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Populations of the Sea
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Many different species of distinct animal groups live intermingled in the same faunal area. Some may have identical habits and requirements, but for the most part the separate species or higher ranks have characteristic limitations, and each has its own function in conditioning the whole complex organic environment, thus influencing the type of species forming the association. This will become increasingly clear in considering the interrelations of the organisms (chapter XVIII).

For an adequate understanding of the intricacies of the fauna, it becomes necessary, therefore, to understand the part played by the separate species or groups of species, and, in order to circumscribe and to interpret the geographic and bathymetric distribution of species, their exact identity must be established. Systematic biology—that is, taxonomy—provides the tools for these purposes and is therefore an indispensable aid toward the desired goal. The field of study is so overwhelmingly large, however, that the many species comprising the primary groups must be investigated in special studies and the results of many different specialists must be integrated to provide a picture of the fauna as a whole. Much still remains to be done in this field, for many sections of the oceans have been only superficially investigated.

A full descriptive treatment of the animals of the sea would require several volumes, but it will suffice for our purpose to list or succinctly to review the primary divisions, including only a few of the secondary divisions that in the development of the study of marine biology and oceanography have assumed more or less importance and that are illustrative of the general field.

In the following synopsis, where the number of species is recorded for any group, the figures have been obtained mainly from Pratt (1935) and Hyman (1940). Illustrations are found mainly in chapter XVII.


Synopsis of the More Important Systematic Groups of Marine Animals



Protozoa are single-celled organisms microscopic or minute in size. The sea bottom harbors many creeping and attached protozoa of the ameboid or ciliate types, but we shall be concerned mainly with the pelagic forms inhabiting the plankton.


Order Dinoflagellata. In the broadest sense, this group contains both animals and plants, it being a borderline group.

Foremost among the protozoa in the economy of the sea are the dinoflagellates, chiefly because of the capacity of many types to carry on photosynthesis. These holophytic members are considered more fully in the discussion on plants, and for oceanographic studies are properly included in the phytoplankton. It will suffice to mention here only Noctiluca (fig. 225g) as an important representative of the holozoic members, none of which have chromatophores. The soft spherical body of Noctiluca is pale pink in color and bears a conspicuous flexible tentacle. The maximum size is only about 1.5 mm, but, when reproducing in profusion by simple cell division, the countless numbers produced may, by their accumulation, impart a pinkish-red color to considerable areas of surface coastal water, and the masses may be blown into conspicuous windrows or patches resembling “tomato soup.” Noctiluca are voracious feeders, engulfing particulate food such as diatoms and other small organisms. This form is also important as a contributor to the luminescence of the sea.


Order Foraminifera. The oceanographic interest of this order (and also, to some extent, of the following order) lies in the skeletal structures produced by its members. In the foraminifera the shells are variously formed, with one or more chambers arranged in a straight line or in a spiral (fig. 225a). Some are provided with many pores for the projection of protoplasmic pseudopodia used in capturing food. The shells are constructed typically of calcium carbonate, but silica and chitin are also used, and in some

benthic arenaceous forms they consist of an agglomeration of foreign materials cemented together. The greatest numbers of individuals are planktonic in life, but upon sinking to the bottom the shells form an important constituent of globigerina ooze, named for the abundant pelagic genus Globigerina (p. 816). Fossil foraminiferal shells are much used in the study of geological strata, being indices of past geological conditions, and are useful in the field of oil geology. A few foraminifera occur in fresh water, but the vast majority occur in the sea, either living on the bottom even at depths of 6000 m or floating freely in the water, preferring the warmer seas. There are over 1200 species, of which about 26 are pelagic. A recent catalogue of foraminifera (Ellis and Messina, 1940) includes some 18,000 living and extinct species.

Order Radiolaria. These are planktonic organisms whose skeletons are composed mainly of silica, but the Acantharia contain acanthin (strontium sulphate), and all types possess an inner capsule of chitin. The siliceous skeletons are formed in the most intricate and widely divergent patterns in the different species and are the most beautiful of all objects found in the sea (fig. 225e,f). Upon sinking and mingling with the bottom sediments, the skeletons become the type constituents of the siliceous radiolarian oozes found most abundantly covering the ocean floor in the deep tropical waters of the Pacific Ocean (fig. 253). There are about 4400 species, all marine.


Suborder Tintinnoinea. These protozoans, commonly called tintinnids, are mostly of extremely small size, varying from 20 μ for Tintinnopsis nana to 640 μ for Cymatocylis robusta. Swimming is accomplished by the beating of a whorl of hairlike cilia at the anterior end. Their loricae, or shells, range in shape from tubular to urn-shaped structures that are secreted in a stereotyped fashion by the animal and may or may not include agglomerated foreign material such as bits of sand, diatom shells, and coccoliths (fig. 225c,d). The tintinnids at times are found in vast numbers, especially in coastal water, where they are important feeders on the smallest plankton, the nannoplankton. Their sensitivity to small changes in environmental conditions makes them fluctuate in numbers with seasonal or other changes. There are 692 known species,

mainly marine. (Kofoid and Campbell, 1929.) Examples: Favella, Tintinnopsis, Tintinnus.


The sponges are multicellular animals, though of simple and loose organization, either with spicules of silica or calcium carbonate imbedded in their bodies for support or with fibrous skeletons made of the horny substance spongin, as in the common commercial sponge. Sponges are all benthic and nearly all marine, only one family occurring in fresh water. In the sea they are to be found in all parts and at all depths, the siliceous forms living largely in the deep sea. Sponges grow attached to the substratum and obtain their food by propelling water through tiny pores in the body wall and filtering out the microorganisms and detritus that may be present. There are about 2500 species, mostly marine.


Coelenterata are tubelike primitive forms with a continuous body wall surrounding a simple digestive cavity with but one opening encircled by tentacles used in capturing food. The group shows a remarkable degree of polymorphism; that is, a single species may present a variety of forms reducible either to the sessile polyp or the swimming medusoid type.

Class Hydrozoa. To this class belong the hydroids commonly found growing in little tufts on rocks and sea weeds along the coast. From these branching polyps are budded the small jellyfish or medusae such as Obelia (fig. 79). The Siphonophora, an order of this class, are characteristic of the open sea and are represented by the beautiful blue Velella (“by-the-wind sailer”) (fig. 226b) and Physalia (the “Portuguese man-of-war”), neither of which possesses a sessile stage. They are planktonic colonial medusae, exhibiting the maximum development of polymorphism of all animals. There are about 2700 species of hydrozoa.

