all oceans. Barnacles frequently attach to living animals such as molluscs, crabs, or lobsters, and one barnacle, the whale barnacle, lives exclusively on whales. Barnacles do not harm the animals on which they attach except by slightly hindering movement, but some specialized barnacles have become parasitic. The most remarkable of these is SacctiUna (^g. 37^, a parasite on crabs. The Sacczdina lar\'a bores through the shell of a crab and pours its cells into the blood of the crab. These cells conglomerate on the underside of the intestine, and then the whole cell mass grows into a rootlike system which permeates the crab's whole body. The part of the Sacculina containing the reproductive organs protrudes from the underside of the crab's abdomen. SacciiUna lives on the body of the crab, but it does not kill the crab, only causing parasitic castration. Eventually, the Sacculina drops off, and the crab regenerates its sex organs. 116 UNDERWATER GUIDE TO MARINE LIFE MALACOSTRACANS: Order Malacostraca The rest of the crustaceans, including crabs, lobsters, sand fleas, shrimp, prawns and others, are included here. In all of them, the appendages on difi^er- ent parts of the body are sharply marked off from each other. The largest appendages are those of the trunk, which are used for locomotion and food- getting and very often bear claws. There are almost always stalked eyes on the head. A carapace covers the head and at least part of the trunk. All members of this group may be referred in general body plan to a central type, that of the shrimp or prawn, in which the body is adapted mainly for swimming. The species that depart from this general body plan do so as a result of a change of habits to burrowing or crawling, which causes a modification of the limbs on which heavy claws may be developed. ISJehalia, a little crustacean common all over the world on shallow rocky or weedy shores (especially near foul water), is the most primitive of this group. It reaches only a half-inch in length. The amphipods are elongated and compressed little crustaceans without a true carapace. These are the beach fleas or sand fleas, and there are great num- bers of them found literally all over the world, living either semiterrestrially on moist beaches or in shallow water. Gammarus is one common genus. The amphipods can swim, walk, and jump. They rarely reach an inch in length, but sometimes they grow larger. The coloration of amphipods often matches that of their background. Except for their movement, they are often difficult to see. Amphipods form a major item of diet for many fishes. One interesting amphipod, the skeleton shrimp, CapreUa, is about a half-inch long and is often seen clinging to seaweeds, especially AgardhieUa, or other suitable substrate by means of its small, posteriorly placed legs. It captures small prey with clawed legs near the mouth in a comical-looking, lurching motion of the whole body. Its acrobatics as it clambers over seaweeds are very amusing to watch. The isopods are small crustaceans that are depressed in body form. The terrestrial sow bugs and pill bugs are examples. Isopods are found on land, in fresh water, over almost all of the sea, and are even sometimes parasitic. Therefore, they qualify as the most widespread of all crustacean groups. Most species are small and found creeping over rocks, in seaweed, or over sand. The mantis shrimps, Squilla, are similar to isopods in being flattened, but they live in burrows in soft bottoms and bear large, raptorial claws, like those of the insect praying mantis, with which they capture prey. Some mantis shrimp reach the large size of almost a foot, but most species are under a third of that size. They are found all over the world. The rest of the malacostracans are grouped under Eucarida, which includes all the crustaceans in which the carapace covers the whole head and trunk and forms a single unit called the "cephalothorax." Krill, or euphausians, are pelagic, small eucaridans that occur in huge swarms in cold seas and form a basic item of the baleen whale's diet. The largest of the eucaridans are the decapods, or species with ten legs, which include the crabs, lobsters, shrimps, and prawn. Some decapods are armed with large claws and some are not. Among those that do not have claws is the sand bug, Hifpa, which lives a rough and tumble life in sandy shores, where it constantly has to battle the force of the breaking surf. The THE INVERTEBRATE LEGIONS 117 sand bug filters plankton brought to it by the waves by means of its feathery antennas. The spiny lobster, PamiUnis, has no claws, but can inflict cuts with the spines that cover its bodv and antennas. It makes a rasping noise when it is seized by rubbing a knob at the base of the antenna on a rough groove on the face below the eyes. Very few of the shrimps and prawns have claws. They eat plankton and smaller bits of organic matter. Very often, these animals travel in large schools, and most of them swim actively, whereas the crabs and lobsters walk on the bottom and swim very little. Perhaps the most famous shrimp is the snapping shrimp, Crangon, which possesses one huge claw. This claw is used to make a snapping or crackling noise that is well known to divers and which became famous during the last war when it was picked up on underwater sound devices. Schools of snapping shrimp are found in shallow, calm water the world over. The marine shrimp, Stenopus, is a very beautiful, clawed shrimp with a white bodv crossed with blue and red iridescent bands and long, filamentous antennas. One of the largest and probably the most famous of all crustaceans is the Atlantic lobster Homarus. It grows to edible size of 1 to 3 pounds in about five years, but lobsters may reach very old age, a weight of up to 35 pounds and 2 feet in length. Homarus is probably the finest-tasting lobster found anywhere. Some crabs have somewhat reduced the abdomen and must hide this fleshy, poorly protected part in a snail shell which is carried about like a house. These are the hermit crabs, which have some remarkable symbiotic relationships with sponges, hydroids, and anemones. The true crabs reduce the abdomen to a little tab that folds under the cephalothorax. They form a large and complex group found the world over. Some true crabs have even taken to terrestrial life in the tropics. Crabs are the scavengers of the sea and will eat any organism they can catch or find dead. Their habits are not as varied as might be supposed. They all have large claws which are used for defense and offense or for manipulating objects. The claws of some are to be a\'oided since they can quite easily crush a finger or a toe. Most crabs display their claws in a threatening gesture whenever they are approached. Some crabs are exceedingly well camouflaged and will not threaten with their claws since they depend on concealment for defense. Crabs are pugnacious and active animals. Some of the most amusing experi- ences under water can result from watching their antics, their battles with one another, and their rapid, sidewise scuttlings about for food. One crab that is often seen about coral reefs is the spiderlike arrow crab, Stenorhynchus (/ig. 36). This bizarre and gangling little fellow makes a humorous picture as it reaches out for food with its blue claws and brings the food to a mouth that is far under its long snout. Arachnids: Class Arachnida — Figure 39 This class is mainly a terrestrial one and includes the economically beneficial, but not well-loved spiders, centipedes, and millipedes. The rightfully disliked ticks are also arachnids. Arachnids were among the largest in size and most dominant of marine animals in the Cambrian Period from 400 to 500 million 118 UNDERWATER GUIDE TO MARINE LIFE SEA STRIDER. HQloborHes--§-i»i. HORSESHOE CRAB Limu.lixs- l&in. SEA SPIDE Nymphon-3in. Fig. 39. Arachnids and insects. Adapted from the Picports of H.M.S. Challenger (^1873—76^ and drawn from life. years ago, before the age of fishes. In Silurian times, 350 million years ago, some scorpionlike arachnids became the first animals to venture onto land. Now, marine arachnids are restricted to a few animals which are either hold- overs from the past or have very specialized habits. The body of arachnids is sharply divided into a cephalothorax (head-body) and an abdomen. There are no antennas and no true mouth parts that could be called jaws, the mouth parts always being of the sucking variety (except for the horseshoe crab). There are only two arachnid groups in the sea. The first of these contains the living fossil, the horseshoe crab, Limuhis, which has existed on earth virtually unchanged for almost 200 million years. It is found on sandy shores over much of the earth, especially in the temperate seas of North America and in the western Pacific. Horseshoe crabs are harmless, but the feet of swimmers have accidentally been pierced by the long tail spine. Very large horseshoe crabs may be over IV2 feet long. They crawl slowly over the bottom, grasping in- vertebrates with their clawed feet. Prey is "chewed" by means of the hard bases of the legs near the mouth. They burrow in soft bottoms and can swim weakly by paddling with their feet. Under the abdomen of the horseshoe crab are a series of leaflike plates. These are the book gills which furnish these animals with oxygen from the water. Commensal flatworms are often found on the leaves of these gills. The sea spiders or pycnogonids are among the most incredible animals found in nature. They are very often of red coloration since several species li\c in rather deep water. They range in size from less than an inch in leg span to over 4 or 5 inches. The body is small, and the abdomen almost nonexistent, but the legs are extremely long and contain branches of the stomach which ordinarily would be in the abdomen if the abdomen were large enough. The mouth parts are of a sucking type; sea spiders live by sucking the juices from THE INVERTEBRATE LEGIONS 119 such plantlike animals as anemones, sponges, hydroids, and sea squirts, in the same way that some insects such as plant lice (aphids) live bv sucking plant juices. Sea spiders are common in all seas, but they are so well camouflaged by their shape and remain so still that they are not very often seen. Insects : Class Insecta — Figure 39 Onlv one group of insects has invaded the sea, the water striders, familiar bugs which shuttle across calm waters of wooded ponds, lakes, or streams. Oceanic water striders, Halohates, are small and tropical in distribution. They are predacious bugs with sucking mouths. They never enter the water but live bv walking on it as a land animal walks on the ground. Just how these little insects have adapted to live on the open expanse of ocean where there is no shelter from waves and weather is not well known. They probablv feed on small flying insects such as midges that fall on the open sea from the air. Thev also may eat diatoms. MOLLUSCS: Phylum Mollusca The name "Mollusca" refers to the fact that these are soft-bodied animals. Molluscs are nonsegmented, but thev have a head with tentacles and usually with eyes. Thev usually move by crawling about on a foot, a broad flattened area on the ventral surface. In the case of bivalves (clams and oysters) and tooth shells, the foot acts as a digging organ and in the case of cephalopods (squids and octopuses), it is formed into tentacles. The mollusc's outer body covering is called the "mantle" and usually secretes a shell which protects the soft body of the inhabitant. The molluscs form a gigantic group with about eightv thousand or more species. However, this great arrav falls rather neatlv into five major groups, each of which forms a rather compact unit, both as to form and general habits. In general, molluscs are rather slow-moving, sluggish animals. The exception to this rule occurs in the octopuses and squids, in which the nervous system is among the best developed of all invertebrates. Chitons: Class Amphineura — Figure 40 These are the most primitive of molluscs. The bodv is shaped like an oval disc and always bears eight hard plates on the back for protection. There are no tentacles. Some rather rare or deep-water chitons are wormlike in shape, but the great majority adhere closely to the body plan described. They are all animals of shallow, rocky shores where they adhere with their muscular foot, often so tightly that they cannot be removed without breaking them. Chitons browse on algae, using their tongues, which bear hard rasping teeth called a "radula," to scrape their food from rocks. Bv day, chitons prefer to rest in shady crannies or dark places on the undersides of rocks, but they wander about freely at night. Chitons defend themselves by rolling into a ball like an armadillo or pill bug. Chitons are found in all waters in the world. Thev are mostly under 3 inches UNDERWATER GUIDE TO MARINE LIFE SPINDLE SHELL bubble: shcll jooth shCll Bu.lloL- 1 in. DentaiKMH-Sln. Fig. 40. Chitons, snails, and tooth shells. Drawn from life. See also Color Plate 10. THE INVERTEBRATE LEGIONS 121 in length, but the Pacific Cryptochiton reaches almost a foot. There are many species off the Pacific Coast of North America. Chitons are less numerous in the Atlantic. Snails: Class Gastropoda — Figure 40 This is the largest class of molluscs by far. The body is short and is usually covered by a single, spiral shell. Locomotion is accomplished by means of the flat, muscular foot upon which the animal glides. Some conchs use the foot to make short jumps rather than as a glider, and the small, planktonic pteropods have 2 winglike feet for swimming at the surface of the sea. All snails have tentacles bearing chemoreceptors and eyes. None of them move rapidly. The mouth has a tongue with a radula. The beautiful shells of snails have caught the attention of collectors. The most primitive of gastropods are the flattened limpets, keyhole limpets, and abalones. These are all animals of tidal or subtidal rocky shores. All scrape algae from rocks with their many-toothed radula or eat dead organic matter, and all are more active by night than by day. The limpets and keyhole limpets are the smallest, never reaching over 4 inches and averaging much smaller. The abalones, however, reach a foot in length and are delicious as food. They are found only in the Pacific and are treated as game animals protected by law on the Pacific Coast of the United States. The limpet, in spite of the primitive nature of its nervous system, shows a remarkable homing behavior. Each limpet has a definite place on a rock to which it adheres when it is uncovered at low tide. It wears a depression in the rock which conforms to the shape of its shell. When covered by water at high tide, the limpet moves away as much as three feet from its rock, "scar" home to browse, returning again at low tide. It finds its way back by remembering the topography of its home rock and always fits itself into the scar. The rachiglossans are snails that have only three large teeth in the radula. These are used to bore through the shells of clams and oysters or other molluscs or perhaps other animals. All of them are carnivorous. The well-known whelk (scungill), Biisycon, is a large species which lays eggs that look like a string of flattened beads. The oyster drills, Urosalpinx, are pests in oyster beds. Murex CColor Plate 10^ is a widespread tropical genus which has a shell that is dis- tinctly laminated and often very spiny at the places where the lamellae overlap. Nassa is a small species that eats other molluscs but also scavenges; it is very common on mud bottom. The taenioglossans make up the great majority of snails. They include carnivorous species such as the moon shell, Natica, which feeds on shellfish and lays eggs in sand collars, common objects cast up on beaches. The slipper shells, Crefidula, and periwinkles, Littorina, are common tidal or subtidal vegetarians of rocky shores. The queen conch, Strombus gigas QColor Plate 10^, forms a home for the little conchfish and is among the largest of snails, often weighing over 5 pounds. It progresses by jumping and is a scavenger. The cowries, Cyffaea CColor Plate JO), are among the most beautiful and most sought after of shells. They are most common in the tropics and not very abundant in North America. The helmet shells. Cassis, include the king conch 122 UNDERWATER GUIDE TO MARINE LIFE or king helmet CColor Plate JO). Helmets prey chiefly on bivalve molluscs and reach a size almost as large as that of the queen conch. The toxoglossans include the poisonous cone shells, Conns (^Color Plate 10^, which have the radula reduced to two long, sharp lancets attached to a poison gland. The cone shells are not known to be dangerous in North America, but some of the South Pacific species are deadly. They should not be handled. Treatment of the poisoning is just like that for snakebite (consisting of lancing the wound in order to encourage bleeding, followed by immediate attention by a physician). Cone shells can always be recognized by their conical shape. The remaining groups of snails all tend to reduce the shell. Pteropods are seldom more than Vi of an inch long, have transparent, papery shells, and are planktonic. Their foot is divided into two large "wings" for purposes of swimming. They are common enough in some places to form bottom oozes from their cast-off shells. The large sea slug or sea hare, Aplysia QcoJor photograph^, reaches almost the size of a football, feeds on algae, and ejects a harmless, inky fluid when it is molested. The smaller plumed sea slugs or nudibranchs CColor Plate W) are probably the most beautifully colored of all invertebrates. Their soft but iridescent colors and flowing movement are extremely beautiful. They grow to a maximum of about 4 inches. Most plumed sea slugs are of cold-water distribution, but some species are also found in the tropics. They are