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The Sea as a Biological Environment
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Classification of the Marine Environment

In order to facilitate a study of the marine environment and its inhabitants, the former may be conveniently divided broadly into primary and secondary biotic divisions based upon physical-chemical attributes or upon the nature of the biota. The boundaries between these biotic divisions, which are diagrammatically shown in fig. 67, are in some instances well defined, but more frequently there is a good deal of overlapping. Thus, although the primary divisions are definitely set off from each other on physical bases, and the typical subdivisions of these habitats can be clearly recognized both biotically and abiotically, yet there are no well-defined boundaries between them.


The main divisions of the marine environment.

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The two primary divisions of the sea are the benthic and the pelagic. The former includes all of the ocean floor, while the latter includes the whole mass of water.

The Benthic Biotic Environment and Its Subdivisions. This division includes all of the bottom terrain from the wave-washed shore line at flood-tide level to the greatest deeps. It supports a characteristic type of life that not only lives upon but contributes to and markedly modifies the character of the bottom. Ekman (1935) discusses the boundaries of the vertical zones from a zoogeographic standpoint, and we follow mainly the scheme employed in his text.


The benthic division may be subdivided into two main systems—namely, the littoral and the deep-sea systems. The dividing line between these has been set at a depth of about 200 m on the arbitrary supposition that this represents the approximate depth of water at the outer edge of the continental shelf (p. 20), and, roughly, also the depth separating the lighted from the dark portion of the sea. The littoral system is subdivided into the eulittoral and the sublittoral zones. The deep-sea system is divided into an upper (archibenthic) and a lower (abyssal-benthic) zone. The limits of the benthic subdivisions are hard to define, and are variously placed by different authors because uniform boundaries that will fit all requirements cannot be drawn. For general biological studies, the different boundaries must be based on the peculiarities of the endemic plant and animal distribution and should follow the region of most distinct faunal and floral change. The biotic zones thus delineated will be characterized by a more or less clearly defined range of external ecological factors which have given character to the population.

The eulittoral zone extends from the high-tide level to a depth of about 40 to 60 m. The lower border is set roughly at the lowest limit at which the more abundant attached plants can grow. The sublittoral zone extends from this level to a depth of about 200 m, or the edge of the continental shelf. The dividing line between these subdivisions varies greatly between extremes, since it is determined by penetration of light sufficient for photosynthesis. It will be relatively shallow in the higher latitudes and deep in the lower latitudes. In the upper part of the eulittoral zone a relatively well-defined tidal or intertidal zone that is bounded by the high- and low-water extremes of the tide is recognized. Some authors confine the eulittoral zone to this narrow section and consider the sublittoral to begin at the low-tide level (cf. Gislen, 1930). The vertical range of the intertidal zone, though rather well defined for any given area, varies greatly in different sections of the world, for it is determined by the tidal range (see chapter XIV). In the upper reaches of the Bay of Fundy the zone may have a vertical range of over 15 m, while in the Gulf of Mexico it is less than 0.7 m, and in areas like the Mediterranean along the southwest coast of Italy the range is yet smaller, only 10 to 30 cm. On exposed coasts subjected to direct ocean waves and swells the upper range is somewhat extended to include a rather well-defined supratidal spray zone with a sparse population of especially resistant forms among which a few animals, such as the isopod Ligyda, appear to be in the process of becoming terrestrial in habit. Many species of animals are found only in the tidal zone and may be limited vertically in maximum distribution even to certain levels within the zone—for example, Ligyda and the gastropods Littorina scutalata, L. planaxis, Acmaea digitalis, and others found at Monterey Bay only above the 0.76-m tidal level (Hewatt, 1937). Thus, in the tidal zone

in which the range in external factors is greatest we find a more restricted vertical range of specific animals than is obvious anywhere else in the benthic region of the sea. Many motile animals, for example crustaceans and fishes, move regularly into the intertidal zone to feed during high tide, and the small pelagic fish known as grunion migrate into the zone during certain high spring tides to deposit their eggs in the sand.

The eulittoral zone gives rise to many biotopes, for it is greatly varied as to type of substratum—for example, rocky, sandy, or muddy—and also as to character of shore line and degree of exposure. The overlying water may be slightly or greatly reduced in salinity. These variations are direct, decisive features controlling the type and abundance of sessile littoral forms (cf. Shelford et al, 1935). The plentiful primary food in this zone is derived from both pelagic and attached plants.

Attempts to establish such zones as Fucus zone, Laminarian zone, and so on, based on the depths at which these plants are characteristically attached, has the disadvantage that the plants are very frequently absent along vast stretches of the coast, owing to unfavorable substratum or other ecological factors; nevertheless, such classification may be of useful local application.

Though the boundary between the sublittoral and the deep-sea systems is set at a depth of 200 m, Ekman's compilations based on the fauna indicate that in most regions the boundary may be located between 200 and 400 m. Light and temperature are important factors, and in high latitudes these factors operate together to shift the boundary into shallower water.

The upper division of the deep-sea system is called the archibenthic, a word introduced by Alexander Agassiz, but the term is unfortunate in that it implies the beginning of the benthos from this region. The zone is also called the continental deep-sea zone, but this gives rise to greater confusion, since the term “continental fauna” sometimes used must include also the littoral fauna unless specifically called continental-slope or deep-sea fauna. The archibenthic zone extends from the sublittoral to a depth between 800 and 1100 m.