Class Scyphozoa. To this class belong the larger medusae with eight notches in the margin of the bell. Here are included the giant jellyfishes, some of which may become 2 m in diameter. A much-suppressed sessile polyp stage is present in the group. The 200 species are entirely marine. Examples: Aurelia, Cyanea.

Class Anthozoa. To this class belong the sea anemones, corals, and alcyonarians. There is no medusoid stage, and many of the polyps are colonial; some, especially the corals, are notable for their precipitation of calcareous skeletal structures, which, through long periods of accumulation, are important in the

building up of coral reefs and similar formations. All 6100 known species of anthozoa are marine.


Ctenophora are small globular or flattened forms of jellylike consistency and with eight meridional rows of fused cilia used in swimming. Some possess a pair of trailing tentacles used in the capturing of food. The abundant globular species are commonly known as “comb jellies” or “sea walnuts” (fig. 226a). There are 80 species, all marine. Numerically important genera are Pleurobrachia and Beroë.


Platyhelminthes are flatworms, a large number of which are found in the sea, either free-living or parasitic.

Class Turbellaria. Nearly all of this class are free-living on the bottom under stones and in crevices, where they move about by means of cilia covering the body.

Class Nemertinea. These are ribbonlike worms sometimes considered as a separate phylum. The benthic species live among rocks, algae, mussels, and so on, or burrow in the bottom, where they capture small organisms by means of a long eversible proboscis. Extraordinary size variations occur, some species being only 5 mm long, while one, Lineus longissimus, may become 25 m in length when extended, and therefore is the longest of the invertebrates; however, its threadlike form contains but little bulk. Fifty-two planktonic species of nemerteans are known, some living at great depths—for example, Pelagonemertes. (Coe, 1926). The planktonic forms are modified, some with caudal and horizontal fins for swimming (fig. 228c). There are about 550 species of nemerteans, of which nearly all are marine.


The thread or round worms occur largely as parasites, but some are found in the plankton, and very large numbers occur in decaying organic detritus on the bottom. There are about 1500 species, many of which are nonmarine.


Class Rotatoria (Rotifera). These are tiny benthic or planktonic organisms provided with rings of cilia for swimming and for gathering food. Vast numbers may occur in the neritic plankton during the warmer seasons. There are about 1200 species of rotifers, of which most are fresh-water inhabitants.



These colonial animals, known as “sea mats” or “moss animals,” form flexible tufts or thin incrustations over the surface of solid objects both in intertidal and deep waters. Below low tide, many species form rigid, erect, latticed or branched colonies. The individual minute animals have calcareous protective skeletons and possess a ring of ciliated tentacles for gathering microscopic food. There are over 3000 species, about 35 of which are nonmarine.


Brachiopoda are ancient sessile animals superficially resembling bivalve molluscs, but the hinged calcareous or horny shells are dorsoventrally situated instead of laterally, as in the molluscs, and the animals gather their food by means of delicate ciliated arms attached within the shell. They grow permanently attached to rocks and shells, usually in the littoral zone below low tide. A few live in burrows. All are marine and all are very abundant as fossils in the Paleozoic and Mesozoic rocks. About 120 living and 3500 fossil species are known.


Phoronidea are wormlike animals, living in membranous tubes in the sand and collecting food by means of ciliated tentacles. There are about 12 marine species.


Chaetognatha include numerous but small (maxima about 75 mm long) holoplanktonic wormlike animals known as “arrow worms” or “glass worms.” They are highly transparent and provided with eyespots, a caudal fin and one or two pairs of lateral fins, and with strong chitinous jaws and teeth for capture of prey. They occur from the surface to great depths and are distributed far to sea in all latitudes. All 30 known species are marine. Sagitta (fig. 228a) is the most abundant genus.


Annelida are true worms with elongated bodies composed of a series of similar segments.

Order Polychaeta. These are marine worms of great abundance provided with many setae and typically with a variety of well-defined head structures such as eyes, tentacles, chitinous jaws, ciliated cirri, and so forth, which are modified in keeping with their habits of life and mode of feeding. They have a wide distribution horizontally and bathymetrically.

For the most part they are benthic, either creeping or burrowing, as in Nereis, Glycera, and Arenicola, or sedentary in calcareous or fibrous tubes, as in Spirorbis and Sebella. Tomopteris is pelagic (fig. 228b). There are in all over 3500 species, nearly all marine.

Order Oligochaeta. These are earthworms, of which only a very few are marine, living near shore.

Class Echiuroidea. These are fleshy marine worms with only one or two pairs of setae. They are unsegmented or indistinctly segmented in the adult. They live in burrows in the mud and sand of the littoral zone. There are about 20 species.


Arthropoda include animals with a segmented, chitinous exoskeleton and with jointed appendages, variously modified for locomotion, feeding, and other activities.

Class Crustacea. Entomostraca. This group, formerly considered a subclass, is of convenience in designating a large assemblage of small, primitive crustacea belonging to several subclasses and orders distinguished from the higher crustacea, or Malacostraca.

Suborder Cladocera. Only a few occur in the sea. Examples: Podon, Evadne, sometimes important in neritic plankton. Very numerous in fresh water.

Order Ostracoda. This order includes more than 2000 species, mostly marine, living in the plankton and on the bottom (fig. 227b).

Order Cirripedia. These are the barnacles which as adults have calcareous shells and live sessilely in all benthic habitats, especially coastal. Some grow attached to drifting objects or upon whales and other animals, or they may form special floats for suspension. There are about 500 species, all marine.

Order Copepoda. Though small in size (about 0.3 mm to 8 mm in length), the copepods bulk large in the animal substance of the sea, for they are by far the most abundant of all crustaceans and usually constitute about 70 per cent of the zooplankton. There are over 6000 species of copedods, found mostly in the sea, where some 750 species are planktonic and extremely numerous. Many others are benthic or parasitic. The three main suborders of free-living forms are Calanoida (fig. 227c), Cyclopoida (fig. 229d), and Harpacticoida (fig. 229a). The first two are mainly pelagic,

the last benthic. Like other Entomostraca and some Malacostraca, they gather food by means of fine bristles on certain appendages (p. 887).

subclass malacostraca. These are the large crustacea, mostly benthic, many with strong claws and biting mouth appendages.