carni\'orous, and some are almost unique in that they eat sponges which are avoided by all other animals but sea spiders. Other species eat anemones or bite the heads off^ hydroids for food. (The hydroids do not seem to be bothered much by this since they quickly grow a new head.) The plumes on the backs of sea slugs are used as respiratory gills and have given these animals the name of "nudibranch" which means "naked gills." One nudibranch, Eolis CColor Plate lOX has extensions of its digestive tract reaching up into its dorsal plumes. The nematocysts of the coelenterates, which it eats, are stored in these exten- sions and are used by Eolis for defense— a completely unique use of the defensive mechanism of one animal by another. Eolis's plumes are brightly and warningly colored, and Eolis is avoided by fishes. Tooth Shells: Class Scaphopoda — Figure 40 These litde animals have a long hollow shell, a radula like the snails, and a burrowing foot like the clams. They never are more than about a half-foot long and are usually half that length. They burrow into sandy or muddy bottoms and feed on microscopic organisms. They are chiefly of cold or deep waters offshore. Bivalves: Class Pelecypoda — Figure 41 These are compressed animals with no head and with two shells or valves united by a hinge. The hinge is elastic, and when the shells are closed by muscles, the hinge is compressed. When the muscles are released, the hinge causes the shells to open. Thus, these animals must exert muscular force to keep their shells closed but exert no force to open them. A starfish is able to tire a bivalve merely by a continued pulling force. When the much stronger bivahe relaxes its muscles out of exhaustion, the starfish is able to eat it. THE INVERTEBRATE LEGIONS 123 PTEROPODS CKallenqer-'if in BOB-TAILED SQUID Roisia- 4 in. ARGONAUT ESe CASE -"t in. Fig. 41. Clams, peropods, cephalopods and hrachiopods. Adapted from the Reports of H.M.S. Challenger (1873-76) and McGoxvan (J 954) and draivn from life. 124 UNDERWATER GUIDE TO MARINE LIFE Bivalves are adapted mainly for protection. Their thick shells are so effective in this respect that this ancient group has evolved very little during the many million years of its existence. It is true that about all they do is feed and breed and secrete a shell. In general, they move only a little, their major response to disturbing stimuli being a closing of the shells. Almost all bivalves are filter-feeders. They have incurrent and excurrent siphons through which sea water enters and leaves the bodv. This water brings oxygen and microscopic, planktonic nutrients to the bivalve. The bivalves form a very compact group. Some, such as the mussel, Mytihis, are sessile and attach themselves to hard substrate by means of tough threads called a "byssus." The jingle shells, Anoniia, have fragile shells and are common in all seas at- tached to all sorts of solid objects in shallow water. The ark shells. Area, have heavy shells, and one species, the bloody clam. Area fexata, is the only mollusc with red blood. Oysters, Ostrea, are also attached to solid substrate. They have the capability of changing from one sex to the other alternately. A few species of oysters form commercially valuable pearls. Pen shells, Pinna, are common, thin-shelled, tropical species which have a sharply triangular shape. They frequently are seen partly buried, pointed end down, in sand. The upper end is very sharp and can inflict deep cuts. Among the nonattached and motile pelecypods are the quahog or cherrystone clams, Venus, which are typical of the many burrowing clams of sandy or gravelly beaches. The soft-shelled clams, Mya, have a thinner shell than the quahog and prefer softer bottoms like soft sand or mud. The same is true of the angel-wing clam, Barnea, and the razor clam, Ensis, which also have fairly delicate shells. Clams with thin shells have, in general, longer svphons and bury themselves deeper than do the harder-shelled species. Thev are also more active and faster diggers than the hard-shells. One species of pelecypods has left the general protectively shelled pattern of adaptation described in previous species. The shipworm, Teredo, is actually an almost naked clam that uses its reduced shell to bore through wood. Like termites, shipworms utilize wood for food. The scallops, Peeten, are actively swimming pelecypods. Thev clap their shells vigorously together to force water out of the body cavity and thus move by a sort of jerky jet propulsion. Scallops have photoreceptors around the edge of the mantle and also have long, sensitive tactile and chemoreceptive tentacles attached to the mantle's edges. All clams and oysters are edible, and some are delicious, but because of their filter-feeding habits, they may at certain times of the year and in some waters become poisonous. Obviously, pelecypods from foul or polluted waters should be avoided. In the summer, toxic bacteria and protozoans mav become plentiful enough in some waters to cause pelecypods, with these microorganisms in their digestive tracts, to become poisonous. It is also said that during the breeding season some clams and oysters are toxic due to the accumulation of sex products in them. At any time of year, these animals spoil quicklv. They should be eaten only when absolutely fresh. It is generally a good rule to eat pelecypods only during months which have an "r" in them; the cooler months, September through April. THE INVERTEBRATE LEGIONS 125 Squids and Octopuses: Class Cephalopoda {"head foot") — Figure 41 In this group, there is a definite tendency to lose the shell and to become active and predatory. Some cuttle fishes, Sefia, still have a shell known as "cuttlebone," but it is entirely internal. The common squids, Loligo, have a fragile pen-shaped shell buried in their flesh and the octopuses are without shells. In all cephalopods the foot is developed into a circlet of tentacles around the mouth. The mouth has a parrotlike beak, and the bites of cephalopods are poisonous, sometimes dangerously so. Although these animals are not often inclined to bite, their means of defense centered elsewhere, they should be handled only with care. While the tentacles are not dangerous, the bites from octopuses have been known to result in death. Defense consists of protective coloration coupled, in the case of octopuses, with secretive habits or, in the case of squids, with speed of swimming. When molested, all cephalopods have the habit of emitting a smoke screen of dark inky fluid and then making a hastv retreat. In some deep sea squids, the ink is luminescent. Cephalopods are the largest, speediest, and most active of invertebrates. The giant squid, Architeuthis, is a favorite food of the sperm whale, lives in the deep sea, and reaches a length of about 50 feet including the long tentacles. Some octopuses have an armspread of over 12 feet. Octopuses are shy animals, but large ones are less shy than small ones. The nervous system of cephalopods is extremely well developed. Their giant nerve fibers conduct nervous impulses at a very rapid rate. Their eyes are very much like mammalian eyes, and these animals see very well. These features provide cephalopods with the extremely fast reflexes necessary for active predatory animals, but they do not necessarily imply intelligence, for which integration of senses and actions is needed (Chapter 2). In fact, the intelligence of these animals, like their ferocity, is often overestimated. Cephalopods are the original inventors of jet propulsion. They draw water into the mantle cavity and forcibly eject it from a siphon. They can change direction by moving the siphon, but they usually swim backwards. The color changes of cephalopods are the most rapid and striking to be seen in the entire animal kingdom. By means of rapidly redistributing pigments in their pigment-bearing cells or chromatophores, they can cause complete change of color or even waves of color to pass over their bodies in a matter of seconds. Most cephalopods match background by means of color change. They also change colors when feeding or in response to stimuli leading to fear. The tentacles of cephalopods are organs used for food-catching, movement (octopuses crawl), and reproduction. These tentacles bear suckers, the power of which is remarkable. Even a small octopus of only a few feet in tentacle spread can hold onto a rock so as to be almost immovable. Care should be taken by the diver not to let an octopus grasp him and a rock at the same time. This the octopus will not do as an aggressive action, but only if it is first molested. During copulation, sperm is transferred to female cephalopods with one specialized tentacle which usually has the suckers reduced in number and size. In some octopuses, namely the paper argonaut, Argonauta, and Philonexis, the 126 UNDERWATER GUIDE TO MARINE LIFE sperm-bearing arm is actually detached from the male and placed in the mantle cavity of the female. This arm was originally thought to be a separate animal, a worm, and was named Hectocotylus. Most octopuses do not lose their sexual arms during copulation, however. Cephalopods can be divided into two groups on the basis of how many arms they have. The squids have ten arms, two of which are much longer than the rest. These are very swift-swimming animals which are common near shore or at sea the world over. They eat fishes and invertebrates. The common squid, Loligo, is a common shore form and reaches a length of Wi feet. Sepiotenthis (^color fhotograph^ is found in the West Indies and is very beautifully colored. Both of these show preference for traveling in schools in precise military formation. Sometimes as many as several dozen may be seen lined up swimming together. The octopuses have eight tentacles, usually all of equal length, and are of bottom-living habits, eating other invertebrates such as molluscs and crustaceans mostly. Some are small and pelagic and fairly good swimmers. Others are extremely flattened and sluggish, spending most of their time attached to rocks. But most are shy and nocturnal, though active, bottom animals. Klingel (1944) gives a sympathetic account of the common octopus, Octofiis (Color Plate 10^ He says that the best way to spot the home of one of these shy animals is to take note of the presence of a pile of shells and stones which the octopus accumulates near its home. After feeding, the octopus has the peculiar habit of drawing hard objects to its body. Octopuses show fear by means of a livid white color phase, but they also have color phases composed of virtually every shade or pattern imaginable. Ink is used both for a defensive smoke screen and for recognition of one octopus by another at night. The paper argonaut, Argonmita, is an octopus, the female of which forms a beautiful papery shell as an egg case. The argonaut is an animal of rather deep tropical waters, but it appears at the surface when it is breeding. BRAGHIOPODS: Phylum Brachiopoda— Figure 41 This phylum is mentioned chiefly because one species holds the geological longevity record for all animals; Lingiila has been in existence for 400 million years. Brachiopods are not now very varied, but were among the most common of animals many million years ago. They look like bivalve molluscs since they have two shells and filter-feed on plankton, but they are actually quite diff^erent anatomically. The lamp shells are brachiopods that grow attached to rocks by a stalk. Brachiopods are fairly common in deep water offshore, but a few are found in shallow waters. ECHINODERMS: Phylum Echinodermata ("spiny-skinned") The starfishes and their allies form an important group evolutionarily speaking since they have a larva that looks very much like the larva of some prochordates. Therefore, the chordates (including vertebrates) and the echino- derms are thought to have common ancestry. Adult echinoderms are characterized by the tube feet by means of which THE INVERTEBRATE LEGIONS 127 thev move. These are little tubes that have suction discs at their ends and w^hich operate by means of an internal bulb that pumps water into the muscular foot, causing it to expand, and that sucks water from the foot, causing it to contract. The strength of several hundred feet working together is considerable. There are five groups of these spiny-skinned animals. Starfishes: Class Asterida — Figure 42 As the name implies, these are star-shaped animals. They are especially diverse on the west coast of North America where the sun star, with a diameter of up to 2^2 feet, is the largest starfish known. All starfishes are carnivorous on bivalve molluscs. They setde over a bivalve and exert a steady pull on the bivalve's shells. Eventuallv, the bivalve tires, and the starfish then extends its stomach into the bivalve and digests the bivalve right in its shell. Starfishes are not very variable in form. (The very common large West Indian star, Oreaster, is shown in figure 43.) Some starfishes are blue, and others are red, brown, or even bright yellow. All can regenerate arms if these are lost. Brittle Stars: Class Ophiurida — Figure 42 These are the most acti\'e of the echinoderms, able to move quite rapidly with their long serpentlike arms. In fact, they are often called "serpent stars." They are fragile and may even purposefully break arms off when they are handled, a defense mechanism that occupies the predator's attention while the brittle star makes its escape. In the middle of the underside of the body is a mouth with serrate, toothlike edges. These are used to sift out and chew the small pieces of organic matter on which these animals feed. Most species are nocturnal, and some are very common in deep water. (A typical brittle star is shown in Color Plate 10.) The basket star or Gorgon's head, Gorgoncephalus, is an extraordinary and very sluggish brittle star that has finely branched arms. It is common in the tropics and reaches a width of 2 feet, but because of its sedentary habits and protective seaweedlike shape, it often goes unnoticed. It may be found adhering to corals or clumped in the branches of sea plume gorgonians, where it catches small, planktonic animals with its tentaclelike arms. Sea Urchins and Sand Dollars: Class Echinida — Figure 42 The bodies of these animals are rounded and spinv in the case of sea urchins and flattened and smooth in the case of sand dollars. There is a shell formed of calcareous plates and on the underside is a five-toothed chewing apparatus called "Aristotle's lantern" with which these animals scrape their algae food from hard surfaces or chew organic matter. These animals move slowly by means of tube feet and depend on their spines or hard shells for protection, for their meat is tasty, forming an excellent chum for fishes. Sea urchins and sand dollars range in all waters of the world over all types of bottom. The long spines of the needle-spined sea urchins, Diadeina, common in all warm seas, are very sharp and brittle and easily pierce flesh, gloves, or even shoes. The spines break UNDERWATER GUIDE TO MARINE LIFE BASKET 5TAR &GorqoncephaluS- Vo 2 f i- Fig. 42. Echinoderms. Drawn from life. THE INVERTEBRATE LEGIONS 129 ^4 / / Fig. 43. Top. The heche-de-mer, Holothuria, averaging about eight inches, is an extremely sluggish mud-eater. The pearlpsh uses the heche-de-mer' s cloaca for a home Cphotograph courtesy John A. Moore^. Bottom. The West Indian star, Oreaster, is commonly found on sandy, weedy hottom among plants like turtle grass, Thalassia, and the alga, the merman's shaving brush, Penicillus. -^'/ off in the flesh and Hberate a puipHsh fluid which stains the wound. The sensa- tion is painful, but the effects are not generally long-lasting. The bits of spine that cannot be pulled out soon dissolve in the flesh. Most sea urchins do not bear such sharp, long spines, but sharp-spined species should not be handled barehanded. Some that have short spines have poison glands associated with these spines. These are the flexible urchins, one of which, Aerosovia, has a sting reported by Phillips and Brady (1953) to be as painful as that of the Portuguese man-of-war. Luckily, this urchin keeps to rather deep water. 