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    Bird Populations
    Bird Populations
    Bird Populations
    Ebook1,089 pages51 hours

    Bird Populations

    By Ian Newton

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    In the latest addition to the New Naturalist series, Ian Newton explores bird populations and what causes their fluctuation – food supplies, competitors, predators, parasites, pathogens and human activity.

    The combination of a rapidly expanding human population, a predominantly utilitarian attitude to land, central government policy on land use and increasing mechanisation have combined to promote massive changes in land use in recent decades than at any previous comparable period. These developments have in turn brought huge changes in bird habitats and populations; some species dependent on the old landscapes declined, while others have increased. Over the same period, changing public attitudes to wildlife allowed previously scarce bird species to recover from past onslaughts, while climate warming has promoted further changes.

    In this seminal new work, Ian Newton sets out to explain why different bird species are distributed as they are, and changed over the years in the way that they have. The regular watching and study of birds now provides a source of recreation and pleasure for very large numbers of people, while continued monitoring of bird numbers can also alert us to impending environmental problems. For all of us, a world with fewer birds would be a poorer place.

    LanguageEnglish
    PublisherHarperCollins UK
    Release dateAug 29, 2013
    ISBN9780007527991
    Bird Populations
    Author

    Ian Newton

    Dr. Ian Newton is respected world-wide both as a biologist with a special interest and expertise in this subject and as a communicator. He is a seasoned and popular keynote speaker at National and International meetings, and his talks are often the high point of conferences. Ian Newton was born and raised in north Derbyshire. He attended Chesterfield Boys Grammar School, followed by the universities of Bristol and Oxford. He has been interested in birds since boyhood, and as a teenager developed a particular fascination with finches, which later led to doctoral and post-doctoral studies on these birds. Later in life he became known for his penetrating field studies of bird populations, notably on raptors. He is now a senior ecologist with the Natural Environment Research Council and visiting professor of ornithology at the University of Oxford.

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      Book preview

      Bird Populations - Ian Newton

      CHAPTER 1

      Preview

      THIS BOOK IS ABOUT BIRD NUMBERS . It discusses why particular species are as numerous as they are, and not more or less numerous, and why some species are increasing while others are decreasing. It is concerned with the various things that influence bird abundance, such as food and other resources, competitors, predators, parasites and pathogens, and also various human impacts. These matters are of interest to anyone with a concern for birds, whether naturalists or researchers, or people involved in managing bird populations for reasons of conservation, crop protection or hunting. This initial chapter introduces some of the main ideas in bird population ecology, and outlines some of the aspects developed in detail in later chapters.

      Much of what we know about the population processes of animals comes from studies on birds. The reasons are not hard to find, for compared to many other animals, birds are relatively easy to study. Most species are active by day; they are conspicuous and can be recognised by their appearance, songs or calls, which makes them easy to detect and count. In addition, most birds can be trapped and marked with leg rings or other tags. They then become identifiable as individuals, enabling their movements and life histories to be followed, and their survival rates to be calculated. Many species, too, are large enough to carry radio-transmitters or other electronic devices, so that their day-to-day activities and movements can be recorded. When tracked using satellites or geolocator tags, individuals can also be followed on their migrations, wherever on earth they travel. Hardly any other kinds of animals show such an obliging combination of characteristics.

      In addition, most bird species rear their young in nests which can be found by watching the adults or searching likely places. Their individual breeding rates can then be measured in ways that are not possible for most other creatures, which show no parental care and in which the young from different parents soon intermingle. The highly developed social structure of birds has also enabled the role of territorial and other aggressive behaviour to be studied in greater detail than for most other animals, revealing its role in population regulation.

      All these features combine to make birds popular with their human observers, and ideal subjects for study, whether by professional ornithologists based in universities and research institutes or by others for personal interest. Organisations such as the British Trust for Ornithology (BTO) rely on the combined voluntary efforts of bird-watchers to monitor changes in the numbers and distributions of birds, both countrywide and long-term. Not surprisingly, then, the population ecology of birds has been studied in greater depth, and by more participants, than that of any other group of organisms, and the general principles that have emerged have wider relevance.

      BIRD NUMBERS

      About 260 bird species occur regularly in Britain and Ireland at the present time, and a similar number occur as rare visitors or vagrants. Among the regulars, some species (144) occur here throughout the year, others as summer visitors (54) and yet others as winter visitors (142), or as passage migrants (122) which travel through the country twice each year on their migrations. Many species occur in more than one category, and for part of each year, two or more different populations of the same species are represented in the British Isles (Newton 2010).

      The species that breed here vary from widespread and abundant, numbering several millions, to extremely local and rare, with fewer than 100 individuals. Among land-birds, the most numerous breeding species include the Blackbird and Wren, both of which number more than ten million individuals in the breeding season, and are now more plentiful than the House Sparrow, which over most of the country has suffered a marked decline in recent decades. Some species give the impression of huge numbers because they gather in great concentrations. The Starling is no longer one of the commonest breeding birds in Britain, but can still be seen in big numbers at its winter roosts, comprising tens of thousands of individuals. Formerly roosts were even larger, frequently containing over 100,000 individuals and occasionally more than a million. Seabirds gather in large breeding colonies, yet spread out to forage over wide areas of open sea, where their average densities can be quite low. At the other extreme, the rarest regular breeding birds in the British Isles, such as the Red-backed Shrike and Purple Sandpiper, probably number no more than a few pairs in any one year.