Order Mysidacea. There are about 300 species, mostly marine, living on or near the bottom.

Order Cumacea. About 400 species of this order are known; nearly all are marine, benthic.

Order Euphausiacea. These are commonly known as “krill,” and in some regions are very abundant in the plankton and near or on the bottom. Some attain a length of about 50 mm, and may at times be the major constituent of the zooplankton. There are 85 known species, all marine. Examples: Euphausia, Meganyctiphanes (fig. 227a).

Order Amphipoda. There are about 3000 species, nearly all marine, in various habitats.

Order Isopoda. Over 3000 species are known; they are mostly marine, living on the bottom and on vegetation or burrowing in wood. Examples: Limnoria, Munnopsis (figs. 77 and 221).

Order Stomatopoda. This order contains about 200 species, all marine, benthic, most common in shallow water of lower latitudes.

Order Decapoda. Decapoda include crabs, lobsters, shrimps. They are widely distributed in both the pelagic and benthic regions. Most of the over 8000 species are marine.

Class Arachnoida. This class is well represented in the sea by a number of marine mites, over 400 species of sea spiders or pycnogonids, and 5 species of Limulus, the king crab. All are benthic.

Class Insecta. Only one insect is submarine during its whole life; a few others live on the foreshore or skip over the surface in search of food. Example: Halobates.


The molluscs are noted particularly for their construction of an infinite variety of calcareous shells encasing the body and for the structural modifications that have taken place in the soft parts known as the foot and the mantle. These modifications are associated with the method of locomotion and capture of food.


Class Amphineura. The chitons are all flat, benthic animals creeping with the aid of a broad, flat foot. There are about 630 species, all marine.

Class Scaphopoda. Tusk shells live in the bottom mud from shallow water to depths of over 5000 m. All 200 known species are marine.

Class Gastropoda. In most types there is a spiral shell, and the foot is used in creeping. In this and the preceding classes a rasplike radula is a characteristic food-gathering organ. Some gastropods are holoplanktonic and may be without shells. These are the marine pteropods and heteropods (about 90 species of each) with the foot modified for swimming (fig. 228d,f). The latter are especially characteristic of the oceanic waters of the lower latitudes. There are about 49,000 species in the class, mostly marine.

Class Pelecypoda. The clams, oysters, and mussels have a hatchet-shaped foot which in many is used for digging. All are benthic, usually sessile or burrowing in mud, rock, or wood. The soft parts are enclosed within hinged shells and the food is conveyed to the mouth by means of ciliary action setting up water currents, sometimes through long siphons. There are about 11,000 species, of which about four fifths are marine.

Class Cephalopoda. In the squids, devilfish, and so forth, the foot is divided to form arms used in capture of prey. In keeping with their active, predacious habits, the eyes are usually well developed, but blind deep-sea forms occur. In Nautilus and related forms there is a well-developed shell. Cephalopods are either benthic or pelagic, some living at great depths. The giant squid, Architeuthis princeps, having a body girth of nearly 1 m and attaining a total length of about 16 m, is the largest of all invertebrates. There are about 400 species, all marine.


Echinodermata are animals with calcareous plates forming a more or less rigid skeleton, or with scattered plates and spicules embedded in the body wall. Many are provided with spines. All are marine, and all but a few sea cucumbers are benthic.

Class Holothuroidea. The sea cucumbers are mainly benthic. only members of the order Pelagiothurida being planktonic. There are over 650 species, some living in abyssal regions.

Class Asteroidea. The sea stars are among the most conspicuous of shore animals, but they live also at very great depths. About 1100 species are known.


Class Ophuroidea. There are more than 1600 species of brittle stars, with a wide horizontal and bathymetric distribution.

Class Echinoidea. There are about 600 species of sea urchins and sand dollars, a few of which live in deep water.

Class Crinoidea. About 800 species of sea lilies and sea feathers are known, with the center of distribution in the East Indian waters, but they also occur in many other waters. The former live mainly in the deep sea and are anchored by long stalks. The latter occur mainly at shallower depths and are without stalks. The class is a vanishing remnant of a formerly abundant group that has left more than 2000 fossil species.


Chordata are animals which in some stage of their life have gill slits and a skeletal axis known as a notochord.

Subphylum Tunicata. These are primitive chordates; of about 700 species, all are marine.

Class Larvacea (Appendicularia). These are small planktonic forms, sometimes abundant. Examples: Oikopleura (fig. 228e), Fritillaria.

Class Ascidiacea. These are sessile ascidians such as Ciona and Culeolus.

Class Thaliacea. This class is made up of pelagic tunicates that float singly or in chains; they may be very abundant at the surface in the warmer waters. Examples: Salpa, Doliolum.

Other protochordates are the wormlike Enteropneusta and the fishlike Cephalochorda, both of which are found burrowing in mud and sand.


Subphylum Vertebrata. This group includes animals with vertebrae. All but the classes Aves and Mammalia are cold-blooded.

Class Cyclostomata. The hagfishes and lampreys are fishlike forms but without paired fins. They have a circular sucking mouth without jaws. The former are all marine, while the latter live both in the sea and in fresh water.

Class Elasmobranchii. These primitive fishes—the sharks, rays, and chimaeras with a cartilaginous endoskeleton—have paired fins and a lower jaw. In this group are many large forms such as the giant manta and the whale shark, the largest of all fishes, which becomes about 16 m long. Nearly all are marine.

Class Pisces. This class includes the true fishes, with a bony endoskeleton, paired fins, and an operculum covering the gills. They are characteristically streamlined for great swimming

speed, but a considerable variety of structural modifications occurs. Like the above class, they are mostly carnivorous and highly rapacious. Most fishes are marine, and some are benthic, but the majority are pelagic, living in both shallow and abyssal depths.

