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    Seashore
    Seashore
    Seashore
    Ebook645 pages12 hours

    Seashore

    By Peter Hayward

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    A comprehensive, authoritative account of the natural history of the seashore, from earliest times to the present day. This edition is exclusive to newnaturalists.com

    The seashore, with its endlessly changing tides, is one of the most fluctuating physical environments on the planet. Home to an abundance of animal and plant life, it is also one of the richest habitats the naturalist can explore. Here in Britain, we are fortunate to have a long and varied coastline, and our relatively large tidal ranges mean that our seashore offers a wide range of coastal habitats, including mud, sand, shingle and rock. In New Naturalist Seashore, Peter Hayward looks at:

    • Resident and migrant species, including fish, barnacles, limpets, winkles, sponges, algae, lichens and sea grasses
    • The effects of tourism and pollution on these habitats
    • The geology of the British Isles, with its sinking and rising coastlines
    • The responses and adaptations of plant and animal life to a changing physical environment

    This narrow strip of beach between the land and the sea that we call the seashore, has always attracted man, in the early years as a source of food, and in Victorian times as a rich habitat that the early naturalists would explore. In this fascinating addition to the highly regarded New Naturalist series, Peter Hayward brings the natural history of the seashore right up to date.

    LanguageEnglish
    PublisherHarperCollins UK
    Release dateAug 19, 2010
    ISBN9780007406043
    Seashore
    Author

    Peter Hayward

    Peter J. Hayward is Senior Lecturer in marine biology at the University of Wales Swansea. He is editor, co-author or author of 13 books on marine biology, including the Handbook of the Marine Fauna of North-West Europe and Collins Pocket Guide to the Sea Shore of Britain and Northern Europe. He has published around 100 papers on the marine Bryozoa, which are his particular research interest. He is zoological editor of the Journal of Natural History, and a frequent contributor to BBC Wildlife.

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      Seashore - Peter Hayward

      1

      Introducing the Seashore

      The seashore can be defined as a strip of land forming the frontier between land and sea. It is not a static frontier, but one that retreats and advances twice daily, under the tide-generating influence of the moon and the sun. The upper limit of the tide, the high-water mark, can be regarded as the shoreline; between it and the extreme lowest level of the tide lies the seashore, a transitional zone between two major ecosystems. It is neither land nor sea, but its environment is modulated by physical factors particular to each, and by others unique to this border zone. It is an exciting place to visit, and to follow a retreating tide as it reveals the sea’s fringes and the astonishing diversity of living organisms found there.

      The British Isles, from Shetland to Scilly, are distributed across 1,250 kilometres of the temperate northeast Atlantic, or 11 degrees of latitude. From eastern England to western Ireland they range 900 kilometres, or 13 degrees of longitude. Nowhere in Britain or Ireland is more than 120 kilometres from a shoreline, whether it be along an open oceanic coast, a deep, sheltered sea loch, or a broad estuary. The coasts of Britain are long and deeply dissected so that the proportion of shoreline to land area is exceptionally high, in northwest Europe probably only exceeded by that of Norway. There is no accurate computation of the length of Britain’s coastline, but for England, Wales and Scotland, and including that of all the Scottish islands, it has been estimated at more than 14,000 kilometres. The extent of seashore uncovered by a retreating tide is dependent upon coastal topography and the vertical range of the tide, both of which vary greatly around Britain’s coasts. Cliff-bound coasts with a moderate tidal range of just 3 or 4 metres will be bordered by only a narrow strip of seashore, while lowland areas adjacent to broad, sheltered inlets usually experience tidal ranges of more than double that magnitude and the sea may retreat for several kilometres at each low tide. The total area of seashore fringing all the islands of Britain has probably never been estimated, but it must be reckoned in terms of hundreds of thousands of hectares. It is not always appreciated how fortunate these islands are in possessing this immense area of natural habitat. Above the limit of the tide lies a supralittoral zone, of varying width, populated by terrestrial plants and animals especially adapted to maritime living, together with a few vagrants from the sea adapted for life among or just above shoreline debris. Along tideless coastlines these maritime communities are the only evidence of the sea’s proximity. There is no seashore, and while the shallower coastal waters may be biologically rich they lack those animals and plants specialised for intertidal living, which may be overwhelmingly abundant on tidal coasts. Britain’s geographical situation, on the northwest edge of the European landmass, fringing the Atlantic Ocean, and its convoluted, varied coastlines, together provide the conditions for maximum modification of tidal currents, resulting in local tidal ranges of an often impressive magnitude, and correspondingly extensive stretches of seashore.