130 UNDERWATER GUIDE TO MARINE LIFE Sea Lilies: Class Crinida — Figure 42 These are almost exclusively deep-water animals and are among the most common of animals found there. Deep-water sea lilies are stalked and very plantlike. There are a few sea lilies that break off their stalks and swim by waving their arms. One of these is the sea feather or feather star, Antedon, and it sometimes ranges to waters only about 100 feet deep. It is of temperate distribution. Sea Cucumbers: Class Holothurida — Figure 42 The onlv echinoderms that have lost their spinv or hard skin are the sea cucumbers, which have an appearance exactly like their name. They exist in shallow and deep waters all over the world and may be either chunky and muscular or elongate and delicate. A ring of tentacles encircles the mouth. The tentacles are tactile and chemoreceptive organs. Tentacles also serve to shovel mud and sand, the sea cucumber's source of food, into the mouth. Some sea cucumbers have the defense mechanism of eviscerating themselves through the mouth by means of a violent contraction of the body when they are disturbed. The sea cucumber soon regenerates the internal organs. The beche-de-mer, Holothnria (/ig. 43), is a common form found in tropical seas the world over. (Its commensal relationship with the pearlfish is described in Chapter 10 under that fish.) INVERTEBRATE CHORDATES: Phylum Chordata This is the phylum to which the vertebrates belong, but there are some invertebrate members of this phylum. These invertebrates are the species that have no backbone, hence are truly invertebrates. But they do have other internal, anatomical characters that vertebrates have (including a dorsal, tubular nervous system), and these place the invertebrate chordates or prochordates in the same phylum as vertebrates. All of the prochordates are filter-feeders, and most of them are small. Cephalochordates : Class Cephalochordata — Figure 44 These are the lowest forms of fishlike animals. Avifhioxus and its close allies are small, up to 3 inches or so, and are common buried in clean, sandy shores of much of the world. They lie in the sand with the mouth protruding, filtering plankton from sea water by means of a large basketlike pharynx. Hemichordates : Class Hemichordata — Figure 44 The acorn worms, Balanoglossus, and others, live in holes in soft-bottomed shores and feed, as do other prochordates, by filtering sea water. The acornlike "head," which is used both in burrowing and feeding, separates them from other wormlike animals. Some members of this class are small, sessile, and colonial. They live on the sea bottom and are not likely to be encountered by the diver. THE INVERTEBRATE LEGIONS 131 Tunicates: Class Tunicata — Figure 44 The sea squirts are the most common prochordates and are most hkely to be encountered. They are Httle, round, or elongate, vaselike animals that have two openings, an incurrent opening for taking in water, and an excurrent one for passing water out. Thus, they resemble clams in this respect and filter-feed exactly as clams do. Ciona and Molgula are two common, shallow-water sea squirts that live attached in groups to rocks or pilings in shallow water. The name "sea squirt" is derived from the habit that these animals have of squirting water when they are handled. There are several almost transparent members of this class that might be mistaken for comb jellyfish. Like comb jellyfish, they are pelagic and swarm on the open sea. Some of them are solitary and others are colonial and all are filter feeders. Salpa is one solitary form of fairly large size. Pyrosoma lives in the tropics, is colonial, and has living in it a luminescent bacteria. Large masses of Pyrosoma, when stimulated, emit light strong enough to read by. Young (1950) reports that this light may be a protective device since fishes that seize these animals drop them when their lights go on. Avv\pVi ioxuS-2 in. r> Fig. 44. Prochordates. Drawn from life. SALPIO SECTION THREE CHAPTER 8 THE LOWER FISHES— Lampreys, Sharks, Rays, and Ratfishes The vertebrates— fishes, amphibians, reptiles, birds, and mammals— compose the dominant and most advanced of the chordates (Chapter 7). This is an extremely diverse group, united by the possession of at least traces of a backbone composed of separate vertebrae. They also possess a cranium which encloses and protects the specialized anterior end of the central nerve chord, the brain. It is through the specialization of the brain and the allied sense organs (eves, nose, ears, etc.) that the advanced vertebrates are able to respond more delicately to subtle modifications of their environment than any other animals. Thus they have become the dominant living organisms on the earth and in the sea. It is not at all clear where the vertebrates came from. Probably some of the prochordates had common ancestors with some vertebrates, but this occurred so long ago (at least 500 million years) that the story is not preserved in fossil form for us to interpret today. THE JAWLESS VERTEBRATES: Glass Agnatha {"imthout jaws") This group, widespread in the temperate zones of both hemispheres, includes descendants of a once prominent array of fishes called "ostracoderms" which lived over 300 million years ago. The ancient forms were mud-eating, bottom-living marine animals which looked more conventionally fishlike than their eellike descendants, the lampreys and hagfishes of today. These have survived probably because they took up a very specialized mode of life— parasitism. It is a matter of definition whether to call the lampreys and hagfishes "parasites," "carnivores," or "scavengers." Theoretically, the parasitic habit begins with a stage in which an animal merely partakes of part of the living host's tissues but lives a largely independent life. The mosquito is an example. Gradually, as evolution progresses, the dependence on the host increases until the parasite is virtually completely dependent on the host. Such is the case with the tapeworm. Lampreys and hagfishes are a borderline case. They are specialized in feeding B2 THE LOWER FISHES 133 methods but are not so host-dependent as to have become nearly as anatomically altered as the tapeworm. Some biologists prefer that they be called "specialized carnivores," but since they feed on living prey, the term "parasite" is better. As is so often the case with parasites, the body form has tended to become simple and degenerate, in this case elongate and with a poorly developed skeleton of cartilage. The hagfishes tend to be scavengers, too, since they feed on dead or dying prey. The Agnatha are distinct from all succeeding vertebrates because of their lack of paired fins and jaws. In the modern forms the mouth is merely a round disc with a hole in the middle containing a rasping tongue. This mouth acts as a powerful sucking mechanism. Hence, these forms are sometimes called the "Cyclostomata" ("round mouth"). With this mouth they are able to do such things as grasp and bore into their fish prey, build a nest of stones, anchor themselves to rocks, pilings, or even boats, and grasp their mates while breeding. Because of the lack of paired fins, they are not very good swimmers. Though they can dash for prey at considerable speed, their steering is not as good as that of fish with more fins and shorter bodies. One does not see them swimming about nearly as much as one sees them attached to some item of substrate waiting to strike, almost snakelike, at some unlucky fish that happens to pass their way. The two groups of agnathans are distinct, and each attacks its prey in a quite different way. LAMPREYS: Family Petromyzontidae There are usually seven gill openings on each side, and there are no barbels on the snout. The eves are large, and the dorsal fin is discontinuous. Lampreys swim vigorously but not too efficiently, and are active, voracious animals. They attach themselves to fishes, preferably at the softer throat, side, and belly regions, and scrape away the skin with the rasping tongue in order to suck the blood and eat some of the flesh. Their saliva is an anticoagulant. Very often this either kills the prey outright or so weakens it as to cause delayed death. Larger fishes do recover. They prefer the soft-scaled fishes. Hard scales seem to be protection against attack. Not much is known of lampreys in the sea, though they spend most of their lives there. They are most commonly observed in streams in the spring, where they go to breed after a certain unknown number of maturing years in the sea. In this respect, they are like salmon and are said to be anadromous. During their journeys up rivers, they perform remarkable acrobatics in climbing rapids and low falls with the aid of their sucking mouths. In the riffles, a nest is scooped out of gravel and large numbers of small eggs are laid. These hatch into Ammocoetes larvae which are nonparasitic but look much like the adults. After spending three to five years in fresh water, having attained a 6-inch length, the Ammocoetes go to the sea to mature, and the breeding cycle begins again. Lampreys may become landlocked as they have in the Great Lakes, where they have all but destroyed the lake trout and whitefish industries. In spite of a repulsive appearance, they are edible and quite tasty. Formerly they were a popular food in New England and in Europe. Lampreys are found in all temperate seas. 134 UNDERWATER GUIDE TO MARINE LIFE Fig. 45. Sea lamprey. SEA lamprey: Petromyzon marinus Size: Up to 3 feet. Weight: Up to 3 pounds. Distribution: The North Atlantic of Europe and the United States south to Chesapeake Bay and rarely to Florida. Identification: Same as for the family. Habits: This lamprey is known to attack members of the cod and mackerel families. Probably it also preys on other soft-scaled fish. Fish are sometimes caught by fishermen which bear the scars of such attack. They have not been recorded attacking warm-blooded prey such as man, but if one does attach to an arm or a leg, so strong is the grip, it cannot be pulled off. However, raising the animal out of the water will cause it to release its hold. Similar Species: The Pacific lamprey, Entospheniis tridentatiis, is common from Alaska to southern California. It is landlocked in Goose and Clear Lakes in California. Its prey mainly consists of salmon and steelhead trout. This species is very similar to the East Coast form. SLIME EELS OR HAGFISHES: Family Myxinidae These are distinguished from lamprevs by the four pairs of short barbels on the head, lack of eyes, a somewhat smaller size, and a continuous dorsal fin. MOUTH Fig. 46. Pacific hagfsh. THE LOWER FISHES 135 The prey is usually dead or dying fishes though it is possible that healthy fishes may sometimes be taken. Thus, they may be considered a sort of cross between a parasite and a scavenger. They bore into the prey, usually at the neck with the tongue, or they may even enter the body through the mouth. Once inside, the prey is eaten from inside out and only a sack of skin is left. Great losses to fishermen have occurred when hagfishes have entered the bodies of fish caught in sill nets or on set lines. Over a hundred hagfishes have been retriexed from the bodv of a single fish. They seem to locate prey by smell rather than by sight as do lampreys. They are mainly nocturnal and lie buried in the mud most of the time. Hagfishes are sometimes caught on lines themselves. They swallow a hook deeply, even as far as the anus. These fishes do not enter fresh water to breed. Small numbers of large, shelled, adhesive eggs are laid in the sea where they sink and stick to the first object they touch. The name "slime eel" is derived from the fact that they can exude huge, dripping quantities of mucus from the skin when caught. A large one can fill a two gallon bucket with slime exuded from the mucus glands on the sides. Their distribution is world-wide in temperate seas, and they seem to prefer cool water. PACIFIC hagfish: Bdellostoma stouti Size: Up to 2 feet. Distrihution: From southeastern Alaska to southern California. Identification: Same as for the family. Habits: This hagfish is especially abundant around Monterey Bay at depths of 10 to 20 fathoms. It preys chiefly on disabled or netted rockfish, Sehastodes, and various flounder. About twenty large eggs are deposited on rock or shell bottom in early summer, but breeding may occur at any time of year. These eggs are oblong and connected together by hooks at their ends. Similar Species: The slime eel (hagfish), Myxine glutinosa, is the Atlantic form, found from the arctic south to Cape Hatteras. It keeps to deeper waters at the southern end of its range and reaches 18 inches in length. THE SHARKS AND RAYS: Class Chondrichthyes {"cartilage fish"), Subclass Elasmobranchi {"plated gills") This subclass omits the strange ratfishes which will be discussed later. It is a very ancient group which has been in existence for over 300 million years. With all that time available for their evolution, it would be expected that sharks and rays would be very diverse in body form and habits, but thev are not. Evidently their body plan has been a successful one, and it has not been varied nearly as much as the bony fishes for instance. There are but two basic body plans, a cylindrical one as seen in sharks and a flattened one as seen in rays. This group is a mysterious one. The underwater swimmer should avail himself of every opportunity to observe its species since the habits of very few are well known. For this reason, the groups will be considered here in some detail. Furthermore, because of the great amount of talk, based on little knowl- 136 UNDERWATER GUIDE TO MARINE LIFE 2'"' DORSAL FIN PRE CAUDAL PIT LATERAL LINE LABIAL FUHROy CAUDAL (tail) fin PELVIC FIN Fig. 47 . The external anatomy of sharks and rays. edge, about some species, particularly sharks, one has to be careful in his judgment about what to believe. This is a marine group with the exception of very few species. The entire skeleton is made up of gristle or cartilage. An enormous advance has been made over agnathans in that sharks and rays possess true jaws and paired fins (the pectorals and pelvics), characteristics which give great versatility in habits and motion. The best field characteristics are (1) the possession of several gill slits, (2) the tail in which the upper lobe is much the largest (heterocercal tail), (3) the position of the mouth under a pronounced snout, and (4) the lack of true scales of the type seen in bony fishes. Sharks and rays possess scales of a verv interesting and basic type. These are called the "dermal denticles" or "placoid scales" and are anatomically very similar to the teeth of all vertebrates (with the exception of agnathans) in that they possess enamel, dentine, and pulp cavity. The scales are scattered over the body of sharks and ravs and are usually small, giving, however, a certain roughness to the skin. Very large sharks have large scales which can produce cuts on a diver. Formerly, carpenters capitalized on this roughness and used sharkskin, or shagreen, as sandpaper until the advent of more modern abrasives. In some parts of the body, these scales are greatly enlarged, on the bodies of skates, Raja, for instance. The sting ray's spine, the sawfish's saw teeth, and, most notably, the teeth in the mouth of all sharks and ravs are all evolutionarily modified and enlarged placoid scales. Even we humans owe what few teeth we have left to their origin as placoid scales. The teeth of sharks and rays are modified greatly for a wide variety of functions and are constantly replaced from the inside of the jaws as the functional teeth on the outside of the jaws are worn down or lost Qfig. 48). A few species have several rows of teeth functioning at once, particularly the shell-crushing kinds such as Port Jackson sharks and eagle rays. THE LOWER FISHES 137 Fig. 48. Top. One of the authors is holding a small nurse shark, Ginglymostoma. The barbels, used for locating prey by taste and touch, are clearly shown. In spite of the docile nature of this shark, it has occasionally been known to bite. Bottom. The inside of the jaws of the tiger shark, Galeocerdo, are lined with several rows of teeth. Those at the bottom of the picture replace those at the top as the latter are worn down. 138 UNDERWATER GUIDE TO MARINE LIFE Most sharks and rays possess a hole just behind the eye. This is the spiracle and is exceedingly important to' the bottom-living species for the purpose of breathing. In the normal breathing pattern of fishes, water enters the mouth, passes the gills where oxygen is extracted, and is expelled through the gill slits. In bottom-living forms, notably rays, it is not possible for water to enter the mouth for the simple reason that in so doing the animal, with its underslung mouth, would gulp more sand and mud than water. Therefore, water enters the spiracle into the throat and gill cavity. Pelagic and mid-water species such as mackerel sharks do not need the spiracle and may lose it completely. In man a vestigial spiracle is seen in the Eustachian tube, which connects the ear and throat cavities. We can understand something of the durabilitv of the sharks and ravs over the centuries by examining their senses and body form. While it is true that sharks and rays possess very limited brain power, some senses are remarkably developed. The sense of smell is powerful and is the chief means of finding food. The lateral line, also important in finding food, is sensitive to low frequency water vibration and "hears" in water in the same way, perhaps, as our ears perceive air vibrations. The ampullas of Lorenzini on the snout sense hydrostatic pressure and perhaps also temperature change. Thousands of tiny sensory crypts on the skin act like taste buds. The ear, entirely internal, gives the animal a sense of position and balance. It is no wonder that, with such equipment for smelling, "hearing," and "feeling" water, sharks and rays have little need of eyesight. The eyes are quite capable of defining images, but their extreme nearsightedness is well known. It has been experimentally shown that a dogfish will not react by sight to an object until it is only one foot away. Nevertheless, sight should by no means be underestimated until more is known about it. One "sense" that is poorly developed, it seems, is that of response to pain. In body form, sharks and rays are also well adapted. They are admirably streamlined and beautiful to watch under water. Though none are as fast or as maneuverable as the best among the bony fishes, the advance over lampreys and hagfishes is great. One shark, Carcharinus, a ground shark, was clocked at 40 mph over a very short distance (Budker, 1947). The fins of typical sharks are beautifully designed hydrodynamically but fairly rigid in contrast to those of bony fishes and are thus used mainly as planing devices like the fins on a submarine. The vertical fins, dorsals and anal, prevent yawing and rolling. The paired fins, chiefly the pectorals which are moderately movable, give a dynamic or changing pitch equilibrium. In other words, they are used to point the shark's body up or down. The tail fin propels the shark and also turns it left or right. The upper lobe of the tail fin is longer, which tends to drive the head down. To counteract this, the head is flattened and tends to point the body upwards. If we add up all of these factors, we see that sharks have, in spite of fin rigidity, remarkable stability in all dimensions, even though maneuverability and some speed are sacrificed; sharks, for instance, cannot stop suddenly but can only come to a gradual halt. Stability