Class Reptilia. This class is represented in the sea by snakes and turtles. They breathe air and are therefore inhabitants of surface waters. The turtles frequent the shore to deposit their eggs on sandy beaches; the snakes bring forth living young and are therefore less dependent upon the shore. The sea snakes are found in the Indo-West Pacific and in tropical waters of America. They grow to a length of from 1 to 2 m or more and some are very poisonous. The sea turtles occur in tropical and subtropical seas. They have paddlelike limbs for swimming, and some grow to great size. The leathery turtle, for example, which is the largest of the class, may attain a weight of 1000 pounds.

Class Aves. A great number of birds are dependent upon the sea for food. Some of these frequent the land only for nesting and rearing of young. Typical examples are the albatrosses, petrels, cormorants, and auks.

Class Mammalia. These are warm-blooded, air-breathing animals with hair and mammary glands.

Order Carnivora. The marine members of this order are the sea otters and, to a lesser degree, the polar bears. The sea otters occur only in small numbers and only along the west coast of North America, where they were formerly hunted commercially to the very verge of extinction. Recently, under rigid protection, they have recuperated to an encouraging degree. The polar bears are confined to the Arctic region, usually on or near floating ice

Order Pinnipedia. Pinnipedia include seals and walruses, nearly all marine. The limbs are finlike, in adaptation to the aquatic existence. There are three families: (1) Otariidae include the eared seals, sea lions, and fur seals. Small external ears are present and the hind limbs can be rotated forward. (2) Phocidae are the hair seals without external ears and with hind limbs incapable of rotation forward. (3) Odobenidae include the walruses, with greatly elongated canine teeth in the upper jaw. They are confined to the Arctic.

Order Sirenia. Sirenia are heavy-bodied mammals with a flat tail and with forelimbs modified as paddles. Hind limbs are wanting. They live near shores in warm waters,

where they browse upon vegetation. They are not numerous. Examples: sea cows, manatees, and dugongs.

Order Cetacea. This order includes whales and dolphins, highly modified for aquatic life by a streamlined body and finlike forelimbs and tail. The hind limbs are wanting.

Suborder Mysticeti. These are the baleen, or whalebone, whales, with a series of long plates of baleen suspended in the mouth (fig. 76a). The frayed ends of these are used in screening out plankton food. Examples: fin whale, humpbacked whale, and blue whale. The last named is the largest of all animals, growing to a maximum length of about 34 m and weighing 294,000 pounds.


a, the blue whale—a whalebone whale; b, the sperm whale—a toothed whale.

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Suborder Odontoceti. Odontoceti are the toothed whales. This group includes (1) sperm whales with teeth only in the lower jaw (fig. 76b) and (2) the numerous dolphins and porpoises with teeth in both jaws.

Reproduction and Life Cycles in Marine Animals

In any comprehensive study of oceanography wherein biological activities are implicated or in the study of any population or individual species in relation to environmental factors, it is necessary to take into consideration the nature of the life cycles of the organisms involved. Only thus can the biological activities and the methods whereby the race is maintained through countless numbers of generations be fully understood. The utility of life-cycle studies in such practical fields as economic entomology, parasitology, and fisheries has been abundantly demonstrated. Through a knowledge of the methods of reproduction and through recognition of the various developmental stages of marine animals the investigator has at hand valuable means of aiding the interpretation

of the fluctuations that occur in a given population, of understanding the vertical or horizontal migrations, and of tracing the methods and routes of distribution, because all of these phenomena are closely bound to phases in the life cycle of the organisms.

It should be pointed out here that, in dealing with the propagation of any individual species in relation to its distribution, we must distinguish broadly between (1) reproductive distribution and (2) sterile distribution. Reproductive distribution is associated with areas where environmental conditions are favorable to maturation, spawning, and larval development. Such areas may be called areas of reproduction, or nursery areas. Sterile distribution is associated with areas in which the submature or adult individuals may live and some spawning may take place, but in which the eggs fail to hatch or the larvae do not survive, so that the area must be restocked periodically by invasion of postlarval stages produced elsewhere.

In a study of the life cycles of marine animals, one is impressed particularly with three facts: (1) the preponderance of animals which, though sessile, creeping, or burrowing in the adult stage, possess a free-swimming period during the early stages of life; (2) the enormous numbers of young that are produced by both pelagic and benthic animals; and (3) the fundamental similarity of the larvae of different invertebrate groups. We shall be concerned only with the first two.

In a superficial survey of populations, it is mainly the larger, more conspicuous adult animals that are seen, yet, from the standpoint of numbers, vastly more starfish, barnacles, clams, crabs, fish, and so on, are represented in the microscopic, feebly swimming larval stages than in the adult stages. Most of these larvae do not survive to assume the adult habit, but, instead, serve as nourishment for other organisms, swimming or sessile, or are in some manner destroyed through action of the physical or chemical environment.

Types of Reproduction. In reproduction, animals are either oviparous or viviparous. The oviparous forms deposit eggs that develop outside the mother's body, while in the viviparous forms the young are nourished by the mother and are born alive in a postembryonic state. An intermediate condition exists in the ovoviviparous forms, where the eggs are incubated and hatched within the body, as in certain sharks, perch, and blennies. The term larviparous is sometimes used to indicate that larval stages are born. An embryo derives its nourishment from the yolk of the egg or directly from the mother, whereas typically a larva is morphologically adapted with mouth and digestive tract for the purpose of seeking its own nourishment. Later it will be seen how important this fact is in the life of many marine animals.

By far the greater number of animals of the sea are oviparous, and it is among these that the extraordinarily large numbers of eggs are

produced. The number of eggs produced by the female of any species is associated, however, with the degree of parental care or other protection accorded the eggs and larvae following fertilization. The greater the care, the fewer the eggs produced. Most spawning consists of casting the eggs freely into the water, where they are fortuitously fertilized by spermatozoa that have also been extruded into the water. In these instances, enormous numbers of eggs are shed each breeding season. The following examples of the numbers of eggs produced by single individuals will illustrate the extraordinary fecundity that is attained:

American oyster 115,000,000 Pacific halibut 3,500,000
Sea hare (Tethys) 478,000,000 Cod 4,400,000
Teredo navalis, more than 2,000,000 Sunfish (Mola) 300,000,000

Parental care of eggs and larvae.

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It has long been recognized, however, that the exceedingly great number of eggs produced by some species is not directly correlated with the number of adults that are found. The large numbers of eggs and larvae produced are, instead, a measure of the tremendous toll paid by these species in order to assure survival of enough individuals to carry on the race.