      Together with its remarkably long coastline, Britain is unusual in its great range of geology and topography. Every age and almost every major kind of rock occurs here, from the unconsolidated Pleistocene gravels of southeast England to the hard Lewisian gneisses of northwest Scotland, which are among the oldest rocks in the world. In general the age and hardness of the basement rock increase along a southeast–northwest gradient. The coasts of East Anglia and north Kent are largely formed of Caenozoic sediments and glacial deposits. These are enfolded by progressively harder Cretaceous and Jurassic rocks that outcrop as chalk, limestone and shale cliffs along the Channel coast, and northwards from the Wash. The even harder Devonian and Carboniferous rocks form the cliffscapes of much of Devon, Cornwall, South Wales, and southern and northeastern Scotland. Highland Scotland largely consists of Precambrian, metamorphic or igneous rocks, which form some of the most spectacular coastlines in Britain. Rock type is the first and major determinant of coastal topography. The hardness of the rock, its chemical composition and physical structure, the degree to which it is folded or faulted, all affect the rate at which it erodes. Soft shales, clays and loose gravels crumble readily before the onslaught of waves, and generally result in low profile, rapidly retreating cliffs. In chalks and limestone, erosion is hastened by chemical solution along bedding planes; chalk tends to collapse precipitately, often giving near vertical cliffs, while harder Carboniferous limestones resist physical processes of erosion longer, and evolve into complex, gullied shores. The hardest of all, the Cornish granites and Scottish gneisses, seem eternal and the sea’s erosive power appears simply to polish them, giving smooth, rounded blocks and boulders along cliffs that scarcely seem to have changed since they were first formed. And yet the coastline is in a continuous state of change. In addition to changes wrought by the power of the sea, the structure of the coast has been influenced by changes in sea level, which has risen and fallen throughout the history of the British Isles. Today, about one third of Britain’s coastline is more or less stable, and has been for some tens of thousands of years, but one third is rising while another third is actively sinking. Thus, the coastline seen today is complicated by the traces, often very obvious, of older shorelines and seashores.

      The nature of the intertidal marine environment at a particular location obviously depends on rock type and topography, and is mediated by aspect, that is, whether it faces an open oceanic coast or is sheltered to a varying degree. Exposed, eroding coastlines provide rocky habitats and gravel banks, but the products of erosion find their way onto sheltered depositing coastlines, where they form a variety of unconsolidated, or mobile, shores of muds, sands and fine gravels. Given the wonderful variety of geology and topography around Britain, and remembering its geographical position, it is no wonder that every kind of intertidal marine habitat can be found along its coasts.

      Every seashore naturalist has rejoiced in the extraordinarily rich diversity of plants and animals to be found between the tides. Rocky coasts are usually clothed by a dense but narrow band of large brown seaweeds; intertidal rock surfaces are plastered with sheets of barnacles or mussels; damp or shaded surfaces bear clumps of sea squirts and mounds of sponges; surf-washed platforms are covered with a tight turf of small red algae. Among all this can be found a host of small and large animals; along the edges of the tide one sees groups or shoals of fish, and above it flocks of feeding birds. It is evidently a highly productive habitat, yet one that changes seasonally. The summer profusion found between the tidemarks seems to vanish with the advent of winter, to reappear just as abruptly in the spring. This phenomenon demonstrates an important aspect of the seashore environment, as a specially modified marine habitat. Coastal waters are generally shallow and sunshine penetrates to the sea floor; they are thus well lit and warm up swiftly in the spring. Conversely, their very shallowness, together with their proximity to land, ensures that they cool rapidly in the winter. Deep-water bodies, away from continental masses, tend to retain heat, warm and cool more slowly, and show less marked differences between summer and winter temperatures.

      The constant movement of the sea along the coast is responsible for conditioning the coastal environment in other important ways. Ocean swells, tidal currents and wind-driven waves continually stir and overturn inshore waters. Oxygen is thereby dissolved far more rapidly than by diffusion through a calm water/air interface and particulate matter and nutrient salts are resuspended with every tide and storm, making them constantly available to the shallow coastal fauna and flora. Light, warmth, oxygen and dissolved nutrients provide all the conditions for fast and sustained plant growth; they stimulate the annual explosive growth of planktonic algae, the phytoplankton, and support the perennial forest of attached macroalgae. These are the twin bases of coastal ecology, providing the most important sources of energy for coastal organisms.