is desired because it reduces to a minimum the amount of effort needed to keep the fish going in any one direction. It makes all motion extremely smooth and graceful. Certainly, sharks are as graceful as any animals that swim. Stability is also THE LOWER FISHES 139 desirable for another reason. Sharks are heavier than water and possess no swim bladder as do the bony fishes. They must swim slowly but constantly or they will sink. Some species have become so attuned to constant motion that they use it to help circulate water through the gills and will suffocate if held stationary. This constant motion of the active species makes them appear menacing. The fastest sharks are those in which the bodv is shortened and compact and the tail lunate, that is, has nearly equal lobes. These are the mackerel sharks, family Isuridae. One shark familv, the hammerheads or Sphyrnidae, has made an attempt to increase maneuverability by extreme flattening and widening of the head which is then used as a forward rudder. Another attempt in increased maneuverability resulted in the flattened rays, where the pectoral fin has been greatly expanded and has gradually taken over all locomotion. All sharks and ravs have internal fertilization, which is uncommon in bonv fishes. The males possess claspers on the pelvic fins which are inserted into the females. A few sharks lay eggs and are said to be oviparous. Most are ovovivip- arous, the eggs hatching in the female's body and the young being born alive shortly thereafter. A few sharks are truly viviparous as are mammals and have a "placenta" by means of which the females nourish the pups as they grow. Some ovoviviparous rays have a sort of internal feeding in which the female's "uterus" secretes a milkv substance for the young. Sharks and ravs produce few young, the recorded extremes in numbers varying from one to eighty-two, and perhaps averaging below twenty. All sharks and rays are carnivorous. A great variety of food is eaten, but the sharks are mainly fish-eaters and the rays mainly mollusc- and crustacean-eaters. A few are cannibalistic. Tiger sharks are especiallv liable to eat other sharks and rays when these are disabled or hooked. Hammerheads like to eat rays. Gudger (1932) reports finding fifty-four spines of the eagle ray, Aetohates narinari, and sting ray, Dasyatis say, in the head and jaws of one large hammerhead. There is a record of the skate, Raja, eating other small rays. The Sharks: Order Pleurotremata {"side openings") The typical shark is commonly visualized as a large, ferocious, swift predator with very little on its single-track mind other than the general destruction of all life of the sea, including man. We shall soon see how incorrect this vision is, for while it is true that some sharks are among the largest and most voracious of animals, most sharks do not conform to such a pattern. In general, the sharks are fairly stout and subcylindrical of body, but several are almost as flattened as rays and even parallel the rays in habits. Others are long and slim. They range in size from 18 inches to nearly 60 feet. Several dwell in the deep sea, and some pelagic species are brightlv luminescent. Even though we must lay aside the stereotyped idea of what a shark looks like, there are three characteristics that all sharks possess: (1) five to seven gill slits on the sides just back of the head (hence the name "Pleurotremata"), (2) pectoral fins that may be large or small but are never connected to the head, and (3) definite eyehds. In all of these traits they diff^er distinctly from rays. 140 UNDERWATER GUIDE TO MARINE LIFE The various shaiks are best told apart by shape and action. Except for a few species, they offer little in the way of pattern or coloration as an aid to identification. Sharks are predominantly fish-eaters, but they are neither fast nor agile enough to catch fast healthy fish so a great part of their diet consists of the disabled or slower fishes. They prefer the easy mark, a wounded fish or a straggler. (Exceptions to this are the fast and active members of the family Isuridae, the mackerel sharks.) Sharks, contrary to popular account, do not have to turn on their sides or backs to bite, even though they occasionally do so. We shall examine the various sharks' diets under later species discussion, but one item of diet deserves more than fleeting attention. This concerns the ever- discussed question: Do sharks attack man? The answer is an undeniable yes. But we must be careful in this assertion to be attentive to the circumstances under which attack is most likely to occur and which species might be responsible. There is a great tendency to overdramatize the possibility of attack. This was particularly damaging to the morale of service men deployed in "shark waters" during the last war. There is an equal tendency toward debunking. Having met sharks and not having been chewed to bits, some people, especially under- water swimmers, have made the extreme assertion that sharks are not dangerous at all. The authors have heard some underwater swimmers claim that sharks are cowards. This seems to be based on the fact that these unfortunate people had once or even several times met a shark which happened to run when aggressively approached. Sharks seem to hunt their prey in a manner somewhat as follows. The first stimulus to feed comes from two sources: smell (olfaction) and, to a lesser extent, reception of low frequency water vibration (through the lateral line). These have the effect of increasing the speed of the shark's movements, causing it to make short hunting turns and figure eights which are of an exploratory, food-seeking nature. These responses reinforce the original olfactory stimulus and also notify other sharks of the presence of food through the stimulus of low frequency vibration. Gradually, as the shark or sharks approach the food, the exploratory turns become more and more direct until the food is struck. Sight may play a part in these last stages. Contact with food is the final stimulus which brings on the "feeding mood." At this stage, sharks are extremely aroused, active, dangerous, and not discriminating in taste. Tiger sharks, Galeocerdo, have been known to become so excited when in the "feeding mood" as to bite blindly at any object including sacks of coal, old shoes, and tin cans. The sharks that are implicated in attack on man are: 1. White Shark or Man-Eater, Carcharodon carcharias. This is the sole deliberate attacker and is always dangerous. It does not seek out man but is liable to attack because it habitually eats large-sized prey. It is aggressive, and its triangular, trenchant teeth (to 2 inches long in a 15-foot specimen) are extremely efficient. Large ones can swallow objects as large as a whole man. Fortunately, this shark is not common. THE LOWER FISHES 141 2. Tiger Shark, Galeocerdo ciivier. This common species is more nocturnal than most sharks. It reaches large size, has large, trenchant teeth, and is especially dangerous when in the "feeding mood." Tiger sharks commonly congregate about areas where refuse is dumped at sea. 3. Hammerhead Sharks, Sphyrna species. This maneuverable species grows to large size and is possibly guided to food more quickly than other sharks because of the widely separated nostrils. The teeth are rather small. 4. Mako Sharks, hums species. These are the fastest sharks that swim and are among the most aggressive. They may be dangerous under certain circumstances even though they are almost exclusively fish-eaters. 5. Lemon Shark, Hy-poprion hrevirostris. It seems that this is a disagreeable and unpredictable species that may, when large, be dangerous. 6. The Ground Sharks, Black-Tipped Sharks, etc., Carcharinus species. Any large members of this genus may be dangerous. The teeth are large and trenchant. 7. Sand Shark, Carchnrias taunis. There are some doubtful records of persons being bitten by this shark. 8. Blue Shark, Prionace glauca. This is a pelagic species, long reputed to be dangerous. There are few records of attack.' 9. Leopard Shark, Triakis semifasciata. This is a small shark which has attacked divers on the Pacific coast. The conditions under which shark attack is likely are not very well known. Springer (1943), in the best work yet published on this subject, has thoroughly studied and compiled a list of well-authorized cases of attack. The following analysis is largely adapted from him and from Budker (1947): 1. Influence of Time of Day. There is an increase of frequency of attack with increasing darkness. The bulk of attacks occur at dusk, but records of attack are known for almost every time of day and night. Increased frequency of attack at dusk is probably due to decreased visibility with darkness and increased shark activity then. 2. Influence of Water Clarity. As is the case with dusk, decreased visibility means that sharks increase dependency on their lateral line for feeding. 3. Influence of Water Depth. The danger seems to be slightly greater at or near the surface. Sharks are in shallower water at night rather than during the day. Nevertheless, there seems to be little or no danger in shallow waters inshore of surf or inside of coral barriers. Sharks must at least have access to deeper waters through channels or inlets. Sharks as a whole are most abundant in fairly shallow waters not far offshore. 4. Influence of Temperature and Geographical Location. Springer has mapped the cases of shark attack on a world-wide basis, and it is apparent that sharks are most dangerous in tropical and subtropical waters in temperatures of over 66° Fahrenheit. The incidence of attack is far higher in southeastern Australia than anywhere else in the world. The beaches of Sydney are fenced in, so great is the danger of attack thought to be. Otherwise, shark attack does not seem to be concentrated at any one place. 142 UNDERWATER GUIDE TO MARINE LIFE 5. Influence cf Movement. Sharks seem to be attracted by inert objects at the water's surface. During the last war, it was urged that men in the water stay in a group and keep moving slowly. The effect of movement on sharks is poorly understood. 6. Influence of Various Scents. a. Refuse. Continuous dumping in any one area will soon train sharks to visit there frequently. Many food scents are attractive to sharks and dumping grounds are often danger spots to be avoided. b. Blood. This is the strongest attractant of sharks. Almost any shark may be dangerous in the presence of blood, which serves to drive them into a frenzy leading up to the dangerous "feeding mood." De Witt (1955) records an instance of a 3-foot leopard shark, Triakis, attacking a diver when blood was in the water. Fast (1955) states that there is "a strong possibility that with the present increase of spearfishing along our [Pacific] coast, the number of attacks by sharks will increase. Those individuals who indulge in this sport should . . . avoid seepage of blood, fish juices, or other attractants." c. Man. Sharks seem neither attracted nor repelled by the scent of man. d. Shark Repellents. During the last World War it was found that decomposing shark flesh had a repellent effect on sharks. Further investigation by the United States government during World War II led to the development of cupric acetate cakes, which can have up to 100 per cent effectiveness (but usually less) in repelling sharks from baited lines for 3 to 4 hours. Springer lists several of these experiments in which baited lines, some repellent-treated and some not, were used to fish for sharks. The repellent lines were vastly less attractive to the sharks, and most bites on repellent lines were made before the repellent had had a chance to dissolve in the surrounding water. There are no proven color-cloud repellents as is sometimes supposed, but sound may occasionally serve to repel sharks. The definitive work on the effect of sound on fishes is yet to be done. To these factors we add several suggestions for an advisable course of action when the swimmer is in the presence of sharks under water. 1. Observe all you can about the shark's behavior. Try to identify the shark. Identification is most important. The swimmer's actions should be very different if he meets a white shark than if he meets a nurse shark. 2. Don't panic. Keep moving in a fishlike way, smoothly and regularly. Try to stick fairly close to protected places, like coral heads or rock outcrops. 3. If there are several of you, stay close together. 4. Avoid blood and offal in the water. 5. If attacked, fight back. This may serve to drive the shark away. The most efficient way to disable a shark is to grasp a pectoral fin tightly as the shark passes by and then to sht open its belly or gills with a knife. This, however, is not a job for amateurs. 6. Don't attack sharks. They may look docile and timid and present an easy target, but a shark, if attacked, cornered, or wounded is very liable to THE LOWER FISHES 143 turn on the swimmer. To draw a shark's blood in the water would be most foolish. This would only serve to incite this shark and draw others to the scene. Under water, sharks possess a puzzling mixture of ferocity and timidity. Their large size, abundance in shallow and comfortably warm waters, and the many unknown facets of their behavior make them ideal subjects for observation and study. With the elimination of doubt, overconfidence, and hysteria, the underwater swimmer is in the unique position of being able to study these creatures firsthand. The Primitive Sharks There is but one family of these protoselachians ("shark forerunners") remaining in the seas today. It is the sole survivor of the ancient stock that gave rise to modern sharks and is distinguished from them in its primitive jaw suspension, dentition, and other anatomical characters. PORT JACKSON SHARKS: Family Heterodontidae The Latin name of the family refers to the fact that the teeth are of two kinds; those in the front of the mouth are small and pointed for grasping and those in the back are flattened like molars for crushing. There are two large dorsal fins, both preceded by a stout spine. The family contains only one genus of wide distribution, but not occurring in the Atlantic. Fig. 49. Port Jackson shark. 144 UNDERWATER GUIDE TO MARINE LIFE PORT JACKSON SHARK (horned shark) : Heterodontus francisci Size: Up to 4 feet. Distrihtition: Point Conception to the Gulf of California. Identification: Same as for the family. Habits: This is a fairly common shark living near kelp in shallow or deep water. It keeps to the bottom and eats hard-shelled invertebrates which it crushes with its strong molar teeth. The large and unmistakable eggs may sometimes be found wedged in among rocks near the low tide zone. The Modern Sharks These sharks are divided into three large groups. The rays are derived from them. Notidanoids: Superfamily Notidanoidea These most primitive of modern sharks comprise a rare and poorly known group. They reach large size and are mostly in moderately deep water of 100 to 400 fathoms, only occasionally coming within the depth range of the diver. The single dorsal fin, long, cylindrical body, and the presence of six to seven gill slits are good field marks. They are widespread in the temperate and tropical waters of the world. COMB-TOOTHED OR COW SHARKS: Family Hexanchidae The teeth of these sharks are unique, being completely unlike in the two jaws. These large sharks are found in all warm oceans but are not common, it appears. They are ovoviviparous. Identification is made easy by the presence of but one dorsal fin. TEETH x^ ^^S-50. Cow shark. cow SHARK (six-GiLLED SHARK, GRisET, MUD shark) : Hexanchiis griseus Size: Up to 15 feet, exceptionally to 26 feet. Matures at 6 feet. Weight: Up to 1,300 pounds. Distribution: All warm seas. In the Atlantic from Cape Hatteras to the Caribbean. In the Pacific from southern California to British Columbia. Not rare in Cuban waters and off San Diego. Identification: Dark brown or gray with a light streak along the lateral line. There are six gill slits. THE LOWER FISHES 145 Habits: This is primarily a deep-water form living at over 50 fathoms. It gets closer to the surface in northern waters and at night. Probably it is a voracious predator on large fishes such as dolphin, swordfish, marlin, and other sharks. Except when feeding, it is probably a sluggish fish. Those who have eaten the flesh report it to have a strong purgative eff"ect. Similar Species: The perlo or seven-gilled shark (Heftranchiis ferlo) is another widespread, large, voracious fish which looks like the cow shark, but is without the light, lateral line stripe and has seven gill slits. Galeoids: Superfamily Galeoidea This is the group of the great majority of modern sharks. All have five gill slits and two dorsal fins which are not preceded by a spine. Because these are active sharks, there is a tendency to reduce or even lose the spiracle. In separating the families, the size and placement of the dorsal fins and gill slits are of importance. CARPET SHARKS OR NURSE SHARKS: Family Ginglymostomidae The two large dorsal fins are both posterior to the pelvics. Gill slits three, four, and five are over the pectoral fins and four and five are so close together as to appear as one. The mouth is preceded by two short barbels and is connected to the nostrils by a short groove. As befits bottom-living, sluggish sharks, the lower tail lobe is poorly developed. These sharks are ovoviviparous. NURSE SHARK (gata) : Ginglymostoma cirratum— Figure 48 Size: Averages 5 feet. Commonly up to 8 to 10 feet. Exceptionally to 14 feet. Distribution: Rhode Island to Brazil, but not commonly north of Cape Hatteras. Also on the western coast of Mexico. Identification: The young have small black spots, which disappear with maturation, scattered over the dingy brown body. The head is large and flat, and the eyes small. Habits: This is a sluggish, inoff^ensive, and rather fearless shark which is very common in shallow tropical waters. It is often seen lying in a protected hole or cove or under a ledge near coral heads. Pairs come into very shallow water in order to mate. At such times they loll about with their dorsal fins out of water. It is quite a sport among local boys to grasp them by the pectoral fins in order to secure a ride. When mating, the male holds the female by a pectoral fin with his teeth. Thus, female nurse sharks are likely to look a bit ragged about the pectorals after a few mating seasons. This shark, although normally inoffensive, will definitely bite when annoyed. Fig. 51. Nurse shark. 