In the marine population as a whole, very little parental protection is given to the offspring in the larval stages, and frequently even the eggs are given no care, yet hundreds of examples can be cited wherein varying degrees of protection are afforded the embryonic stages and sometimes the larvae as well. Many of the larger crustacea retain the developing eggs attached by secretions to hairlike structures on the abdominal appendages. Some annelids produce viscid secretions for attaching the eggs to setae or to the body wall, while others retain the young up to a well-developed larval stage in special brood pouches, as in Spirorbis (fig. 77). Many other invertebrates provide brood pouches—for example,

the isopod wood gribble, Limnoria, in which chitinous flaps cover the eggs and young. In many copepods the developing eggs are retained in membranous sacs (fig. 77). Among the vertebrates may be mentioned the male pipe fish, which carries the developing young in a special groove on the ventral side of his body. Other animals guard their eggs by hovering over them, as in the slipper shell, Crepidula, or the little six-rayed starfish, Leptasterias hexactis, which continues protection even beyond the larval stage. Many other examples of parental care are discussed by Wilson (1935).


Some types of egg cases for protection of eggs and larvae.

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It is clear that in these instances of parental protection the need for large numbers of eggs is somewhat diminished. Nevertheless, when a relatively long, helpless, pelagic stage follows the protected period of incubation, many larvae must still be produced. Thus, for instance, the blue crab, which, though protecting the eggs till the young are hatched, has pelagic larvae and is said to carry over two million eggs (Truitt, 1939). In contrast, Limnoria produces a maximum of only about-twenty-five eggs, but retains these within a pouch until the young are able to burrow into the wood where they were born. Thus they escape the hazardous pelagic life of larvae. In this animal the hazards of a swimming existence are met not by the very young but by submature specimens which by short migrations attempt to establish themselves in less crowded situations prior to breeding (Johnson, 1935). The fact that some animals produce more eggs than others and at the same time offer more parental care must indicate that factors operate to destroy more developing young in one than in the other.

Finally should be mentioned the common method of depositing eggs in masses or protective capsules of various types (fig. 78), thus diminishing loss from excessive dispersal and other hazards of a floating existence during the period of incubation. The capsules are also sometimes

watched over in the manner of the octopus, which keeps her eggs free from enemies. Additional examples are given in the discussion of life cycles.

Types of Development. There are two main types of development: (1) direct and (2) indirect. The direct development in oviparous species is associated with eggs of considerable yolk content, such as are found in the fishes, cephalopods, some nemerteans, crustaceans, and others. The newly hatched young are similar to the parent except for size.

Direct development is common among the deep-sea benthic animals, and this habit appears to be an advantageous adaptation. The slowly moving currents of great depths are of less importance in the dispersal of larvae than are the stronger currents of shallow water. The micro-planktonic life so characteristic of surface layers and from which the pelagic larvae of littoral animals directly derive their food has no counter-part in the deep, and hence it is imperative that the young produced be able to feed directly upon the bottom detritus. The possibility of deep-sea larvae swimming from great depths to the surface, where food is plentiful, and later returning to the bottom appears to be impracticable in nature. The young of some mid-depth pelagic forms—for example, Cyclothone among the fishes and Acanthephyra among the prawns—do however live nearer the surface, where food is more plentiful, than where the adults are commonly to be found (Hjort, 1912).

Benthic animals of Arctic and Antarctic regions also commonly possess no pelagic larval stages. Hjort (1912) and Murray (1913) consider this a probable explanation of the great local concentrations of certain boreal and arctic benthic animals, because the direct development results in the young remaining in the area in which they are born. Brief pelagic larval stages following protection during incubation and absence of dispersing currents lead also to local adult concentrations.

The indirect development is associated with a type of egg with little yolk (that is, alecithal), and hence a self-sustaining larva must develop quickly or the organism dies. This type of development is characteristic of marine invertebrates, which usually cast their eggs free in the water or carry them through the incubative period in special brood pouches. Larval stages appear before the full character of the species to which they belong becomes established. Many of these—for example, the pluteus larvae of the Echinoidea and the Ophuroidea—when first discovered were described as distinct kinds of animal, only to be found later to be the young of already well-known species. The locomotor organs of most of the larvae are cilia (see below for exceptions) which by their rhythmic beating propel the animal slowly through the water at a rate just sufficient to keep them in suspension. The great similarity of structure exhibited by the larvae of some groups suggests a common origin for the groups which as adults are structurally very dissimilar.


Typical Life Cycles. The life cycles of many species have not been investigated, but the principal features in the life history of the major groups have been established. We shall here review only the groups of most immediate interest in general oceanographic studies.

In the protozoa, reproduction is mainly by binary fission, whereby the animals divide to form two separate animals, these in turn dividing after growth. Under favorable conditions, this method makes possible a production of great masses of individuals, as is often witnessed in such forms as Noctiluca. Gametes are also formed in this animal as a result of multiple fission. These unite in pairs, but their further development is unknown. In foraminifera, and possibly also in radiolaria, there is a cyclical alternation of generations in which sexual and asexual phases alternate and give rise to morphologically different individuals (Myers, 1936).

In the tintinnids, in which transverse binary fission occurs, the anterior daughter escapes from the lorica, while the posterior daughter retains the old lorica (Kofoid, 1930).

The sponges reproduce asexually by budding or fragmentation, and sexually by union of gametes, the latter resulting in a free-swimming, flagellated larva, the amphiblastula, which, after a period of swimming, settles to the bottom and grows to form the adult sponge. Asexually produced units, known as gemmules, possessing a heavy protective covering are produced by some sponges as a means of survival during adverse periods. Reproduction by formation of gemmules occurs principally among the fresh-water sponges, but some marine forms also produce gemmules.

In the coelenterates, both sexual and asexual reproduction are important features in the life cycle. The union of germ cells results in a free-swimming, ciliated, planula larva about 1 mm long (fig. 80c). The planulae, though lacking the mouth and enteron of typical larvae, may live for a sufficiently long period on yolk food to bring about dispersal of sessile coelenterates such as the corals and anemones. Vaughan (1919) found the pelagic period of corals to be from one day to two or three weeks. Upon settling to a hard bottom the planulae of corals and other Anthozoa develop a mouth and tentacles for feeding, and later the reproductive organs are formed. In some there is also active asexual reproduction by fission and budding. Large coral colonies are thus initiated from a single individual. It is the skeletons of these asexually produced individuals which form the large coral heads, some of which are 3 m or more in diameter and contain many thousands of individual polyps. The length of time required for formation of such colonies has been investigated by Vaughan (1919), who found that a coral colony (Porites asteroides) 50 mm in diameter may be formed in four years.