      Seashores are thus highly productive during particular periods of the year and it is not surprising that a diverse assemblage of animals exists to exploit this productivity. Many of the species are permanent seashore residents, barnacles, mussels, limpets, winkles, tubeworms and many others, adapted for seashore living. But every spring the array of species present is enormously increased by seasonal migrants which move inshore to take advantage of the abundance of food available, to fuel their own reproductive cycles. Numerous species of fish, for example, take up residence between the tides for the duration of the summer months. Others merely visit for a brief period in spring, depositing their eggs where they will be safe from predators, or where growing juveniles can find a ready supply of food. Crabs move inshore each year to feed and mate; impregnated females move offshore again at the end of the summer, but their larvae, spending their first few months of life in the plankton, return to the shore as they metamorphose and there grow rapidly through their first summer.

      Against all the advantages of coastal living must be set the severe disadvantage posed by the twice-daily retreat of the sea, exposing the inhabitants to the dangers of sunshine and wind, or frost and rain, and to an additional suite of predators. Respiration must slow or stop during this period, water loss must be avoided or controlled, and temperature change must be endured. On sandy shores the preponderance of the inhabitants retreats into burrows or moves downshore with the falling tide. In deep, low-shore rock pools, animals and plants may be able to withstand the minor fluctuation in temperature experienced during their brief exposure, but higher up the shore pools may be isolated from the main body of seawater for hours at a time. Temperatures may then rise, or fall, sharply; oxygen is swiftly depleted in warm, shallow pools and pH may fluctuate uncomfortably, and rainwater may dangerously freshen the pool environment. Physiological stress is thus characteristic of intertidal living. Temperature in particular may fluctuate more widely and more swiftly than in either of the adjacent marine and terrestrial environments. Low tide on a hot summer day may be so dangerous for shore crabs that they must use water reserves to cool themselves, balancing the risk of dehydration against the threat of heat stress. Air frost is equally damaging and the winter pool retreats occupied by some shore dwellers need to be sufficiently deep to avoid the risk of drying out. Yet even a casual survey of a rocky shore will suggest that plant and animal communities have nevertheless adapted to these often inimical conditions. The most obvious evidence is the clearly displayed zonation of the brown seaweeds, each species flourishing upshore to the limit of its physiological tolerance.

      The flora of the intertidal zone and adjacent shallow coastal waters is entirely dominated by macroalgae. Fungi can be found, but are generally inconspicuous; beds of seagrasses, true vascular plants, colonise shallow sea floors of fine muddy sand, and some tropical reef flat or lagoonal habitats are monopolised by cyanophytes, or blue-green algae. On temperate rocky shores the highest part of the intertidal and, especially on wave-exposed shores, the spray-washed fringe above high-water mark, are usually colonised by lichens. However, between and just below the tides, especially along temperate coastlines, is where the algae flourish, and achieve their most extraordinary expression of form and diversity. Algae are rarely important or conspicuous in terrestrial habitats, yet they are of immense importance in the sea, forming the basis of almost all marine food chains. The microscopic, non-cellular plants that constitute the phytoplankton are undoubtedly of primary significance; but, although their total standing crop can scarcely be computed, they are almost unnoticeable except in time of bloom, when their overwhelming abundance colours the water. It is only in shallow waters within the depth limit penetrated by sunlight that algae occur as attached, multicellular, perennial plants. They are then analogous to terrestrial vegetation, forming forests and understorey, shrubberies and turfs of close-packed plants, each firmly secured to the sea floor or seashore by a branching, root-like holdfast. However, there the resemblance ends. Although algae photosynthesise, they must derive all the nutrients necessary for growth by absorption through the surfaces of their fronds; their holdfasts function only to secure them, and unlike the roots of terrestrial plants draw nothing from the substratum they are attached to. Considering that the intertidal zone and sublittoral fringe constitute a tiny portion of the marine realm, relative to the immense area and volume of the oceans, it is all the more remarkable that the algae achieve such a wide diversity of size, form and reproductive biology in such a small compass. However, given the productive capacity of this narrow strip of habitat, it is not so surprising that growth rate and standing crop of coastal macroalgal populations can be staggeringly high. As a consequence the amount of dissolved organic material, originating as algal mucus, and decayed particulate matter produced by the algal zone each year is immense.

      It has been frequently stated, but is always worth repeating, that examples of almost every major class of almost every animal phylum can be found between the tides, or at least close to low-water mark of spring tides. It is interesting to note the exceptions: cephalochordates and pogonophorans live offshore, and do not occur on the seashore; chaetognaths (arrow worms) and ctenophores (comb jellies) are entirely planktonic, although the latter are commonly found stranded at low water on sandy shores. These four small phyla aside, 23 other animal phyla have intertidal or shallow coastal representatives, and 14 of these can be found, often in a wide taxonomic variety, on a good day’s field work. The task of charting the diversity of marine animals has scarcely begun and comparisons between marine, freshwater and terrestrial habitats are rather pointless. Yet it is worth remarking that the overwhelming majority of the species presently described from land and fresh water belong to the insects and the vertebrates, with a few minor groups of molluscs and crustaceans reasonably well represented. It is just not possible to experience the same broad systematic and evolutionary spectrum one finds on the seashore anywhere else.