146 UNDERWATER GUIDE TO MARINE LIFE Nurse sharks nose about coral and seaweed in search of food which consists of a great variety of small fishes and invertebrates. At times, they eat refuse, amounting almost to scavengers. Several rows of small teeth in each jaw form a crushing or holding dentition. CAT SHARKS: Family Scyliorhinidae The two dorsal fins are nearly equal in size and posteriorly placed, the first being behind or over the pelvics. The tail fin is much like that of the nurse sharks with a poorly developed lower lobe. The fourth and fifth gill slits are over the pectoral, and the spiracles are fairly large. This group is rather uniform with many small, cat-eyed, prettily patterned species, most of which live in deep tropical and temperate waters beyond the range of the diver. Cat sharks are most common in the South Pacific and Indian Oceans, but are world-wide in distribution. The common spotted dogfishes, Scyliorhiniis species, of Europe belong in this family. They are oviparous as far as is known. SWELL shark: Cephaloscyllium uter Size: Up to 3 feet. Distrihiition: Monterey Bay to Acapulco, Mexico. Identification: Ground color is brownish yellow. Barred with black and spotted with black and white. Habits: This is the only cat shark which is common inshore. It may be seen in kelp beds particularly. It presumably eats a wide variety of animal foods, especially fish, and can swallow surprisingly large objects with its capacious mouth. The common name is derived from its habit of inflating like a puffer when molested. The flesh will cause diarrhea if eaten. SAND SHARKS: Family Carchariidae The two dorsal fins and the anal fin are of nearly equal size and the posterior margin of the first dorsal is just over the anterior margin of the pelvic. Fig. 52. Swell shark. Fig. 53. Sand shark YOUN6 ^JOCrORPlSH COLOR PLATE 7 HO(jf=iSM COLOR I'l.ATK N THE LOWER FISHES 147 All of the gill slits are anterior to the pectoral fin. There is only one genus in the family- The species are ovoviviparous and are cosmopolitan in warm waters. SAND SHARK (sAND TIGER ) : Carcharias taurus Size: Averages 4 to 6 feet. Up to 9 feet. Weight: Up to 250 pounds. Distribution: Maine to Brazil. Most common in warm temperate waters and rare in the tropics. Identification: Grayish brown above, fading to white below. The young have light spots and blotches which disappear with age. Habits: This is the most common large shark swimming over sandy beaches. It is fairly sluggish but can move at good speed when the occasion demands. It feeds principally at night on fishes and squids and occasional large crustaceans, the sharp, pointed teeth serving admirably to hold such slippery prey. They may feed in large schools in very shallow water. Although there are no records of attacks on man, this shark is to be respected. MACKEREL SHARKS: Family Isuridae These are unquestionably the most active, swiftest, and most voracious of sharks. The modifications toward swiftness are good field marks. These are (1) a compact and torpedo-shaped body, (2) a high first dorsal fin placed at the highest point on the back, (3) reduced second dorsal and anal fins of nearly equal size, (4) a narrow caudal peduncle reinforced with lateral keels, and (5) a lunate tail in which the lobes are of nearly equal size. In many of these respects, they resemble the mackerels and spearfishes which are among the very swiftest of bony fishes. The gill slits are anterior to the pectoral fin. There is no spiracle. The coloration is characteristic and like that of most pelagic fishes being a sea-blue above and white beneath, an example of obliterative coloration (Chapter 2). These are primarily fish-eaters and are found in all temperate and tropical seas. They are ovoviviparous. MAKO (sharp-nosed MACKEREL shark) : Isurus oxyrhincus .Size: Averages 6 feet. Up to 12 feet and possibly a little larger. Weight: Averages 150 pounds. Up to 1,200 pounds or more. Distribution: Tropical and warm temperate Atlantic. Common in the Bahamas. In summer to New England. Identification: Anal slightly posterior to second dorsal. Coloration is slate-blue above, abruptly changing to dirty white below. The snout is conical and sharply pointed. The teeth are slender and recurved for holding fish. Habits: This may be the swiftest shark that swims. It is pelagic and pursues swift, schooling fish such as mackerel and herring for food. Occas'onal squids and sea turtles are taken. This fish is best known for its habit of leaping spectacularly from the water especially when hooked, but it is most often seen basking at the ocean's surface, solitary or in small schools, with the large dorsal fin pro- truding. It can be dangerous, particularly around boats. It's near relative, the bonito shark of the Pacific, Isurus glaucus, has been known to attack men and 148 UNDERWATER GUIDE TO MARINE LIFE Fig. 54. Mako shark. Fig. 55. White shark. boats. The young are few (one to five) and large at birth (20 pounds). This is an excellent food fish. Similar Sfecies: The Pacific bonito shark or Pacific mako, Isiiriis glaucus, may be identical to the Atlantic form. It is found from Monterey Bav to the south and over the entire South Pacific. The porbeagles and mackerel sharks (Laiinza species) replace the makos in the Atlantic and Pacific in water below 60° Fahrenheit. These are stouter sharks than the makos and have the second dorsal and anal directly opposite each other. The generic name Lamna, is derived from a Greek word depicting a horrible, man-eating fish, and was used to intimidate naughtv children. WHITE SHARK (man-eater) : Carchawdon carcharias Size: Averages 8 to 12 feet. Rare over 15 feet. Recorded up to 36V2 feet. Weight: At 21 feet, one weighed 7,100 pounds and its liver weighed 1,005 pounds. Distribution: Nowhere common. Most common near southeastern Australia. Occurs in all warm temperate and tropical seas. Identification: Similar to mako and mackerel sharks, but the second dorsal and anal fins are even more reduced; the teeth are triangular with serrate edoes and the shape stouter. Coloration is slate-grav with a black axillar spot. Over 15 feet, the color fades to lead-white. THE LOWER FISHES 149 Habits: This may be the most voracious thing that swims with the exception of the killer whale, Orca. It is active, fast, and very powerful and is responsible for most of the authenticated reports of attacks on man. Though principally pelagic, it will enter very shallow water. It has been known to attack boats without provocation or especially when hooked or harpooned. The white shark feeds on large fishes, sea turtles, and e\en sea lions. One shark was found with a 100-pound sea lion nearly intact in its stomach. Another specimen of 15 feet was found to contain a large Newfoundland dog. Another ISVz-footer contained two 6-foot sharks. The young are few in number and large, being over 100 pounds in weight when born. Shnilar Species: One recently extinct Carcharadon is worth mention because of its spectacular size. Only the teeth have been found, but they indicate a body length of over 60 feet. A full-grown man could have stood upright in this huge fish's mouth. BASKING SHARKS: Family Cetorhinidae These are the largest fishes found in temperate waters. They are much like mackerel sharks in form, but the gill slits are very, long, the mouth huge, the snout short, and the teeth minute and very numerous. The skin is very rough and abrasive. They are ovoviviparous. There is but one species of circumpolar distribution. BASKING SHARK (bone shark) : Cetorhimis maximiis Size: Commonly to 30 feet. Exceptionally to over 40 feet. Weight: Commonly to 8,600 pounds. Distribution: Temperate seas of both hemispheres. South to Cape Hatteras and California. Not arctic but gets into very cold water. There is a center of abundance to the southwest of Iceland. Identification: The color is a light gray to brown, lighter on underside. Habits: The common name, basking shark, is derived from the habit of basking at the water's surface with the enormous dorsal fin and even part of the back out of water. At such times, it is very easy to approach this shark in a boat or in the water. Nevertheless, it should be remembered that this huge fish is 150 UNDERWATER GUIDE TO MARINE LIFE extremely powerful. One stroke of the tail can easily stave in a small boat. These fish travel singly, in small groups, or in large schools of up to a hundred. At times, several may travel in tandem and have been mistaken for sea serpents. As is often the case with many giant fishes, these are plankton feeders, having fine gill rakers with which to strain their food from the water. When feeding, they travel open-mouthed through the water. Excellent pictures of this may be seen in the motion picture Men of Arran. Basking sharks sometimes leap from the water. They have been decimated by oil hunters; a single liver yields from sixty to six hundred gallons of oil. They are usually seen only in summer when the plankton supply is good, retreating to deeper water and possibly not feeding in winter. WHALE SHARKS: Family Rhineodontidae There is no fish that approaches the whale shark in size except the basking shark. The last two gill slits are over the pectoral fin, and the first dorsal is fairly far back over the small pelvics. The tail fin is huge. The mouth is large and not preceded by a snout. There is only one species of cosmopolitan tropical distribution. Fig. 57. Whale shark. WHALE shark: Rhincodon typiis Size: To 60 feet or more. Very rare over 30 feet. Weight: A 38-foot whale shark weighed 26,594 pounds. Distribution: All tropical seas. Most numerous off the Philippines and the coasts of southern California and western Mexico. Identification: The coloration is a unique gray to red or green crossed with yellowish stripes and spots. Three prominent ridges run down the sides. Habits: This is a very inoffensive fish, so sluggish that it has been rammed by boats on several occasions. Despite this, it can probably travel swiftly. The feeding habits, teeth, and gill rakers are much Hke those of the basking shark. It also basks on the surface and may school, though it is usually solitary. It sometimes gulps plankton, small fish, or squids while standing vertically beneath the school of food. THRESHER SHARKS: Family Alopidae These are large sharks with a large, extended upper tail lobe. Otherwise they resemble mackerel sharks. THE LOWER FISHES 151 Fig. 58. Thresher shark. THRESHER SHARK (fOX SHARk) : AlofiaS VulfeS Size: Matures at 14 feet. Rarely to 20 feet. Weight: Matures at 500 pounds. Rarely to 1,000 pounds. Distribution: Cosmopolitan in all warm seas. North to the state of Washington and to New England in summer. Identification: Blackish to slate and brown, sometimes with a metallic luster. The teeth are rather small. Habits: This is a pelagic species of ferocious appearance, but it is quite harm- less and is almost exclusively a fish-eater. It provides one of the very few examples of cooperative feeding among fishes, another being found among the puffers. Several sharks herd moderate-sized, schooling fishes such as mackerel, herring, or bonito into tight groups with the aid of their tails. Attack is thus made easier. They may also use the tail to stun prey. The thresher is swift and voracious. One was found off Scodand containing a bushel of fish in its stomach. Contrary to rumor, threshers do not attack whales. It is a good food fish. SMOOTH DOGFISHES: Family Triakidae The first dorsal is large and just behind the pectoral fin. The second dorsal is reduced in size, but it is not as reduced as in the similar requiem sharks, Carcharinidae, so is noticeably larger than the anal fin. The lower lobe of the tail fin is poorly developed. The last two gill slits are over the pectoral, and the body is slender and tapering. These fish are very much like the requiem sharks and are characteristic of shallow, warm temperate, and some tropical waters the world over. However, thev tend to small size and are considered harmless. There are both ovoviviparous and truly viviparous species. The flesh of most species is good as food. SMOOTH dogfish: Miistehis canis Size: Averages 2V2 to 3V2 feet. Up to 5 feet. Distribution: Winters from Cape Hatteras to Chesapeake Bay. Arrives in New England from May to June when the bottom water warms to about 45° Fahrenheit. Rare in tropics, but known to Brazil. Identification: Plain gray to brown. Can change shade from dark gray to pale, pearly white when over a light bottom. The change takes two days for maximum effect. This is one of the few sharks able to change color. Habits: This, like all smooth dogfishes, is a shallow water fish rarely straying beyond the 10-fathom limit except in the southern part of its range. It is most common in bays and may enter brackish and fresh water. The teeth are arranged 152 UNDERWATER GUIDE TO MARINE LIFE Fig. 59. Smooth dogfish. in a pavement and are adapted for shell crushing. Thus, it is a crab-eater primarily, though a few lobsters are taken. It also takes small fishes, squids, and some molluscs and worms. It may scavenge. This is the fish on which experi- ments on shark feeding methods have been done. Smell is the chief aid for find- ing food, the shark moving its head from side to side and therefore sensing food alternately more strongly with one nostril than the other until the direction of the food is detected. It has been shown that plugging one nostril causes the fish to take much longer to find food. This shark is viviparous. As many as twenty-seven foot-long young are born in June and July. The gestation period is ten months. Similar Species: The gray smooth hound, Mnstelus californicus, is found from northern California to Baja California and the sicklefin smooth hound, Mustalus lunulatus, from San Diego to Colombia. Both are very much like the East Coast species in habits and appearance. Fig. 60. Leopard shark. LEOPARD shark: Triakis semifasciata Size: Females mature at Wi feet and grow to 5 feet. Males grow to 3 feet. Distribution: Oregon to Baja California. Identification: The pattern of black crossbars and spots is unmistakable. Habits: This is a bay and shallow-water shark similar to the smooth dogfish. Although small, this shark is easily incited by the presence of blood and will bite. Similar Species: The brown smooth hound Triakis henlei, looks very much like the Mustelus species. It is very common in San Francisco Bay and other enclosed California bays. THE LOWER FISHES 153 REQUIEM SHARKS: Family Carcharinidae These sharks form the bulk of "typical" modern sharks and are characterized by a large first dorsal fin which is usually far anterior to the pelvic. The second dorsal is much reduced, just above and of a similar size to the anal. Added field marks are the position of the fourth and fifth gill slits over the pectoral fins and the characteristically shaped caudal fins, which is preceded by well-developed precaudal pits. These fishes all have a well-developed third eyelid or nictitating membrane which serves to keep the eyes free of foreign matter (fig. 64^ Spiracles are small or missing. The teeth are bladelike, triangular, and trenchant, and only one row is functional at a time. Requiem sharks are ovoviviparous or viviparous and characteristic of warm temperate to tropical seas the world over. TIGER shark: Galeocerdo cuvier Size: Common at 12 feet. Said to reach 30 feet, but largest recorded in Atlantic was 18 feet. Weight: Common at 850 pounds. Distribution: Common in the Caribbean. North to Cape Hatteras and rarely to Cape Cod. On the West Coast to southern California. Cosmopolitan in warm temperate and tropical seas. Identification: The snout is very blunt and rounded and the labial furrow prominent. The sides are blotched and striped, but this pattern disappears with age. The teeth are large, few in number, and characteristic (fig. 48^. Hahits: This is one of the most voracious sharks, having extremely catholic tastes in food. It eats sea turtles, sea lions, fishes large and small, crustaceans, offal, and other sharks. It also preys on sting rays, the spines having little effect on the shark. One, when captured, vomited the arm of a man who had been murdered at sea and dismembered. This is the most abundant large shark of the Caribbean and seems to be attracted by the cast-off offal of slaughterhouses. In such dumping areas, it may be dangerous. There are records of its having attacked man, though these are few even in areas of shark abundance. In spite of the foregoing, this is a sluggish shark, normally becoming active only when aroused to the "feeding mood." One expects to see tiger sharks in shallow water near land, but they may be encountered almost anywhere, even in harbors or the deep sea. It is the most prolific shark, normally bearing thirty to fifty living young. The largest number of young ever recorded in a brood, eighty-two, was from a female of this species. The young are small, only 18 inches long at birth. e^ Fig. 61. Tiger shark. 