The remarkable life histories of many jellyfish of the class Hydrozoa offer the examples of alternation of generations that are used in all zoological texts. The jellyfish, or medusa, stage (fig. 79) of such forms as Obelia is either male or female, and the eggs are cast free in the water, where they are fertilized and develop into planula larvae. The planula soon settles on the bottom to form the sessile polyp, or hydroid, stage. From special structures on the polyp, asexually produced buds become separated as swimming medusae, thus completing the cycle. The alternation of sessile and pelagic generations is a factor of great significance in the distribution not only of the hydrozoa such as Obelia, but also of other forms—for instance, the large scyphozoan Aurelia, which, though varying in details of life history, possesses stages similar to those in Obelia. The sessile generation is the chief link instrumental in restricting all stages of such animals to the neritic waters, generally to the proximity of shores and shoals with a suitable substratum of rocks, shells, or larger plants for attachment of the planula larvae. Bigelow (1938) found that in the open sea off Bermuda only about 3 per cent of the medusae caught at a distance of 10 miles from shore were of the type with a fixed stage in their life history. The degree of dispersal is, of course, dependent upon the speed and direction of the water currents prevailing. Some swimming jellyfish—for example, Aglantha digitalis and other members of the order Trachylina—are not dependent upon a sessile stage because daughter medusae develop directly from the pelagic stages.


The life cycle of a typical hydrozoan jellyfish, Obelia.

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Open-sea colonial coelenterates—for example, Velella or Physalia—are representatives of the “blue-sea fauna.” Their life cycle is adapted to offshore life by elimination of the sessile stage. The planula larva gives rise to a medusiform stage from which the complicated colony arises.

The Ctenophores are all hermaphroditic, and the eggs are usually shed into the water, where, upon fertilization, they grow by direct development into free-swimming larvae. Gastrodes, a parasite in Salpa, produces a typical planula larva.

The great importance of annelids, especially in the littoral benthic fauna, warrants their inclusion in this brief study of life histories. At certain seasons the voracious swimming larvae of annelids are a major

constituent of the temporary plankton and a factor to be considered in the control of phytoplankton production in neritic waters at these times (p. 772). Most marine annelids are unisexual, in contrast to the hermaphroditic terrestrial forms. Both sexual and asexual reproduction occur, but, when great numbers of pelagic larvae are found in the plankton, they result from the shedding of many eggs free in the water, where they are fertilized and develop into ciliated, swimming, trochophore larvae (fig. 80b), which are soon transformed into miniature worms with three or more segments before they desert the plankton (fig. 224a) There are many modifications of the trochophore, its fundamental structure being reflected in the larvae of a number of animals, especially the molluscs, the nemerteans, and other flatworms.


Some characteristic marine larvae. a, nauplius larva of the copepod Labidocera; b, trochophore larva of the annelid Nereis agassizi; c, planula larva of coelenterates; d, zoea larva of the crab Pachygrapsus; e, veliger larva of the clam; f, bipinnaria larva of starfish; g, the cod larva with yolk sac.

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The larvae of many benthic annelids enter the plankton only after they have completed their early stages under some means of special protection. For example, in the little tube worm, Spirorbis, and related forms, they develop in a special brood pouch beneath the operculum, while in some Polynoe, or scale worms, they are sheltered by the dorsal, flaplike elytra, and in yet other instances the eggs are deposited in attached or demersal gelatinoid masses (fig. 78), where the developing embryos and larvae enjoy some degree of protection. In Spirorbis the trochophore stage is passed in the brood pouch and the older larvae may assume the sessile habit after only twenty-four to thirty-six hours in the plankton. If a suitable substratum is not available, the pelagic stage is somewhat prolonged. Nereis agassizi spawns the eggs free

in the water, and the pelagic stage may be of twelve to sixteen or more days' duration. In N. vexillosa the eggs are spawned in masses (fig. 78).

The chaetognaths, or arrow worms, are hermaphroditic but not self-fertilizing. The eggs, which are fertilized internally, are shed into the water, where they develop directly into free-swimming larvae not unlike the adults.

Most crustaceans, in which we are particularly interested because of their prominence in some phases of oceanographic studies, pass through several distinct pelagic larval stages. The common initial crustacean larva is the nauplius (fig. 80a), bearing three pairs of appendages used for both swimming and feeding.

In copepods the sexes are separate, in some species the ratio of adult males to females being strikingly unequal at all times, while in other species the inequalities are seasonal, the males being more abundant at the onset of breeding but later diminishing in numbers more rapidly than the females after the breeding season (Damas, 1905, Farran, 1927, Campbell, 1934). The most important of the planktonic copepods spawn the fertilized eggs free in the water, yet many littoral species and some important pelagic species—for instance, Oithona, Paraeuchaeta, and others—carry the eggs in brood sacs through the period of incubation (fig. 77). In both cases the eggs hatch to typical self-sustaining nauplii. Paraeuchaeta is somewhat of an exception, for it develops from a heavily yolked egg and does not feed in the naupliar stage (Nicholls, 1934). In copepods there are normally six successive naupliar stages separated by definite moulting of the chitinous skin. The hard exoskeleton of crustaceans does not grow, and must therefore be shed or moulted periodically as the animal becomes too large for the encasement. In many crustaceans the number of moults may be variable, but in copepods there are a fixed number of stages, each separated by a moult. At the termination of the sixth naupliar stage a complete metamorphosis occurs from which emerges Stage I of six successive copepodid (copepodite) stages. Copepodid Stage VI is the adult, and, during spring reproduction in waters of the latitude of the British Isles, maturity may be reached in a period of about twenty-eight days in Calanus finmarchicus, but it is much delayed in the autumn-winter generation or in populations of more northern waters.