      Almost all the fauna found on the seashore has marine evolutionary origins. Some are at the extreme upper limit of their distribution and may be uncommon on the shore, occurring only infrequently at the lowest level of the tide and in the shallow sublittoral. Many small whelk species, burrowing bivalves and sea stars, for example, are more common offshore, and range to at least the edge of the continental shelf, at about 200 metres. Many more are quite at home in shallow waters and range to a varying degree, and in varying abundance, onto the seashore. Among these are numerous species of brittlestar, mud-dwelling bristle worms, shrimps and prawns, and the teeming fauna associated with seaweeds, which includes a hundred or so species of small gastropod. A significant proportion represents species, or larger taxonomic groups, which have fully marine evolutionary origins, but which are adapted to an entirely intertidal existence. These include limpets, winkles and top shells, few species of which range far below the lower limits of the tide, and, of course, the acorn barnacles which thrive in vast numbers on the upper reaches of the shore. The Amphipoda includes genera and families almost entirely limited to intertidal living, especially as part of the infauna of sandy beaches or among strandline debris. The gammarid amphipods have almost completely forsaken the seashore; a majority of species in that family is found in brackish or freshwater habitats and very few lead a marine existence. Strictly intertidal species are of especial interest in that their biology and ecology reflect their precarious existence on the sea’s margins. They display often remarkable behavioural adaptations, which ensure that they find and remain in their optimum microhabitat, or at their preferred tidal level, and their reproductive behaviour and life cycles are usually closely attuned to seasonal cycles of growth and productivity on the seashore.

      Amongst all the marine inhabitants of the seashore there also exists a small but interesting minority of animal species whose evolutionary backgrounds are essentially terrestrial. A large number of insect species, many arachnids and a few millipedes and centipedes, live along the strandline, in and among seaweed debris. Some may be equally common in similar, damp habitats inland, but others are particularly adapted to seaside living. Some seaweed flies spend a large part of their life cycle, usually as larvae, actually between the tides. They are still air breathing, but survive high-tide periods in air pockets deep within rock crevices.

      The mites are the most adventurous of this terrestrial element and a large number of species, representing dozens of different families, has evolved equipped for intertidal living. One family, the Halacaridae, appropriately called the ‘sea mites’, is completely marine in distribution and includes many sublittoral species, some of which venture into the deep sea. The molluscs include a few air-breathing pulmonate gastropods, relatives of a group that long ago adapted to terrestrial living to the extent of losing the gill, instead employing the mantle cavity as a lung. These returned migrants continue to be air breathing. They generally live on the upper and middle reaches of the shore, moving downwards to feed as the tide recedes, but always retreating again on the flood. The most well known of these are the pulmonate sea slugs; Onchidella celtica is the sole representative of these in Britain. Pulmonate snails and limpets are most common on tropical and subtropical seashores, but on western coasts of Britain a few small species of pulmonate snail can be found on rocky or marshy shores.

      Coastlines and seashores have profoundly influenced human cultural evolution and history. The coast has been both a major barrier and a migration pathway, and an important source of food on sometimes otherwise inhospitable shores. In northern Europe, as indeed in other parts of the world, the migrations of early Neolithic peoples can often be plotted by shell mounds left along their routes. However, if a very broad generalisation may be permitted, it seems that, except on the smallest islands, migrating peoples rapidly move inland. Coastal fishing communities seem most often to have been rather isolated from inland communities. Celtic Britain no doubt had small coastal settlements that depended on the sea and the seashore for part of their food. However, it was probably in Roman times, in southern England, that commercial fisheries were first established for oysters, flatfish and coastal species such as mullet and bass. Inshore fishing was certainly flourishing and well organised by the late twelfth century. The Magna Carta included provisions for the protection of fisheries, and through the Middle Ages numerous laws and statutes were enacted regulating the equipment to be used in the catching of fish, the areas to be fished and the type and number of fish to be taken. While they served mostly to protect the interests of fishermen and fish merchants, and especially those with legal right to take river fish, these laws now seem radically environmental in their objectives. At various times it was forbidden to obstruct the movement of salmon by the construction of weirs; small mesh nets were outlawed as likely to trap and destroy fish fry, and the use of dredges was considered to be damaging to the sea floor. Many traditional fisheries were conducted from the seashore, employing fish weirs, or stationary traps built in tidal regions. Basket traps, and fixed nets and lines were also used, as were long seine nets, dragged along the beach or deployed from small boats. An iron-mouthed oyster dredge seems to have been in wide use in southern England by the end of the thirteenth century and, by the mid-fourteenth century, the increasing use of a wide-mouthed bottom trawl, which seems to have evolved through a fusion of the oyster dredge with the seine net, was causing such concern that Parliament petitioned Edward III to abolish it.