The abyssal-benthic zone comprises all of the deep-sea benthic system below the archibenthic zone. It is a region of relatively uniform conditions. Temperatures are uniformly low, from 5° to −1°C, and solar light is wanting. There are no seasons, and hence the seasonal biological phenomena associated with the littoral zone are suppressed. Stagnant conditions do not prevail in the open ocean, however, for there is ample circulation to supply well-aerated water resulting from deep vertical movements in the high latitudes (p. 138). No plants are produced, and the extent to which autotrophic bacteria play a part in the manufacture of food is not known. The animals are carnivorous, feeding mainly upon organic detritus which in its initial organic state must have originated

in the plants of the surface waters. The abyssal zone, though not sharply marked off at its upper limits from the archibenthic zone, has its own characteristic population, as will be brought out in a following chapter.

The benthic environment from shore seaward to abyssal depths is covered, to a greater or less degree, by sedimentary deposits that may be classified as terrigenous deposits, organic or pelagic oozes, and red clay. A detailed discussion of the deposits will be found in chapter XX, and the nature of the distribution is shown in fig. 253. As far as the biology of benthic animals is concerned, the most important features of these oozes are their physical consistencies and the amount of digestible organic material they contain. Most deep-sea benthic forms are detritus eaters and mainly dependent, therefore, upon the rain of pelagic organisms that falls to the bottom. The production of pelagic food usually decreases markedly with increasing distance from the coast, and the amount reaching the bottom in areas of very deep water is further reduced by its disintegration while sinking. Hence, the littoral muds are most rich in food, and the red clay at great depths and far from shore is the poorest. This difference is reflected in the number of animals actually collected from different areas (cf. p. 806).

The Pelagic Environment and Its Subdivisions. The pelagic division includes all of the ocean waters covering the benthic division. Horizontally, the pelagic division is subdivided into an open-sea (oceanic) province, and an inshore (neritic) province.

Vertically, the oceanic province has an upper lighted zone and a lower dark zone with no well-marked boundary between the two. For convenience the boundary is arbitrarily set at 200 m, since this would correspond with the arbitrarily set depth for the edge of the continental shelf and at the same time place the littoral system and the neritic province in areas definitely within the lighted portion. Actually, light changes gradually in both quantity and quality from the very surface downward to depths where it is no longer detectable (p. 82), and this depth varies with latitude, season, amount of suspended material, living or dead, and therefore also with distance from shore. These variables of the pelagic environment are of profound importance to the population of the sea, as will be pointed out later.

The outstanding features of the oceanic province are the broad spatial expanses and the great ranges of depth. As distinguished from the neritic province the waters are as a rule very transparent, with little or no detritus of terrestrial origin. These waters are predominantly blue in color and support the blue surface fauna to be discussed more fully in chapter XVII. Although solar light penetrates relatively deeper than in inshore waters, the great depth of the water included in this province results in complete elimination of solar light in the deeper portion

of the province, and as a result only carnivores and detritus feeders can exist in the very deep layers.

The chemical composition of the offshore water is relatively stable. Salinity is uniformly high, with only small fluctuations in space and time (p. 123), and plant nutrients are frequently relatively low in the upper layer and only slowly replaced.

The vertical border separating the neritic province from the oceanic is set at the edge of the continental shelf; hence all water of depths shallower than 200 m would fall within the neritic province, which accordingly may extend far seaward in instances where the continental shelf is broad, as off the east coast of the United States, or be very narrow, as off the west coast of South America.

Although biologically and chemically the border between the oceanic and the neritic provinces is not strictly definable, yet as we approach the coast the plant and animal life takes on characteristics not found in the typically oceanic province where “blue-sea” forms prevail. The chemical constituents of the sea water in the neritic province are more variable than in the oceanic. Salinities are usually lower, sometimes markedly, and undergo seasonal or sporadic fluctuations such that many of the inhabitants are more or less euryhaline in nature—that is, able to endure wide ranges of salinity. River water may bring in nutrients and may also exert a stabilizing influence on the turbulent motion, being at times, therefore, instrumental in initiating plant growth in the upper layers (p. 789). Plant nutrients, nitrates, phosphorus, and so on are more readily available in the shallower inshore water because of the greater possibility of return by vertical currents after they have been regenerated from the disintegrating organisms on the bottom or in the deeper water (chapter VII). This factor is of the utmost importance to production of diatoms, foremost of the primary food of the sea. Therefore, per unit area of the sea, the neritic province is far more productive than the oceanic province and is consequently the region of greatest importance to marine life in general. Here fish of greatest economic importance are taken, not only because of greater availability, but also because it is their natural habitat.

Other Biotic Units. The above classification of the marine environments is based mainly on broad geographical, physical, chemical, and biological characteristics that circumscribe more or less clearly the separate zones. Within each of these extensive zones we observe many and varied sets of ecological conditions resulting from differences in substratum, proximity to shore, depth and chemical-physical condition of the water, and so forth.

The primary “topographic” unit used in ecological classification of the environment is the biotope, or niche, which is defined as “an area of which the principal habitat conditions and the living forms which are

adapted to them are uniform” (Hesse, Allee, and Schmidt, 1937). Since in any given type of biotope the habitat conditions make specific demands on the inhabitants, it follows that an analogous development of the inhabitants is frequently reflected in the population, and those not fitted for the habitat are eliminated from it. Obviously, some organisms are not so narrowly bound to the biotope as are others of more specialized nature. Thus, within a biotope may be found some generalized forms such as certain cephalopods and fishes that wander more or less freely from one type of biotope to another. The more specialized a biotope becomes with respect to living conditions, the more uniform will the inhabitants become, so that only a few species with large numbers of individuals may exist. The smaller habitat anomalies found within the biotope are called facies. The number of organisms that can live in any given biotope may in special cases be determined by available suitable space, but more frequently it will depend upon the food supply that may be produced within the biotope or be carried to it from outside by currents. The community of forms in a biotope is called a biocoenosis.

Biotopes having certain characteristics in common—for example, proximity to the coast or estuarine locality—are united into larger divisions known as biochores.

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The Sea as a Biological Environment
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