In Calanus finmarchicus, by far the most thoroughly investigated of all pelagic copepods, it has long been known (Gran, 1902, and others) that the animals spend the winter months in the deeper water layers. The breeding of this species occurs in spring and summer in boreal waters, and there are two or more successive generations, each of which, apparently, may bear more than one brood. The generation arising from the first spring spawning appears to mature quickly, spawn, and die. The last generation produced in autumn is a relatively long-lived

one, because it is this generation which carries the stock over the winter period. The winter stock is found in relatively deep water and is of uniform composition, consisting of copepodids in Stage IV or V. The stock is much reduced during the winter, but with the return of spring the animals pass into Stage VI (the adult stage) and spawn in surface waters, to produce the first generation of the season. Figure 81, from Nicholls (1933), shows changes in the percentage composition of the population, indicating three main periods of spawning in the Clyde Sea area.

The life cycle of Calanus finmarchicus appears to be quite characteristic of other members of that important genus and perhaps of other related genera as well, but very few pelagic copepods have been adequately investigated, and considerable variation can be expected. We shall not enter into the remarkable life histories of the parasitic copepods, but an example of a typically free-living littoral form—that is, Tisbe furcata—is instructive for comparison with Calanus. Tisbe furcata carries the eggs through the incubative period in brood sacs. There are six naupliar and six copepodid stages, as in Calanus but the time required for development from egg to mating maturity may be as little as ten days, and filled egg sacs are carried by females of the new generation in fourteen days after hatching. During maximum production, one female may produce at least seven or eight broods at about five- to eight-day intervals.


Successive generations of Calanus finmarchicus (from Nicholls).

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In the euphasiids, another group of outstanding importance in the economy of the sea, the method of reproduction is not unlike that occurring in many copepods, in that usually the eggs are shed in the water, but the succession of generations is not rapid and the life span is of greater length. For Euphausia superba, in Antarctic waters, the time required to reach sexual maturity is estimated by Ruud (1932) to be two years. Some investigations indicate that normally the animals live in the immediate vicinity of the bottom and that during spawning they congregate in swarms and ascend to deposit the eggs in surface-water layers. Here, while slowly sinking, the eggs hatch to typical naupliar larvae, which are followed by successive stages of distinctive larvae, the older of which may return for a time to the surface layers. In

Thysanoessa inermis the stages are, in order, two nauplius, one metanauplius, three calyptopis, fourteen furcilia, twelve cyrtopia, and the adult (Lebour, 1926). Some species—for example, Nyctiphanes couchii—carry the eggs in a brood pouch through the incubative and naupliar periods.

Among most other crustaceans the eggs are carried through incubation attached to appendages or in brood pouches of various types. In heavily yolked eggs, as in the common crab and related forms, the nauplius stage is passed within the egg, and the developmental stage emerging from the egg is known as a zoea larva (fig. 80d), of which there may be several separate stages. The weakly swimming zoea may drift in the plankton for several weeks before changing to the megalopa stage and settling on the bottom. The lobster produces a special type of pelagic larva. the phyllosoma, which, with its leaflike body, is especially adapted to float in the plankton (fig. 229g). In barnacles, the larvae escaping from the mantle cavity within the shell of the sessile adult are typical nauplii which, after living a pelagic existence for a few weeks, are transformed into what are called cypris larvae (fig. 2241). The cypris larvae soon settle to the bottom and, upon attachment to a solid surface, metamorphose to assume the adult state.

In most bivalve (pelecypod) molluscs the sexes are separate, although some are hermaphroditic, and others—for example, certain species of oysters—are unisexual but change their sex alternately from one to the other. This phenomenon is also observed in other marine animals (Coe, 1940). Fertilization of the eggs frequently takes place after the eggs have been shed into the water, but commonly the eggs are retained within the brood pouch formed by the gills, in which case the spermatozoa are taken in through the inhalant siphon with the stream of water that is kept flowing over the gills. A modified trochophore larva is first formed, and from this a later larva, the veliger, results (fig. 80e). After a period of swimming the veliger settles to the bottom.

Gastropods are often hermaphroditic. Fertilization among them is commonly internal, and their eggs are frequently deposited in gelatinous or membranous cases attached to rocks or sea weed. Tiny floating cases containing several eggs are sometimes formed, as in Littorina (fig. 78). The trochophore and part of the veliger stage are passed within the egg case.

Among the echinoderms the sexes are separate and the eggs are usually spawned into the water, where fertilization occurs. Some (Asterina) lay demersal eggs which, because of their viscid nature, adhere to rocks and other objects. In other echinoderms, especially deep-sea and polar species, the eggs are fertilized and retained in brood pouches, where they undergo early development. Development is indirect in all cases, but the forms with most heavily yolked eggs do not have pelagic

larvae. Most echinoderms, however, do possess pelagic larval stages that may be of several weeks' duration.

The metamorphosis resulting in the adult state is as complete as that experienced by the butterfly, and it is not surprising that, before the parentage of the pelagic larvae was known, they were considered as distinct animals unrelated to the adult (figs. 80f and 224f and j). The characteristic larvae are bipinnaria (sea stars), echinopluteus (sea urchins), ophiopluteus (brittle stars), auricularia (sea cucumbers). The echinoderm larvae are of only moderate interest in the economy of the sea, but great biological interest is attached to the probable significance of some in showing a relationship to the most primitive chordates.

Many fishes—for example the cod, mackerel, halibut, and sardine—shed their eggs into the water, where fertilization takes place and the developing larvae are nourished by the yolk of the floating eggs (fig. 80g). The herring deposits viscid eggs which, upon sinking to the bottom in shallow water, become attached to solid objects. The gobies, blennies, sculpins, and others attach their eggs to solid objects or lay them on the bottom, where the male may stand guard over them until hatched. The grunion buries its eggs in the sand of wave-washed beaches during periods of high spring tide. Here the eggs remain for a period of about two weeks, when the next series of high tides washes them out and stimulates their hatching (Thompson, 1919, Clark, 1925). In sharks and rays, fertilization is internal, and either the young are born alive or the nonbuoyant eggs are deposited in leathery cases known as “mermaids, purses” (fig. 78, p. 317).