      Offshore fishing became well established throughout northern Europe from the late fourteenth century and while local, traditional fishing techniques persisted, especially in the case of inshore or estuarine species such as oysters, whitebait or flatfish, the commercial significance of seashore fisheries more or less disappeared over the following centuries. Local demand perhaps kept alive the tradition of gathering cockles, mussels, winkles and shrimps from intertidal areas; today these are once more commercially important fisheries. Shellfish culture, of mussels and oysters, and salmon ranching, are now well established in Britain. At Arcachon, in western France, mussels are traditionally grown in tidal areas, but the best results are obtained when both oysters and mussels are grown in the shallow subtidal, off the bottom and away from the attention of predators. Both salmon enclosures and rafts supporting shellfish cages need to be moored in sheltered areas and although the seashore is therefore physically unaffected, the introduction of these new elements must eventually influence its ecology.

      Another new form of commercial exploitation of the seashore results from the burgeoning leisure industry. Sea angling is a major leisure pursuit and it has an apparently limitless demand for live, natural bait. Lugworm habitats in some parts of Britain have been devastated by professional bait-diggers, and intertidal populations of razor fish seem also to be declining rapidly. Rocky shores, also, suffer from the attention of bait collectors, and the continual overturning of lower shore rocks and boulders in search of soft crabs not only destroys the attached plants and animals growing on both upper and lower surfaces, but also the fauna living in the sediment beneath. In some regions local bylaws already regulate shore collecting of crabs, lobsters, cockles, and even lugworms, and it is certain that in the near future the commercial exploitation of all intertidal animals will have to be severely limited, or even prohibited, if the habitat is to survive. Seaweeds have been exploited in many ways, both for local usage and, at various times, as a resource for larger, national industries. A few species continue to be gathered in small quantities for human food; laver bread, or Porphyra, is eaten in South Wales, and in parts of Ireland and Scotland dulse, Palmaria, and carrageen, Chondrus, are still used as seasonings or salad dressings. The British, however, have never developed the same enthusiasm for seaweed displayed by the Japanese. Large brown seaweeds are traditionally used as agricultural fertilisers, and in fact have always been critically important to people farming the poor, sandy soils of the Hebrides, and parts of western Ireland. They also provided good feeding for livestock, especially sheep, although the small, hardy breeds developed for seashore grazing are not widely kept now. Kelps and wracks have been important sources of soda, potash, iodine and agar, to the extent that considerable industries once depended upon them. Today, they have an increasing significance to the food and chemical industries as a source of alginates. The world demand for these substances is presently supplied mostly by the giant kelps of the new world, but northwest Europe has always provided a proportion of the demand and it is quite probable that the kelp forests of northern Britain will soon be again the focus of commercial exploitation. In Japan and parts of China many red, brown and green seaweeds are cultivated, grown on rafts or lattice structures in shallow, coastal embayments. This is infinitely preferable to stripping the seashore of its natural vegetation, and as a bonus increases the natural habitat available to coastal animals.