154 UNDERWATER GUIDE TO MARINE LIFE Fig. 62. Blue shark. BLUE shark: Prionace gJaiica Size: Commonly 9 to 12 feet. Rarely to 20 feet. Weight: 164 pounds at 9 feet. Distribution: Cosmopolitan in warm temperate and tropical seas. North to British Columbia in the Pacific and Nova Scotia in the Atlantic. Not found in shallow-water areas such as the Gulf of Mexico. Identification: The coloration is a beautiful indigo blue, fading to pure white below. The body is slim, the pectorals very long, and the snout very long and pointed. The first dorsal is rather far back for a requiem shark. Habits: This is the most common pelagic shark, held in fear by sailors and in contempt by whalers. It is normally sluggish except when aroused. There are few recorded attacks by this species on man, but it should not be counted as harmless. It is most commonly seen at the surface, basking with the dorsal fin out of water or following after ships in search of food. For the latter reason, they have been named chiens de mer by the French. When hunting prey, they are active and have been observed to repeatedly blink with the third eyelid. They are most active at night. The blue shark's greed is practically legendary. It feeds on all sorts of small pelagic fishes and squids. Frequently, it gathers in great numbers about whaling ships to share in the carcass. At such times, when the sea is clouded by blood, the sharks flounder all about the whale. Occasionally, the whalers strike the sharks with their blubber spades and horribly mutilate them, but even when so wounded, they go right on feeding until some of their fellows dispose of them. There is a record of one captured blue shark, which had had its liver removed, trying to catch a mackerel after being thrown back in the water. Another gutted blue shark was recaught on a hook baited with its own intestines. This shark is viviparous and rather prolific, producing as manv as fiftv-four pups at almost any time of year. LEMON SHARK (yellow shark) : Ncgaprion brevirostris Size: Matures at 7 feet. To 1 1 feet rarely. Weight: 265 pounds at 9Vi feet. Distribution: Cape Hatteras to Brazil. Most abundant in the Caribbean. Also in the South Pacific and Indian Oceans. Identification: The first dorsal is small for a requiem shark and the second dorsal rather large. The color is yellowish brown, fading to a distinct yellow on the underside. The snout is broad and rounded. THE LOWER FISHES 155 Fig. 6?>. Lemon shark. Hahits: This is a strictly inshore shark and may be seen around harbors and sounds. It even enters such fresh waters as the Amazon. It is mainly a fish-eater. It is one of the most common of Caribbean sharks and is not to be trusted, its dis- position supposedly being erratic. This shark is viviparous, bearing young in the spring and summer. GROUND sharks: Carcharinus species This is the largest genus of sharks, containing many species that are difficult to identify even by experts. Therefore, they will be considered as a group. Their distribution is world-wide in warm temperate and tropical waters. Identification: As a group, the ground sharks are bluish or grayish to brownish, generally lighter on the belly. The first dorsal is very high and placed forward. The second dorsal is over the anal and, like the anal, is small. The upper pre- caudal pit is very large. There is no spiracle. Hahits: These are inshore and reef-inhabiting sharks of fish-eating habits largely, but they will take many foods. The larger ones are not to be trusted. The teeth are trenchant and large. When the cry "shark" goes up along inshore waters, the cause is usually a ground shark. They are curious fish and have approached the authors to within a foot, returning several times for repeated "looks," but seldom gave trouble. In fact, they usually fled when approached, but that is no guarantee that they invariably do so. One, the Lake Nicaragua shark, Carcharinus nicaragiiensis, is the only habitually fresh-water shark and has been known to attack people. A list of the most common species with hints for identification follows: East Coast: 1. Cub Shark (Bull Shark, Fish Shark, Southern Ground Shark): C. Leiicas This species is never found far from land. It frequently travels in groups. The diet is extremely varied, this shark can be dangerous at times. The snout is very short, broad, and rounded for the genus. It attains a length of 10 to 12 feet, though rare over 8 feet. From Cape Hatteras to Brazil and strays to Massachusetts. 2. Small Black-Tipped Shark (Spinner Shark): C. linihatus The eyes are large, and the pectoral, dorsal, anal and lower caudal fins are tipped with black. It is rather small, averaging 6 feet and reaching 9 feet. This is an active, swift, largely pelagic species. It may school at the 156 UNDERWATER GUIDE TO MARINE LIFE NICTITATIN& MEM6RAIME Fig. 64. New York ground shark. surface and frequently leaps from the water, turning as it does so. A tropical species which strays to Massachusetts. 3. White-Tipped Shark: C. longiniamis This shark has a very broad and rounded first dorsal tipped with grayish white. The snout is rather short. It may reach 13 feet and is the most pelagic of the ground sharks. Found in the tropical and warm temperate Adantic. 4. New York Ground Shark (Brown Shark, Sand-Bar Shark): C. milherti The first dorsal is very high and placed far forward, and the second dorsal is directly over the anal. It reaches 8 feet and 200 pounds. This is the most abundant ground shark in the shallow waters along the mid- Atlantic Coast. It may enter river mouths and is the only large shark com- monly found in Long Island Sound. It also enters the canals of Venice. It is largely a bottom feeder on all sorts of fishes and invertebrates. The young are born in shoal waters in the north from June to August. From southern New England to Brazil. Also in the Mediterranean and off west Africa. West Coast: 5. Bay Shark: C. lamieUa This is a large shark of up to 15 feet, most common in bays and shallow waters. The fins are not dark-tipped. From southern California to Mazatlan, Mexico. 6. Gambuso: C. aziireus This shark reaches 10 feet, is more pelagic than the former, and has dark-tipped fins. SHARP-NOSED SHARK: ScoUodofi teuae-novae Size: Averages from 2 to 2^2 feet up to 3 feet. Weight: Averages 8 to 10 pounds. THE LOWER FISHES 15"; Distribution: Common in the Caribbean and Gulf of Mexico and in summer to Cape Hatteras. Straggles to New York. Identification: This shark is a southern counterpart, ecologically speaking, of the smooth dogfish. The small size of the second dorsal fin, lack of spiracles, short gill slits, and prominent labial furrow serve to identify it. The color is brown to gray with the dorsal and tail fin edged in black. The body is quite slim. Habits: A typical beach, surf, and estuary species, this shark has never been found more than one or two miles from land. It eats a wide variety of small in\'ertebrates and fishes and has not endeared itself to fishermen because of its habit of preying on their baits. It is viviparous. Similar Sfecies: The Pacific Sharp Nose, Scoliodon longurio, is found from southern California to Panama. It reaches 3Vi feet and 9 pounds. It is rare in scientific collections but should be common in bays in its range and should be looked for. SOUP-FIN (oil shark, tope) : Galeorhiniis zyopterns Size: Females reach 6^2 feet, males up to 6 feet. Weight: Females reach 100 pounds, males 60 pounds. Distribution: Pacific Coast from San Francisco, to Cedros Island. Identification: Similar to the blue shark but has short pectoral fins, no pre- caudal pit, and is brown to gray in color. The dorsal, caudal, and pectoral fins are tipped with black. Habits: This voracious shark eats all sorts of fishes, squids and other inverte- brates, and even carrion. It is the most valuable shark commercially and is sought for its liver which is rich in vitamin A. The flesh is very good, and the fins are used to make the famous sharks-fin soup. This is an inshore species. Fig. 65. Sharp-nosed shark. Fig. 66. Soup-fin shark. 158 UNDERWATER GUIDE TO MARINE LIFE common both at the surface and on the bottom. The htters are large, averaging thirty-five and recorded from six to fifty-two. Gestation may take one year. It is ovoviviparous. HAMMERHEADS: Family Sphyrnidae The widely expanded and flattened head serves to separate these sharks from the otherwise similar requiem sharks. The eyes and nostrils are at or near the ends of the head's extensions. Supposedly, the head, acting like a forward rudder, gives these sharks greater maneuverability than other sharks. The very large and wide-set nostrils are highlv efficient. These sharks are usuallv among the first to arrive when blood is in the water. Hammerheads are found in all warm temperate and tropical seas. hammerhead: Sphyrna zygaena Size: Matures at 7 to 8 feet. Averages 8 to 11 feet. Reaches 15 feet. Weight: 900 pounds at 12 feet. Distribution: All tropical and subtropical seas. North to Cape Hatteras (rarely to Cape Cod) and southern California. Identification: Slate to brownish gray in color. Habits: This is a strong-swimming, voracious, active, surface shark, found at sea or inshore or even in tidal, brackish waters. Like many other sharks, it often basks at the surface with its caudal and dorsal fins protruding. Hammerheads have a definite liking for sting rav meat. They will also eat fishes of various kinds and other sharks as well. Great numbers of sting ray and catfish spines have been found in the heads, jaws, and bodies of hammerheads, apparently causing no discomfort to the shark. This shark is known to be dangerous to men even though the teeth are rather small. The northerly migration of schools of hammerheads in the summer consists mainly of small, 6-foot sharks. They are viviparous, bearing up to thirty-seven young. Similar Sfecies: Sfhyrna zygaena is merely one of three very similar hammer- heads. The great hammerhead, Sfhyrna tudes, is the largest of these. It reaches eONNCT Fig. 67 . Hammerhead shark and bonnet shark. THE LOWER FISHES 159 17 feet and 1,500 pounds. The little 5-foot bonnet shark, Sphyrna tihuro, is found north to New Jersey and San Diego. It is a shallow-water, bay species, and is harmless. The kidney-shaped head separates it from the three tvpical hammerheads. Squaloid Sharks: Superfamily Squoloidea This is the third and last subdivision of the modern sharks. The best field characteristic is the complete lack of the anal fin. They also possess two dorsal fins both of which are sometimes preceded by a stout spine and fine gill slits which are short and entirely in front of the pectoral fins. Spiracles are present and reproduction is ovoviviparous. Presumably, this group gave rise to the rays. SPINY DOGFISHES: Family Squalidae The dorsal fin spines are large and the labial furrows are long. These are small slim sharks of world-wide distribution. Many of them live in deep water, and all have very large eyes. The tail fin has no subterminal notch. SPINY DOGFISH (piKED DOGFISH, grayfish) : Sqiialus acanthhis Size: Averages 2^2 to 3 feet. Up to 4 to 5 feet. The females are the largest. Weight: Averages 7 to 10 pounds. Up to 20 pounds. Distrihution: Subarctic North Atlantic south to Cape Hatteras. North Pacific south to Point Conception. Identification: Slate to brown color with yellowish spots on sides. The shape is very slim and subcylindrical. Hahits: This is neither a swift nor very active shark. Nevertheless it is voracious, feeding avidly on small fishes such as herring, menhaden, and small mackerel. It also eats invertebrates, such as jellyfish, and refuse. It is a cold- water form, characteristic of waters of 43° to 59° Fahrenheit. Both its northern and southern movements are determined by these temperatures, and it is thus like the common mackerel except that its temperature range is a little lower. During the hot months and in the south it is found in deep waters down to 100 fathoms. It is by far the most numerous shark in cooler waters, usually found traveling in huge schools inshore, in brackish water, or a few miles offshore. These schools are constantly on the move, so their appearance is somewhat erratic. From one to fourteen young are born in the late autumn offshore after two years of gestation. When caught, this shark thrashes violently curling its body like a bow and striking out with the slightly poisonous dorsal fin spines. The wounds these produce may be painful. The flesh is good and is usually sold as "grayfish." Fig. 68. Spiny dogfish. 160 UNDERWATER GUIDE TO MARINE LIFE Fig. 69. Greenland shark. SLEEPER SHARKS: Family Dalatiidae The dorsal and pectoral fins are reduced in size, and the gill slits are short. These sharks are as sluggish in appearance as they are in reality. The teeth are very different in the two jaws. Many are inhabitants of the deep sea and have luminous spots. Their distribution is world-wide, and they are ovoviviparous. GREENLAND SHARK (sLEEPER SHARK, GURRY shark) : Somniosiis microcefhaUis Size: Averages 8 to 14 feet. Possibly to 24 feet. The females are larger than the males. Distribution: Arctic seas to Cape Cod. Identification: The color is coffee-brown to dark gray with tinges of violet. There may be faint, darker crossbands. Habits: This is the largest arctic fish and the only arctic shark. It may live in water as cold as 28° Fahrenheit, or as warm as 55° Fahrenheit. It probably lies near or on the bottom, rising only to feed and may be counted as the most sluggish of large sharks, offering no resistance to capture. In spite of this, it seems to be able to catch seals as well as large fishes such as halibut and salmon, though it will also eat crustaceans and jellyfish. It seems to be irresistibly attracted by carrion. Therefore, whalers know it well. In spite of rumor to the contrary, it attacks neither living whales nor men in kayaks, and it is quite harmless. The flesh is intoxicating and poisonous if eaten fresh, but is wholesome if dried or allowed to become partly rotten. Similar Species: The Pacific sleeper shark, Somniosiis j)acificus, is found south to southern California. Its habits are like those of its Atlantic cousin. ANGELFISHES: Family Squatinidae These raylike fishes are shown to be sharks by the possession of gill slits on the side of the "neck," pectorals which are not attached to the head, and free eyelids. They also possess characteristics which are superficially raylike: much flattened body form, enlarged pectoral and pelvic fins, two small dorsals placed far back, a row of spines along the mid-back, and eyes and enlarged spiracles on the top of the head. They are not to be considered the evolutionary link between sharks and rays, but they are midway between those two groups in habits. Thus, they are bottom-living but still use the tail in a sculling, sharklike manner for locomotion, little use being made of the pectorals except for steering. They are ovoviviparous and world-wide in distribution. MONKFiSH (angel shark): Sqtiatina diimeril Size: Matures at 3 to Wi feet. To 5 feet. Weight: At 4 feet weighs 60 pounds. THE LOWER FISHES 161 Fig. 70. Monkfish. Distribution: Summer visitor to mid-Atlantic states. Most common near Chesapeake, south to the Gulf of Mexico and the Caribbean. Not really plentiful anywhere. Identification: Blue to ash-gray above with spots and mottled pattern. May have some reddish tints. Habits: In summer, this is mainly an inshore fish. It lies on the bottom and may pardy cover itself with sand or mud as rays are wont to do. From this van- tage, it ambushes flatfish, skates, mullet, and other fishes. It also eats crustaceans O ' 7 7/ and molluscs, which it crushes with its strong jaws. In winter it strays to water as deep as 700 fathoms. The young are born in early summer inshore in num- bers up to twentv-five at a time. This shark has powerful jaws and may inflict a painful bite if molested. The teeth are small and numerous. The term "angel shark" refers to the angel-wing shape of the pectorals. Similar Species: The California angel shark, Squatina californica, is of similar size to the Atlantic forms. It is found from Alaska to southern California and is rather common in the south. The European monkfish, Squatina squatina, gets much larger, 8 feet and 170 pounds, and is very plentiful in Europe. Rays: Order Hypotremata {"openings beneath") Among the most remarkable of the animals of the sea are the rays, sometimes grouped as the order Batoidei. Most of them are greatly flattened, but just as there are raylike sharks, there are also sharklike rays, so body form is not dis- tinctive. The significant characteristics are ( 1 ) the enlargement and continuation of the pectoral fins onto the head, (2) the position of the five pairs of gill slits on the underside (forced there by the overgrowth of the pectorals), and (3) the absence of lower and usually also of upper eyelids. The rays probably developed from squaloid sharks in response to the develop- ment of expanded pectoral fins for greater mobility. The evolution of the ad- vanced rays (Myliobatoidea) from the primitive types (Pristoidea) was the result of the following steps: (1) the pectorals took over all locomotion, (2) dorsal and caudal fins were lost, (3) the body became flat, (4) the enlarged spiracles and eyes were moved to the top of the head, and (5) the tail became 162 UNDERWATER GUIDE TO MARINE LIFE Fig. 71. One of the authors is photograpJi'nig a large soutliern sti>ig ray, Dasyatis americana, four feet in diameter, which was found covered with sand sleeping on the hottorn. distinct from the body- We see this evolutionary transition in following the changes in body form from the sawfishes to the guitarfishes to skates and tor- pedoes and finally to the eagle rays. This evolutionary trend is an adaptation to bottom-living. But like almost every group of animals, there are departures from the principal theme, and so the rays have also risen from the bottom to take on the more pelagic life that we see in the eagle rays and mantas. Paralleling these evolutionary tendencies are distictive methods of locomo- tion. In the most sharklike and least flattened rays, the sawfishes and guitarfishes, locomotion is