The eggs of fishes fall roughly into two groups, depending upon buoyancy: (1) pelagic and (2) demersal. The demersal eggs sink to the bottom or are deposited there; pelagic eggs float freely in the water and hence greater numbers are produced to overcome the losses inherent with this group. Many fisheries investigations are concerned with the occurrence and dispersal of pelagic eggs and the resulting larvae, for from such studies much information is gleaned regarding the spawning habits and areas of many commercially important fishes (p. 861). In general, the development of fishes can be considered as being direct, there being no general metamorphosis of form to the adult morphology. There is frequently, nonetheless, a marked degree of dissimilarity between larvae and adult, and in some there is a distinct metamorphosis. The Leptocephalus larva of the eel, for instance, was once considered a separate species. Many fishes have definite spawning grounds far removed from their feeding habitat, and the remarkable migrations of such fishes as the eel and the salmon are directly associated with reproductive instincts (pp. 811 and 861).

The reproductive habits of certain deep-sea fishes are of special interest as an indication of adaptation to the environment. In the lightless,

sparsely populated, abyssal depths it is conceivable that individuals of the sexes may become separated to such a degree that fertilization of eggs at spawning becomes highly fortuitous. In some deep-sea fishes this condition is overcome by the male becoming parasitic on the female, being permanently and securely grown to her body as a mere appendage with a united circulatory system (fig. 231c, p. 831).

The mammals of the sea bring forth living young, which are nursed for a period by the mother. Like those of some fishes, the great migratory movements of whales and seals are associated with wanderings to and from favorable breeding grounds. Growth in whales is extremely rapid; sexual maturity may be reached in two years, and one calf may be produced every other year.

The spawning of many marine animals, especially in boreal waters, is of a spontaneous nature, and vast numbers of individuals spawn within a period of a few days, with the result that in such cases the main spawning period is easily ascertained, since great swarms of eggs or larvae appear suddenly in the plankton and are gradually dispersed by water movements. This feature is especially well illustrated by the oyster, certain sea cucumbers (Cucumaria), nereid worms, and barnacles.

The degree of success of (1) spawning or (2) survival of larvae of successive spawning seasons gives rise to an inequality in numerical strength of year classes of adult or juvenile forms constituting any given population. This inequality is best demonstrated by studies of commercial fishes, investigations of which have been most ardently pursued. However, the same inequality must also occur in the populations of any animals with a normal life span sufficiently long for individuals to live through several reproductive seasons as juveniles and adults.

For purposes of illustration we may consider a species with a life span of several years in which the age of individuals can be accurately determined and in which adequately large and inclusive samples are obtainable for comparison. Now, assume a highly successful spawning and larval survival in a moderate population of this species in the breeding season of 1930, a very poor spawning season in 1931, an average degree of spawning and survival of larvae in 1932, and then another highly successful year in 1933. The 1930 year class will, upon investigation of the whole population in 1931, show up as a disproportionately great number of small, one-year-old individuals in relation to the other age groups in the population. In the next year (1932) the two-year-old individuals of the 1930 spawning are still conspicuous in the population, but the smaller number of one-year-old individuals is evidence of a poor spawning or survival for the 1931 reproductive season. Thus, in 1933 and subsequent years the downward trend of numerical strength of the 1930 and 1931 classes can be traced and compared with other year classes—for example, that of the average year 1932 and of the successful year 1933

(fig. 82). As indicated in fig. 82, the 1930 and 1933 spawning produced “dominant year classes.”

From such comparative studies of year classes and with a knowledge of the spawning habits and age groups, means are provided for analysis of probable environmental factors that determine the degree of success of spawning or survival of larvae, because the relative number of individuals entering into any year class must depend mainly on these critical periods. In subsequent years within the normal life span of the species, the reduction of numbers in year classes is not so likely to be of catastrophic nature. It has been pointed out by Hjort et al (1933) that for a given region the average rate of growth of individuals within the different year classes of Norwegian herring is the same for each year class regardless of the relative numerical strength of the classes. This seems to indicate that, in the sea, nature each year provides a sufficiency of food for the survival and growth of the older stages of this fish as represented in the composite commercial catch. The numbers of individuals belonging to the separate year classes may be widely different (as much as 1 to 30), and this difference must then result from some factor or multiplicity of factors operating to destroy the animals during the very early stages of their existence.


Schematic illustration of changes in year class composition of a population.

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Studies of commercially important fish, shellfish, and whales are deeply concerned with analysis of the year classes. For example, in nature there is an equilibrium between the rate at which fish enter the accumulated stock or supply and the rate at which they are withdrawn through natural mortality. Additional removal through fishing exploitation disturbs this equilibrium and may constitute so serious a drain upon the stock that the drain becomes greater than the rate of replenishment and the stock subsequently becomes so depleted that it is no longer

profitably fished. Intensive fishing is evidenced by a decline in the proportion of older and larger specimens entering into the catch and also by a leveling off of abundant year classes. Therefore a record of the trend of the proportion of year classes gives valuable information regarding the toll that fisheries operations are exacting on the accumulated stock in the area being exploited. Such information provides practical aid in formulating conservation policies and in determining the optimum catch practicable to the fishing industry. The theories of fisheries science here involved are of great importance to that branch of marine biology. For a fuller discussion the reader should refer to Hjort et al (1933), Thompson (1937), and other relevant reports.

Year-class analyses of the fish population are also utilized as a basis in arriving at forecasts of the most probable abundance of fish in the next year's catch. Illustrative of this are the investigations of the U. S. Bureau of Fisheries into the fluctuations so characteristic of commercial catches of mackerel. Sette (1931et seq) studied the relative abundance of these fish caught with reference to each year class of the population. The numerical strength of the younger year classes entering into the catch provided a basis for calculating the probable yields that these classes would give in the following year under similar conditions of fishing. Such calculations can be significant only when the downward trend in numbers of the dominant year classes is rather regular. In dealing with migratory fishes, the occurrence of sporadic invasions of populations produced elsewhere or with a different range and whose year-class composition is not known must give rise to unexpected changes in the ratio of the year classes occurring in any one range or locality under investigation. The continued success of such commercial fisheries as the mackerel is determined mainly by the numerical strength of the dominant year classes of the population native to the fishing area. In the above investigation it was found that the big year class of 1923 constituted the main bulk of a declining fisheries yield for a period of years until the industry again experienced a sharp upward incline as the contingents of the 1928 successful spawning entered the catch.

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