      The industrialisation of Britain has had an immense impact on the seashore environment, and future development seems likely to heighten this impact. The history of Britain’s industrial evolution can be viewed from the sea. Ports and dockyards developed naturally around river mouths, as did shipbuilding and primary industries. Canals, and later railways, linked the ports with inland manufactories. Chemical plants, refineries and power stations could be cheaply built on flat coastal lands, away from urban centres, close to water supplies and with the sea convenient for the disposal of effluent. In many regions of Britain the old industries have now gone, often leaving a terribly scarred or damaged environment. Some have been replaced by new industries, enterprise parks, heritage sites or urban developments, while some of the most recent industrial sites, such as petrochemical plants or power stations, continue to expand. The sea has always been viewed as a convenient and practical dump, with the larger rivers providing ideal conduits for the disposal of every kind of rubbish and waste produced by human beings. The sea’s capacity for absorbing trash, breaking it down and recycling it through marine food webs was considered to be limitless, and it was not until the middle years of the twentieth century that a few dissenting opinions were heard. Rachel Carson’s prophetic book Silent Spring (1966) sounded a dire warning of the threat from pesticides, but even she perhaps could not have imagined the looming disasters now threatening immense areas of the world’s seas. The sheer volume of filth produced by northern Europe, and the increasing proportion of persistent, damaging pollutants in urban and industrial effluents, could completely destroy the ecological balance of large areas of the North Sea. A series of human tragedies and dramatic environmental disasters brought marine pollution and its consequences into public focus during the decade following the publication of Carson’s book. It became the object of research and teaching, it promoted well-meaning but largely useless legislation, and provided powerful justification for popular environmental and conservation movements. Alas, the environmental degradation of the sea, and the seashores of the British Isles, has continued relentlessly and one can only echo P. H. Gosse’s lament for the ruined shores of South Devon (in E. W. Gosse, 1907). Perhaps the most critical influence on seashore ecology in the years ahead will be that engendered by proposed barrages across major tidal rivers, such as the Severn, or large shallow embayments, such as Morecambe Bay. Enormous areas of intertidal habitat will be permanently immersed and lost, and both supralittoral and sublittoral habitats may be profoundly altered. The gross consequences for the ecology of these areas might be predictable, and considered affordable, but the ecological complexity of marine habitats is still incompletely understood even on Britain’s well-studied shores, and the ultimate effects of such major interference with coastal ecosystems can only be guessed.

      Notwithstanding the major threat imposed by continued and continuous industrialisation, perhaps the most sustained pressure suffered by the seashore results from the late twentieth-century boom in the business of leisure. It is extraordinary that for so much of its modern history the majority of Britain’s population had no contact with the coast. Fishermen, mariners, dockers and shipbuilders were dependent on the coast and the sea; but it was a place to earn a living, not to visit for pleasure, and most people thus had no occasion to go there. This began to change in the late eighteenth century when fashionable and wealthy people developed a taste for sea bathing as a healthy, though not necessarily pleasurable, activity. Liberalising attitudes to work, rest and recreation among the Victorians eventually opened up the coast to the rest of the British population. Seaside holidays became a tradition by the beginning of the twentieth century, stimulating development and expansion of coastal resorts that continued for more than six decades. Cheap foreign travel eventually led to the decline of many British seaside resorts, but overseas holidays are unlikely to be inexpensive for very much longer and the holidaying British may have to be content with their own coasts. This revived seaside holiday will be rather different from its traditional origins. Small harbour towns in southern Britain, now redundant as fishing ports, or as centres for small-scale shipbuilding, export or as ferry terminals, make ideal yacht havens. Marinas, as they are now termed, are spreading through all the small coastal towns of England, Wales and western Scotland, as well as the coasts of Ireland. Their development demands building aggregates, cheaply obtained just offshore; sustaining them demands power, water, sewage disposal and somewhere to dump rubbish. The dog whelk, Nucella lapillus, was the first observed casualty of this new threat. In southwest England, poisoned by tributyl tin originating from yacht paints, female dog whelks were effectively sterilised and local populations crashed. Although the gross pollutions of primary industries are now gone or contained, it seems that the seashore today faces less obvious but even more insidious threats.

      Naturalists first began to explore Britain’s seashore during the eighteenth century. Popular interest was then largely directed towards seashells, corals or other attractive objects, especially from exotic parts of the world, but the first scientists to study intertidal life became absorbed with the more unfamiliar organisms living there. The hydroids, sea anemones and bryozoans, collectively termed zoophytes, corallines, or simply polyps, attracted considerable debate regarding their nature – were they plants or animals? John Ellis’ work, An Essay towards a Natural History of the Corallines (1755), is a fine example of the many books on this topic that appeared in the latter part of the century. Popular marine biology truly blossomed during the nineteenth century, reaching an apogee in late Victorian times that has still to be surpassed. Seashore studies were seen as a healthy pursuit, an intellectually stimulating pastime, and particularly as a way of appreciating the marvels of God’s creation. A galaxy of scholarly specialists produced a steady flow of fine, illustrated books for the benefit of the growing amateur naturalist market. Apart from small pocket books, such as those of the Revd W. G. Wood, there were monographs published on every phylum and class of the fauna of the seashore and shallow seas of Britain, and of course an equal number devoted to seaweeds. Scientific study of the sea and seashore was largely promoted by these Victorian specialists, who founded clubs and societies, and eventually stimulated the formation of a special group, the Dredging Committee, within the British Association for the Advancement of Science. This group, inspired by the enthusiasm of Edward Forbes, finally established the science of marine biology.