accomplished by powerful sculling motions of the tail as in sharks. The pectoral is used little. In the torpedoes also, propulsion is chiefly caudal, but this is probably because the disc is so inflexible, it being the location of the electric organs. The skates move almost entirely with the pectoral fins, un- dulating waves traveling in this fin from front to rear. Sting rays also use THE LOWER FISHES 163 undulating waves of the pectoral for slow locomotion, but flap their "wings" like a bird more than do the skates when they desire to move fast. The eagle and manta rays flap their wings exclusively and are able to move rapidly in this way. Turning in all rays is accomplished mainly or exclusively by the pectoral fins. Some rays, particularly the "flying" ones are among the most graceful of fishes. Thev are also among the most powerful things that swim as anyone who has hooked one can attest. As befits their largely bottom-living habits, rays are dark on the upper surface while the lower surface remains white and patternless. The upper surface is sometimes patterned to resemble the substrate over which the animal lives. Most rays are rather sluggish and nocturnal and, though powerful swimmers, are not as speedy as sharks or bony fishes. It is expected, therefore, that they would abandon the fish-eating diet of sharks for more easily caught food such as invertebrates. Most of them cannot see their food as they grasp it with their mouths, but must settle over the active food species with their bodies or dig for the less active ones in the bottom. They are the only fishes that are known to settle over and trap food with their bodies, but they do not envelop prey with their wings as has been supposed. Most rays possess many rows of small, sharp, or flattened teeth arranged so as to act in a grasping or crushing manner. Rays vary in size from a few inches to 23 feet (across the pectorals) and are found all over the world from arctic to tropical waters. Some sting rays and sawfishes even enter fresh water. Rays allow close approach since they are not timid animals. By day they are often seen asleep on the bottom. They are, therefore, easy prey for the swimmer, but the authors do not consider it either very good sportsmanship nor verv good sense to molest them. It must be remem- bered that even rather small rays are powerful, and many, if not most of them, can be dangerous, as we shall see in the following pages. There are fi\'e major groups (superfamilies) of rays. Sawfishes: Superfamily Pristoidea These are the most sharklike of the rays. The presence of the extended snout bearing saw teeth is an unmistakable and unique identifying character. This saw can be a very formidable weapon. Sawfishes are ovoviviparous. The saw of newly born sawfish is covered by a sheath in order to protect the female. 164 UNDERWATER GUIDE TO MARINE LIFE This sheath is lost soon after birth. The distribution of sawfishes is world-wide in warm, shallow seas. They enter brackish and fresh water. There is but one family containing a single genus. SAWFISHES: Family Pristidae sawfish: Pristis fectinatus Size: Commonly to 16 feet. Reaches 20 feet or more. Weight: Commonly to 700 pounds. Distribution: Most common off Florida and in the Gulf of Mexico. To Cape Hatteras in summer. Also south to Brazil. Identification: Grayish brown in color. Habits: These bottom-living rays are most plentiful in shallow bays and estuaries, entering brackish and even fresh waters. The shape indicates a sluggish but powerful animal. They rarely rise far from the bottom except to attack schools of small fishes, such as mullet, on which they feed. The saw is used in a sideways slashing manner to stun and impale these fishes, which are later eaten at leisure. The saw also serves for poking about in the sand or mud bottom for various items of food, mostly invertebrates. It is also a powerful weapon of defense. A sawfish will not attack man unprovoked, but even small ones should not be molested. Contrary to rumor, sawfishes do not attack whales, but they may attack fairly large fishes, although rarely. The young are born in summer and autumn after about a year's gestation. They are 2 feet long at birth. Young sawfishes make excellent eating. Guitar Fishes: Superfamily Rhinobatoidea In this group, the transition from sharklike to raylike forms is complete. The disc is either distinctly heart-shaped or round. The tail is always stout, and the dorsal and tail fins are well developed. A uniform shagreen covers the body. The mid-dorsal spines are somewhat enlarged. They are ovoviviparous and found in all warm temperate and tropical seas. GUITARFISHES: Family Rhinobatidae SPOTTED guitarfish: Rhinobatos lentiginosns Size: Averages 2 feet. Reaches 3 feet. The females are larger than the males. Distribution: Most common off Florida. To Yucatan and Cape Hatteras. Identification: Covered with small Hght dots over a brown to ash-gray ground color. The disc is wedge- or heart-shaped and the snout long. Habits: Guitarfishes swim close to the bottom using sculling motions of the tail for propulsion and the pectorals for steering. They may lie half buried in mud or sand and eat small molluscs and crustaceans. They are not often en- countered in water deeper than 5 to 10 fathoms and are even encountered in surf feeding on the gammarid shrimj-)s and sand crabs, Hippa, found there. Similar Species: Three West Coast species show a perfect graded transition to skates in the shape of the disc, though the tail remains stout in all. The shovel-nosed guitarfish, Rliinobatos productus, is much like the East THE LOWER FISHES 165 Coast form. It reaches 4 feet or over and ranges from central California to the Gulf of California. The mottled guitarfish, Zapteryx exasperata, has a shortened snout, wider disc, dark crossbands, and reaches 3 feet in length. It is found from San Diego to Panama. The thornback, Platyrhinoidis triseriata, has a skatelike rounded disc and skate- like spines on the back and shoulders. It reaches 3 feet and is found from central California to Baja California. Skates: Superfamily Rajoidea In these rays and all remaining rays, the tail is sharply marked off from the body, and the pectorals are the chief means of locomotion. Skates also have the dorsal and tail fins greatly reduced in size and the pelvic fins deeply notched so that they look double. The males possess very large claspers. The discs and tail are covered with enlarged prickles in patches which vary according to species. Skates are characteristic of temperate and arctic waters of the world. They are in the tropics, but only in very deep water. Some run up rivers. They are the onlv oviparous rays. The egg cases, called "sea purses' or "sailor's purses," are very common objects on beaches. SKATES: Family Rajidae This is the only family of skates and contains the largest of all elasmobranch genera, Raja, with almost a hundred species. The species of this genus have E&G CASE X-r Figs. 7?) and 74. Common skate Qeft^ and spotted guitarfish (rig^it). 166 UNDERWATER GUIDE TO MARINE LIFE similar habits and some are difficult to identify. Therefore, skates will be considered as one group. skates: Raja species Skates are ground fishes found over sand, mud, gravel, or shell bottoms. They are rather inactive by day, at which time they lie on the bottom, sometimes half buried. To bury themselves, they stir up the bottom with their pectorals and let the sand or mud fall back on them. They depend a good deal on protective coloration to avoid being detected, their grayish or brownish, often spotted or mottled bodies blending in well with the substrate. When alarmed they tend to press their bodies closely to the bottom and may form a suction between their bodies and the bottom which makes them difficult to dislodge. If further alarmed, they may curl themselves into sort of a ball and lash violently with their spiny tails. Some have electric organs on their tails able to deliver half a volt. The adaptive value of such a small shock is not known. The skates are not exclusively bottom fishes. Frequently, they come to the surface to bask in the sun after the manner of many sharks. Skates, like most rays, have many rows of small, pointed or flat teeth in the jaw which form a crushing or grasping dentition. Their prey is quite varied but usually consists of crustaceans, molluscs, worms, and small to medium-sized fishes. With active prey, such as fish, their method of securing food is by ambush. They lie on the bottom until a victim comes their way, then quickly pounce on it, holding it down with their bodies, and then grasping it in their jaws. This method of capture is made necessary by the position of the jaws on the underside. Locomotion is achieved by undulating waves passing from front to rear in the pectoral fin. Skates can swim surprisingly fast, at which time the pectorals are used in a flapping motion like the wings of a bird. The tail may aid in steering. Skates mate with their ventral sides together and are frequently caught in this position. One or both claspers are inserted. The egg capsules or sea purses Cfig. 73) are 3 to 8 inches long and contain one to several eggs. They are laid in the summer and adhere to rocks, shells, algae, etc., hatching in four and one-half to fifteen months. The courtship and mating procedure should be easy to observe by the swimmer. Males are smaller than females by about one-third. Sometimes, several follow a single female and strike each other with the spiny parts of the pectorals. These spiny pectoral patches are also used to grasp the female while mating. Skates make excellent food. Raie au heiirre noire is a French culinary delicacy. Skates reach 6 feet and 60 pounds but are mostly much smaller. A list of some common species follows: East Coast: 1. Common Skate (Litde Skate, Hedgehog Skate): Raja erinacea This is a small skate, reaching only 20 inches in length. The spines arc in a distinctive pattern with no spines of large size directly on the midline. This is the most common New England skate, found in shoal water and to 80 fathoms, usually over a sandy bottom. It ranges from Cape Hattcras to Nova Scotia. 2. Clear-Nosed Skate (Briar Skate, Summer Skate): Raja eglanteria There are two translucent areas on the side of the snout and there is a THE LOWER FISHES 167 prominent row of spines on the midline of the back and tail. The whole tail is covered with thorns of like size. The back is spotted or barred with dark brown. Reaches 2 feet in length. Found in shallow waters from Maine to Florida but common only from Massachusetts to Delaware. 3. Barndoor Skate (Sharp-Nosed Skate): Raja laevis The popular name is derived from its large size, which may attain a length of 5 feet and a weight of 35 pounds. The snout is prolonged and pointed and the body spines small. It is the only skate with pigmented mucus pores on the underside. Found from Newfoundland to Cape Hatteras. West Coast: 4. California Skate: Raja inornata The outer border of the pelvic fin is deeply concave and the color is dark olive to brown with vague mottlings. It reaches 2^2 feet. Found from the Strait of Juan de Fuca in the North Pacific south to Cedros Island, California. 5. Big Skate: Raja hinoculata There is a large ocellar spot on the inner side of each pectoral fin. The outer border of the pelvic is nearly straight. It reaches 6 to 8 feet in length and the egg cases are a foot long and contain up to seven eggs. Found from Alaska to southern California. 6. Long- Nosed Skate: Raja rhina The snout on this skate is very long and tapering. The overall size may reach 4 to 5 feet. Found from Alaska to southern California. Electric Rays: Superfamily Torpedinoidea These are very odd rays possessing an almost circular disc and a stout, some- what sharklike tail on which are situated two large dorsal fins and a large tail fin. The eyes are small; some electric rays are even blind. The body is smooth and peculiarly flabby. The pectoral fins are thick and rather immobile, being the seat of the large electric organs, which may weigh as much as one-sixth of the total weight. Because of the fact that typical muscle nerves go to these organs, they are thought to be a muscle tissue in which the electrical properties inherent in all muscle tissue have been accentuated at the expense of the locomotory properties. They are formed of thousands of hexagonal discs piled in columns and thus look like honeycombs. The resemblance to a voltaic pile is striking. A shock delivered by these rays is really a "train" of twelve to a hundred separate pulses each of which lasts about three hundredths of a second. Voltages delivered vary from eight to 220 depending upon the size of the ray and its species. Several shocks may be delivered in succession, but each one is a little weaker than its predecessor. A rest period is needed to build up the power to its full state again. The shocking power is far less than the really dangerous electric "eel," Electrofhorus, of South American rivers. Still, it is enough to knock down and temporarily disable a full-grown man. The purpose of the shocking power is a matter of some mystery. Almost certainly, it is defensive, and it seems likely that it may also be offensive. Presumably, electric rays are too slow-moving to catch the fishes, such as dogfish 168 UNDERWATER GUIDE TO MARINE LIFE and flatfish, that they are known to eat. They swim feebly with sculling motions of the tail, using the stiff pectorals only sparingly, if at all. Electric rays are among the laziest of fishes. They lie buried in sand or mud most of the time. They are found in shallow and deep waters of all temperate and tropical seas, mostly over soft bottoms. They are ovoviviparous and range in size from a foot in length to 6 feet and 200 pounds. ELECTRIC RAYS: Family Torpedinidae TORPEDO (electric RAY, NUMBFISH, crampfish) : ToYfedo nohiUana Size: Matures at 2 feet. Up to 5 to 6 feet long. Weight: Averages 30 to 75 pounds. Up to 200 pounds. Distribution: Both sides of North Adantic from Nova Scotia to Cape Hatteras (rarely Cuba) and from Scotland to west Africa. Nowhere plentiful. Identification: The disc is very wide and round and about one-half the total length. The color is chocolate to purplish brown, sometimes with obscure dark spots. Habits: Little is known other than the general information given above. This is one of the most electrically powerful of the electric rays. It is known to eat Fig. 75. Torpedo ray. large and even active fish such as salmon, eels, and flatfishes, a support for the argument that electric rays use their electric powers to secure food. Similar Sfecies: The California electric ray, Torpedo californica, is found from British Columbia to southern California and is locally plentiful in moderate depths. It reaches a length of 3 feet and a weight of 50 pounds. The litde electric ray or crampfish, Narcine braziliensis, belongs to a genus of small rays which has oval discs, not nearly as blunt anteriorly as the torpedoes. They have a mottled coloration which blends with the sand, resembling the THE LOWER FISHES 169 coloration of the round sting ray. This species eats invertebrates mostly and can produce shocks of thirty-seven voks. It matures at 9 to 10 inches in length and reaches 18 inches. Found from Florida and Texas south to Brazil. The True Rays: Superfamily Myliobatoidea The remaining rays are grouped in a single superfamily. All have the tail very well marked off from the body, usually like a whip, with one or several dangerous poison spines at its base (exceptions to this are found among butterfly and devil rays). This tail spine can be a very dangerous defensive instrument which can produce excruciating pain. Known remedies are a warm Epsom salts bath or an injection of 5 per cent permanganate of potash solution (Condy's Fluid). The injection of antibiotics or other antiseptics may also be of value. These are principally shallow-water fishes of the tropics and subtropics. Some enter fresh water. The species vary in size from under a foot to 23 feet in width. They are all ovoviviparous. Several species can utter a grunting sound. STING RAYS: Family Dasyatidae The discs are usually rhomboid but may be round (in the round sting rays). The tail is long and whiplike and bears no fins (except for round sting rays). These rays all have a long serrate spine at the base of the tail. This spine is not only poisonous, but it is large enough to cause a dangerous wound by itself. The stingers may vary from an inch or so to over a foot long. If one is lost or worn out another grows in its place so that two or even three may be present at once. The spine appears to be used mostly in defense but may rarely be used off^ensively. It was a matter of debate for many years whether or not the spine actually carried a poison. It was thought that the jagged wounds, and mucus and other foreign matter introduced into the wound, were enough to cause the extreme pain, swelling, cramps, inflammation, gangrene, or, rarely, death. (Stings are not always serious, but the fact that they may be so should be warning enough.) Then it was discovered that the narrow grooves along each edge of the spine contained a glandular, poison-secreting tissue. Furthermore, the poison is of a virulent nature. Poison glands may also be present at the base of the spine. Sting rays are usually not active, lying buried in sand with only the eyes and spiracle showing. However, .they are not nearly as sluggish as torpedoes and may frequently be seen swimming over reefs and sandy bottoms, being capable of considerable speeds. They are mostly crustacean- and mollusc-eaters, but do take fishes also. They commonly stir up the bottom with their pectorals in search of food. At times they are extremely abundant over shallow, sandy bottoms, almost seeming to pave the bottom. Large ones are powerful, but inoffensive. The authors, in order to stir one very large southern sting ray for purposes of photography Qfigs. 