      The oceanographic surveys and expeditions of the final quarter of the nineteenth century were one great consequence of the early Victorian passion for natural history. Another was the establishment, here and in Europe, of shore-based marine stations, such as those at Plymouth, Millport, St. Andrews and Port Erin, where zoologists and botanists could research and teach. Intertidal marine biology and ecology became an important part of diploma and degree courses, and remain so. Initially, the seashore provided a perfect natural laboratory where students could find and experiment with examples of all the most important and diverse animal phyla. The varied life cycles and reproductive habits, feeding strategies, growth forms, associations and symbioses suggested an endless sequence of questions and problems for investigation. Environmental awareness, and the impact of human activity on complex, finely balanced but finite habitats ensures that marine biology, and the ecology of our shores, will remain important areas for research, and become increasingly important as indicators of the health and vitality of the seas around us. Yet, while scientific interest and concern is bound to grow, natural history continues to absorb the interest of a substantial proportion of the British public and there is still much to occupy the amateur naturalist on the seashore. Of the several thousand animal and plant species found on the shore or in shallow coastal seas of Britain, the biology and ecology of very few are known with any detail. Growth rates, size distributions, longevity and age structure of populations, even food preferences and breeding periods, are unrecorded or unknown for very many species. Local recording and monitoring schemes can supply interesting and scientifically important information, and lead the most enthusiastic observers into new fields of ecological and biological research in what is still the most exciting natural habitat in these islands.

      2

      Living on the Seashore

      The physical environment

      The open sea is a remarkably benign environment, and seawater itself is an ideal medium in which to live, grow and reproduce. Water is the major constituent of all living tissues, and organisms living immersed in water avoid the lethal risk of dehydration. Away from coasts, seawater is a constant medium: 96.5 per cent by weight consists of pure water, the other 3.5 per cent consists of dissolved elements, salts and gases. This proportion, 35 grams per 1,000 grams, is termed the total salinity of seawater, and has usually been expressed as parts per thousand, ppt or ‰; salinity measurements are now expressed as ratios related to the conductivity of standard 35 ppt seawater, and the number does not require a unit. The preponderance of solutes, 99.78 per cent, consists of just six ionised compounds, which exist everywhere in exactly the same proportion; sodium (Na+) and chlorine (Cl-), together forming salt, are the most important of these. Salinity varies over the oceans, from as low as 32 in polar seas affected by freshening ice-melt, to as high as 37 in tropical regions with low rainfall and high evaporation; globally it averages around 35. Water is an excellent solvent; negatively charged ionised compounds bond with the hydrogen end of the water molecules while positive ions bond with the oxygen atom. A proportion of the H2O molecules ionise to give hydrogen (H+) and hydroxyl (OH-) ions that may also react with charged ions of other compounds. The balance between H+ and OH- ions determines the relative acidity of water: as H+ ions increase acidity rises, while increasing OH- ions leads to an alkaline medium. Relative acidity of water is measured on a logarithmic scale of 1 to 14: the lower the value, the higher the concentration of H+ ions, and the greater the value, the higher the concentration of OH- ions. Pure water is neutral at pH 7, but dissolved sodium, potassium and calcium impart a slightly alkaline quality to seawater, the pH value of which ranges from 7.5 to 8.4. Seawater is maintained at this narrow pH range through the chemical intervention of carbon dioxide, which dissolves more readily than other gases because of the reactions it initiates as it goes into solution. Dissolved carbon dioxide initially forms carbonic acid, which then dissociates to form hydrogen (H+) and bicarbonate (HCO3) ions; the latter may then further dissociate into H+ and carbonate (CO3-) ions. All three actions are reversible, and the result is a dynamic balance in the concentration of free H+ ions in seawater, and hence also a little-varying pH value. Carbon dioxide is of profound significance in metabolic processes of living organisms; it is utilised in photosynthesis, and is the major product of respiration; in the sea it is additionally critical in the processes of calcification, and the carbonic acid–bicarbonate–carbonate reaction serves to maintain a constant reservoir of the gas in the world ocean.

      Oxygen is equally important to living processes, but dissolves less readily than carbon dioxide; in the upper 10 metres or so of the oceans’ surface layers oxygen concentration is usually less than one-tenth of that of the atmosphere, and it decreases rapidly to a minimum at about 500 metres. Oxygen is swiftly depleted by biological processes, and the minimum oxygen zone reflects this; in fact, below 500 metres, oxygen levels increase again with increasing depth and below 2,000 metres may be as high as surface levels. This is partly attributable to the fact that oxygen dissolves more readily at lower temperatures; cold water is dense, and at the poles intensely cold, oxygen-rich water sinks to the ocean floor and flows equatorwards, providing oxygen for deep benthic (bottom-dwelling) organisms.

      Temperature regulates all living processes, and a further advantage to living in the sea is that ambient temperature fluctuates only minimally, on both daily and seasonal bases, in comparison to terrestrial environments. Water absorbs and retains heat, losing it only very slowly; thus, at any particular latitude the variation in sea surface temperature between day and night is negligible while around the British Isles, for example, the annual range in surface temperatures may be as little as 5°C off western coasts to a maximum of 12°C off the southeastern corner. Temperature decreases with depth but stabilises, so that below 200 metres the annual range may be just a few degrees.

      Oceanic seawater thus provides an environment characterised by remarkably constant physical and chemical attributes. It is, however, deficient in one important aspect: chemical nutrients. The planktonic microalgae that are at the base of marine food chains require carbon dioxide, water and sunlight for photosynthesis, but cannot build organic molecules without the critical addition of nitrates and phosphates, and these are everywhere in short supply. Unlike the other constituents of seawater these two nutrient salts do not exist in fixed ratio and are always rapidly used up by plankton growth as they become available. There is a constant loss of nutrients as plants and animals die and sink to the sea floor. Over the deep ocean these nutrients, once lost, are irrecoverable, and for this reason the surface layers of the open oceans are relatively barren habitats, except for limited areas where diverging currents allow nutrient-rich deep bottom water to upwell. Planktonic marine life only really flourishes on and above the fringing shelves of the continents, where winds create convectional mixing that reaches to the sea’s bottom, stirs up the lowest layers and periodically restores the vital nutrients to the surface. In coastal waters, shallower than 50 metres or so, nutrients are probably never in short supply, the constant motion engendered by tides and winds continually stirring up bottom sediments. Further, close to land, nitrates and phosphates are added to the marine environment in river outflows, and as wind-blown terrestrial sediments.

      With the possible exception of the abyssal sea floor, the richest marine habitats are those adjacent to the continental landmasses, where most marine organisms achieve their greatest abundance and diversity. The biological wealth of an ecosystem can be quantified in several ways, and, apart from species abundance and diversity, for ecologists the most useful measures are biomass, the total living material in a given habitat, and productivity, which estimates the growth and reproductive output of the biological community. In order to render all biological components of a community comparable, biomass is usually expressed as units of organic carbon per unit area or volume. Productivity is similarly expressed – g C/m², g C/m³ or g C/l – per unit of time. The distinction between these two measures is important: a bottom-living community may consist of large, long-lived and slow-growing animals, such as horse mussels, with a rather low and sometimes irregular reproductive output. The biomass (B) of such a community will be large, but the annual productivity (P) comparatively small; the ratio, P/B, between the two values will also be correspondingly low. Other communities may consist of mostly small polychaetes and crustaceans, which grow and reproduce rapidly, often passing through several generations each year. This type of community will have a small biomass but an impressive productivity and a consequently high P/B ratio. In reality, stable bottom communities tend to consist of mixtures of both types of organism and the objective of the ecologist is often to quantify the relative contribution of each type, or even species, to the biomass and productivity of the community.

      Comparisons of productivity values are often instructive. Tropical rainforests are usually cited as the richest biological communities on earth, and their productivity is often astonishingly high, in cases exceeding 3,000 g C/m²/year. Tropical mangroves have equally high productivity levels, and coral reefs, largely through the photosynthesising activity of the corals’ symbiotic zooxanthellae, are also impressive producers. However, temperate seagrasses, such as Zostera, sometimes achieve productivity rates as high as their tropical counterparts, and both coastal salt marshes and the macroalgal belt of north European coasts can have very high productivity levels, rivalling those of tropical forests and coral reefs. This is perhaps not surprising when one considers the growth rates of some seaweeds; the kelp, Saccorhiza polyschides, for example, grows from a microscopic spore to a plant weighed in kilograms in just ten months. In the marine realm, biomass and productivity values demonstrate the profoundly different ecological conditions in the blue oceans and the shallow shelf seas. Productivity values refer usually to primary productivity, which measures the production of organic carbon through photosynthesis. Phytoplankton communities of the nutrient-poor open ocean are sparse, and primary productivity averages no more than 50 g C/m²/year. In the temperate shelf seas of northern Europe, where nutrients are recycled with every winter storm, phytoplankton growth is switched on by increasing temperature and day length every spring, and this spring bloom may result in productivity values of 250 g C/m²/year. The coastal macroalgal communities have the highest primary productivity in temperate seas. Intertidal fucoid algae may achieve 300 g C/m² yearly, while that of the subtidal kelp forests may exceed 1,500 g

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