71 and 76X grasped it by the tip of the tail. It rose, circled slowly, swam a short distance, and sank to the bottom, burying itself by throwing sand over its back with its pectoral fins. This afforded an excellent chance to observe and photograph its exceedingly graceful and beautiful "flight," not forgetting, of course, that this seemingly inoff^ensive beast possessed a formidable weapon. Sting rays are ovoviviparous and found in warm shallow waters the world over. 170 UNDERWATER GUIDE TO MARINE LIFE r, / Fig. 16. Top. The round sting ray, Urolophus, is mottled to resemble its suhstrate. This ray, though small, has a very -powerful sting. Bottom. The southern sting ray, Dasyatis americana, is most easily identified hy the large, finlike fold beneath the tail. VM Hi^^^l m^.-,- "■'""jpipir^ i^^fPH 1 ^'i % WtBt^Kj^^K/ff^^^^^^^^^^^^' '^ THE LOWER FISHES 171 ROUND STING RAY: Urolofhus jamaicensis (^Also known as Urohatis^— Figure 76 Size: Usually 16 inches to 2 feet long. Distribution: Florida to the Caribbean. Sometimes to Cape Hatteras. Identification: The disc is round and light-colored with dark mottlings which blend in with the sandv bottom. The pattern may vary with the nature of the bottom, but this has not been demonstrated. The tail is short and has a caudal fin and highly developed spine. Habits: This is a species of shallow shoal and reef waters. It is not timid. One swam directly under the arm of one of the authors and settled quietly on the sand not two feet in front of him. It is more common than supposed, as swimmers will surelv discover, being found often about reefs in the Bahamas particularlv. It scoops out holes in the sand to expose its invertebrate food. In spite of its small size, the sting is dangerous, said to be more dangerous than other sting rays of much larger size. Similar Species: Members of this genus are common in most tropical waters. The Californian round sting ray, Urolophtis helleri, is very similar to the eastern form and is found from Point Conception to Panama. STiNGAREE: Dasjatis sabina Size: To only 15 inches across, the smallest Atlantic member of its genus. Distribution: Gulf of Mexico to Florida and the Caribbean. To Cape Hatteras and, rarely, to the Chesapeake Bay. Identification: The snout is pointed and rather long, and the pectorals are very rounded. It is yellow-brown to dark brown with a dark line down the midline. The midline of the back possesses prickles. Habits: This ray is more or less strictly tropical and is found in very shallow water. When walking through shallow water, it is well to use a shuffling motion of the feet. This tends to drive the rays ahead if they are present. The sting is probably not as painful as that of the round sting ray. Similar Species: The southern sting ray, Dasyatis americana (_fig. 76'), is the most common large species from Cape Hatteras to the Caribbean. It replaces Fig. 77. Stingaree. 172 UNDERWATER GUIDE TO MARINE LIFE the northern sting ray south of Cape Hatteras and these two rays are very similar in habits. It is easily recognized by the finlike folds under the tail behind the spine, a keel above this fold on the tail, and the angular shape of the pectorals. It averages three feet across the pectorals and probably reaches 4 feet. It often enters very shallow water, at which times the pectoral fins and tail may break the surface. The northern sting ray, Dasyatis centrura, reaches 13 to 14 feet in length and 5 feet in breadth. At Wz feet in breadth, it weighs 180 pounds. It is much like the southern sting ray but has a thorny tail with no keel on top of the tail. It is locally plentiful in shallow waters from Cape Hatteras to Cape Cod. The diamond sting ray, Dasyatis dipteninis, is much like the preceding species. It is found from British Columbia to Central America and is common in bays from San Diego southward. It reaches a length of 6 feet. Fig. 78. Butterfly ray. BUTTERFLY RAY (sAND skate) : Gyuinura viicTiira . Size: Averages 2 feet in breadth. To 4V2 feet across, rarely. Distribution: From Massachusetts to Brazil. Most common from the Gulf of Mexico to the Chesapeake. Identification: The breadth is much greater than the length, and the spineless tail is very short. The color is gray, brown, or even greenish or purplish. It can change its color to match the surroundings somewhat. Habits: This ray behaves much like the Dasyatis species, but is more active, especially in response to tide changes. This ray prefers coastal sandy bottoms and feeds on a wide variety of molluscs and crustaceans. It may enter brackish waters. It bears young from May to August. Similar Species: The giant butterfly ray, GyMiniira altavela, grows to a breadth of 7 feet or more. It is found from Massachusetts to Brazil. This ray has small spines on the tail base. It is known to utter a grunting sound when in distress. The butterfly sting ray, Gymnura marmorata, of Point Conception to Mazat- Ian, has a small sting and reaches a breadth of 5 feet. It is common in shallow bays along beaches. THE LOWER FISHES 173 Figs. 19 and 80. Spotted eagle ray (left) and cow-nosed ray Qright'). EAGLE RAYS: Family Myliobatidae These rays have largely abandoned bottom-living habits, but still feed on the bottom on hard-shelled molluscs and occasional crustaceans. The teeth are well adapted for crushing such resistant fare and are flat and like a pavement in the jaws. When feeding, thev swim near the bottom in order to detect, probably by smell, the currents emitted from clam siphons. Then they dig the clams by flapping their powerful pectorals or by rooting about like a hog with their noses. When not feeding, they swim gracefully and rapidly through the water near the surface and may even leap from the water completely. They rarely rest on the bottom. Because of their active habits, these rays are not as flat as most rays. There is a definite head with an elevated crown, which has the eyes and spiracles on the side rather than on top. The body is heavy and the pectorals powerful and thick. There are one or more poisonous spines at the tail base, but these are so near the body that they cannot be used as efficiently as those of sting rays. Nevertheless, the presence of spines as well as the large size of up to 8 feet across and 600 pounds in weight should be enough to caution the swimmer that he had best not tamper with these normally inoffensive fishes. They are very hard to handle and dangerous if hooked or speared. Eagle ravs travel over deep water but are most common in sandy or muddy bays where shellfish abound. They seem not to enter enclosed bays and estuaries as sting rays do. They are ovoviviparous and found in all warm temperate and tropical seas. SPOTTED EAGLE RAY (SPOTTED DUCK-BILL RAY, LEOPARD RAy) : Aetohatus narinari Size: Averages 3 to 4 feet across. Up to almost 8 feet across. Weight: Up to 500 pounds or more. Distribution: Almost cosmopolitan in warm seas. North to North Carolina (rarely the Chesapeake) and over much of the Indo-Pacific. Identification: The vivid light spots on the dark olive, brown, or black background form a distinctive pattern. The snout is almost like a duck's bill. Habits: These large and spectacular rays are almost exclusively mollusc-eaters. They travel singly, in pairs, or in large schools of hundreds of individuals. A 174 UNDERWATER GUIDE TO MARINE LIFE large school in an oyster bed can very well spell the end of almost the entire bed. Large rays are able to crack thick-shelled molluscs, an indication of the immense power of their jaws. But the jaws are delicate as well as powerful since the meat of the molluscs is separated from the shell so skillfully that very little shell is swallowed. These are powerful swimmers and are known to leap from the water. One specimen only a little over 5 feet across was easily able to drag a 22-foot boat containing three men. They are known to emit a harsh cough when captured. The young are released into the air one at a time by the mother as she leaps out of the water, a spectacular method of childbirth which is seldom observed. Similar S'pecies: The eagle ray, Myliohatus freyninvillei, is smaller than the spotted eagle ray, reaching 5 feet across, and maturing at 2 to 3 feet across. There are no spots, and the snout is not like a duck's bill. It strays to Cape Cod in summer, but lives the year round in the Caribbean. The bat sting ray, Myliohatis californiciis, is very similar to the eagle ray and is common inshore in bays from Oregon to Magdalena Bay. COW-NOSED ray: Kliinoftera qiiadriloha Size: Averages 3 feet across. Up to a width of 7 feet. Weight: Averages 25 to 70 pounds. Distribution: New England. Less common in tropical than in warm temperate zones of both hemispheres. Identification: The peculiar bilobed nose is distinctive. The color is a yellowish brown. Habits: This is the commonest of the eagle rays. Like the rest of them, it is a mollusc-eater. A school of these rays rooting about in clam or oyster beds is reminiscent of a drove of hogs. It is commonest in shoal water and may leap from the surface. The young are produced in spring and summer. Little is known of their habits in spite of their reasonable plentifulness. Cow-nosed rays have the disturbing habit of "rounding" a swimmer, but in doing so have no aggressive intentions. DEVILFISHES: Family Mobulidae These rays have completely abandoned the bottom-living habit and do not even feed there. Thus, the eyes are on the side of the head, and the spiracles are so reduced that respiration is sharklike, most water entering the gills through the mouth. Devilfishes are easily identified by the large cephalic fins that look like devil's horns on either side of the mouth. These are used to funnel and scoop plankton and small fishes into the ray's mouth. A gill sieve filters the food from the water much as in the whale and basking sharks. There are no poisonous tail spines, but the great size and famed power of these fishes should be enough to caution the unwary. Contrary to rumor, these are inoffensive fishes showing little fear of man. Rarely are we treated to a sane view of the manta, for instance. Magazines and movies would rather show them gushing blood from harpoon wounds or making alleged attacks on swimmers. If molested, however, one can hardly blame them for fighting back. The pectoral fins are used to deliver repeated sledge-hammerlike blows. Devilfishes are ovoviviparous and found in all temperate and tropical seas. THE LOWER FISHES 175 LITTLE devilfish: Mohulu hjfostoma Size: Up to 4 to 5 feet in width. Near relatives reach 23 feet in width. Weiolit: Up to 40 pounds. Near relatives reach 3,500 pounds. Distribution: Cape Hatteras to Brazil. Recorded north as far as New York. Identification: The mouth is on the underside beneath the short snout. The color is a blackish brown. The tail is relatively long. Habits: Little devilfish travel in schools and feed by rushing at schools of small fishes or plankton. They drive the prey toward shallow water near shore and may even beach themselves temporarily in their vigorous feeding rushes. They are inveterate leapers. According to Coles (1916) these rays leap clear of the water and land with a great splash. The larger manta also leaps, but usually not completely clear of the water. Females also leap during childbirth. The young are ejected one at a time as the female leaps into the air. If a female devilfish is harpooned, this will often cause a forcible ejection of the young. Little devilfish can utter a musical barking sound. Similar Species: The lesser devilfish, Mobida lucasana, reaches 7 feet in width and is found from Baja California to Central America. The manta, Manta hirostris, has a large mouth terminal on the snout and a shorter tail than the little devilfish. It ranges in all tropical and subtropical seas and is fairly common. It reaches 23 feet in breadth and 3,500 pounds. Contrary to the similar little devilfishes, it usually travels alone, moves slowly as it feeds, and leaps only partly out of the water. It still makes a tremendous noise in its leaps however. Tales of its power in pulling boats are legendary. It likes to bask Fig. 81. Little devilfish.. 176 UNDERWATER GUIDE TO MARINE LIFE in the sun with the upturned "horns" protruding from the water, at which times it is easily approached. The word "manta" means "blanket" in Spanish. GHIMAERAS: Glass Chondrichthyes, Subclass Holocephali These very odd fishes are mostly found in the deep sea and have the long, pointed tail that is characteristic of so many deep-sea fishes. The reason for this is unknown. Chimaeras do not seem to use the tail much for propulsion, this task mainly being the job of the flapping pectorals. Very commonly there are odd appendages on the head of the male which may be of some unknown reproduc- tive function. They have an operculum like bony fishes. Despite their preference for deep waters, chimaeras do get into shallow water. The long dorsal spine is to be avoided. It is poisonous and able to inflict a painful wound. CHIMAERAS: Family Chimaeridae RATFiSH (chimaera) : Hydrolagiis colliei Size: To 3 feet. Distribution: Northwest Alaska to Baja California. Fairly common in shallow water in northern California and northward. Identification: The color is silvery with gold, blue, and green iridescent sheens. Hahits: The egg cases are unique. Very little is known of the habits. Presum- ably, ratfishes eat invertebrates, as the chisellike front teeth and crushing back teeth would indicate. Similar Species: The Atlantic chimaera, Chimaera affinis, reaches 3 feet and is a deep-water fish of waters to the north of Cape Cod. E6S CASE Fig. 82. Ratfish. X^ CHAPTER ^ MASTERS OF THE WATER— Bony Fishes CLASS OSTEIGHTHYES {"bony fish") We have previously mentioned that the primary aims of this book are to give identification of groups (such as famihes) rather than of species and to see into which niche each group falls in the economy of the sea. Perhaps nowhere is the usefulness of this procedure better exemplified than in the case of bony fishes, for the following reasons: 1. Bony fishes are the dominant animals of the sea, receiving far more attention than any other of the sea's inhabitants. They are tremendously numerous and, in contrast to the sharks and rays, extremely diverse both in form and habits. To understand this great diversity, it is best to think not in terms of thousands of species, but in terms of families. 2. By keeping groups in mind, the swimmer will not be completely lost when confronted by strange, new fishes elsewhere in the world. For instance, both the red tai of Japan and Europe's huge dentex become species of porgies, Sparidae, rather than just a couple of unknown, isolated species. 3. The bony fishes are much more difficult to identify than sharks and rays, demanding the use of comparatively complex characters for proper identification. For instance, demoiselles, Pomacentridae, are not too diffi- cult to identify as a group, but species identification is complex and is still a matter of dispute among ichthyologists. It is difficult, if not impossible, to give a list of characters that strictly define bony fishes. All have skulls that are composed of a complex system of bones, but this information is of little use to the underwater swimmer. However, there are two field characters which vary little; the possession of an operculum which covers the gills (if absent, as in some eels, there is never more than one gill opening) and the presence of fins which are supported by bony rays. The fins are very mobile, in contrast to those of the cartilaginous fishes. Usually, also, there is a covering of bony scales which overlap like the shingles on a roof. The movement of fishes is a topic of great interest, to which the observations of the underwater swimmer can add much. The ffight characteristics of birds 177 178 UNDERWATER GUIDE TO MARINE LIFE SPINX DORSAL FIN LATERAL LINE . | SOFT DORSAL FIN KEEL SCUTES OPERCULUM \y y PECTORAL FIN t PELVIC, (ventral) fin THORACIC POSITION r CAUDAL (tail) fin PELVIC (VfNTRAL) FIN ABDOMINAL POSITION Fig. 83. External anatomy of hony fishes. through their medium, air, have been well studied, but fish movement is less understood, though they, too, fly, in a sense, through water. Bony fishes are more efficient swimmers than sharks and rays because of the combination of three factors: (1) the shortened, compact body allowing short, efficient, lateral strokes of the tail (a tendency seen in the mackerel sharks), (2) the possession of an internal hydrostatic organ, the swim bladder, which adjusts the specific gravity of the fish so that stabilization in any depth of water is possible (sharks, as we pointed out earlier, cannot do this), and (3) the great mobility of the fins, resulting in increased maneuverability, and the ability to start and stop suddenly (sharks and rays cannot do this). Fish movement is so intimately related to the habits of fishes and is so useful in identification of groups that we will give it considerable attention in our review of the families. Breder (1926) has set up three main categories of fish movement: 1. Carangiform (like the jack, family Carangidae). The typical fish movement in which propulsion is accomplished by short lateral strokes of the tail, and steering is accomplished largely by the paired fins. 2. Anguilliform (like the eel, family Anguillidae). The long-bodied fishes such as eels progress by a long series of sine waves which pass down the body. (Most sharks, being long-bodied, have a modified and shortened movement of this type.) 3. Ostraciform (like the trunkfishes, family Ostraciidae). Propulsion becomes largely a task of the pectoral, dorsal, and anal fins. The tail is used mainly for steering. Within these very broad categories many variations occur, and these shall be mentioned as they are met, particularly as departure from the typical COMr