      FIG 1. The Blackbird Turdus merula is one of the commonest bird species in Britain and Ireland, occurring anywhere with trees and bushes, from countryside to city centres. (Keith Kirk)

      In general, Britain and Ireland have fewer breeding land-bird species than other comparable areas at similar latitude in continental Europe, and Ireland has notably fewer than Britain, but both islands have more seabirds. These differences can be attributed to the position of our islands, off the main continental landmass but surrounded by highly productive seas. For wintering birds, however, Britain and Ireland hold a relatively large number of species, present in much greater numbers of individuals than comparable parts of continental Europe. This can be attributed to the mild climate, making our islands attractive as a wintering area for birds drawn from a large span of the Arctic, from eastern Canada through Greenland, Iceland and northern Europe to central Siberia. Not all birds from these northern areas winter in Britain and Ireland, but these islands draw wintering birds from a wider swathe of the Arctic than any other equivalent area at comparable latitude.

      FIG 2. Pre-roost gathering of Starlings Sturnus vulgaris. These birds tend to roost in the same places year after year, whether conifer woods, reedbeds, cliffs or city buildings, gathering each evening from the surrounding countryside up to 20 km away. (Keith Kirk)

      FIG 3. The Canada Goose Branta canadensis was introduced from North America in the seventeenth century, but is now one of the most widespread non-native bird species in Britain, estimated at more than 60,000 pairs. (Richard Chandler)

      To the various species native to Britain and Ireland we can add more than a dozen introduced (or escaped) alien species, which now have self-sustaining resident breeding populations (Holling et al. 2012). The most widespread of these aliens include the Pheasant, Red-legged Partridge, Little Owl and Canada Goose, but other aliens currently have more restricted distributions, while new ones are continually being added as they escape from bird collections. These non-native species must be separated from other naturally occurring ones, such as the Capercaillie and White-tailed Eagle, which were reintroduced after their earlier elimination.

      COUNTING BIRDS

      The first step in any study of bird populations is to count the birds reliably. Except for highly localised species, it is seldom possible to study the whole population of any bird. Instead the researcher has usually to work within a defined area, whose avian occupants may form a tiny part of a much wider population. Individuals may move freely in and out of the study area, and the birds which breed there may occupy a wider area, or even a different part of the world, outside the breeding season. Hence, most population studies of birds have been concerned with the numbers found in defined areas at specific times. The general applicability of the conclusions from such localised studies depends partly on how typical is the study area of the bird’s range as a whole, and on whether the area is big enough compared with the scale over which the factors influencing overall population levels operate.

      In recent decades, however, more widespread data on bird populations have become increasingly available. Most of our information on the population trends of the commoner British birds now comes from the annual nationwide monitoring programmes organised by the BTO: the Common Birds Census which began in the 1960s and, after a period of overlap, gave way in 1994 to the Breeding Bird Survey. Both schemes depend on volunteer observers making regular bird counts in particular areas year after year. The Breeding Bird Survey uses a less time-consuming method than its predecessor, and is of more robust statistical design. It attracts more participants, which enables a wider range of randomly selected areas to be sampled in different habitats across the country. The findings on year-to-year population trends can thus be considered to be reasonably representative of the country as a whole, and cannot be considered atypical in the way that studies on any single plot might be. However, not all bird species can be readily sampled in this way, and some rarer, less accessible or nocturnal species are monitored by special surveys, conducted at intervals of several to many years, depending on species. For example, since the 1960s, the Peregrine has been surveyed nationally about every ten years, but as with many other uncommon species, in some localised areas the breeding numbers are checked annually by dedicated enthusiasts.

      For most birds at our latitudes, the annual calendar comprises two distinct phases: a breeding (summer) season, when numbers increase because reproduction exceeds mortality, and a non-breeding (winter) season, when numbers decline because of mortality. Overall numbers therefore reach a seasonal peak immediately after one breeding season and a seasonal trough early in the next, although frequent movements mean that this pattern may not be evident in every patch of habitat. The extent of these seasonal fluctuations depends mainly on the reproductive and mortality rates of the species concerned, being most pronounced in the most prolific species (Fig. 4). Many bird species can be counted most easily while they are breeding, because they are most conspicuous then, and tied for long periods to specific locations where they nest. In consequence, our knowledge of their population changes, from year to year or from place to place, is based entirely on counts of breeding numbers, or on indices of breeding numbers, such as displaying males or nests. Assessments of breeding numbers are of particular value, because it is at the beginning of the breeding season that numbers reach their lowest level of the year, and it is upon breeders that future additions to the population depend. Hence, throughout this book, unless otherwise stated, I shall be concerned primarily with the limitation of bird breeding numbers. This is an important point, because different factors can limit bird numbers at different times of year. Many species suffer high predation of eggs and young, so that predation may limit their post-breeding numbers, as counted in late summer. But with or without this predation, winter food can support only a certain number of individuals so that others starve, in which case food shortage becomes the main factor limiting subsequent breeding numbers. These statements are not true of all birds, but apply to many.

      FIG 4. Changes in the numbers of Great Tits Parus major within four successive years in Marley Wood, Oxford, showing the rise in numbers during each breeding season followed by the decline to the start of the next breeding season. The continuous line joins the numbers of breeding adults each year, and the dashed line follows, respectively, the total of adults plus eggs, the total of adults plus fledged young, and the estimated autumn numbers. (From Perrins 1979)

      In studies of bird populations, it often emerges that heavy losses from one cause at one time of year can be offset by reduced losses from some other later-acting cause. Also, poor breeding can be offset by subsequent good survival, while poor survival can be offset by subsequent good breeding. This is what is meant by population regulation. The idea that different mortality factors are not simply additive but can act in a compensatory manner has proved difficult for some people to grasp, but examples from game and other birds are given later in this book. For such compensation to occur, some aspect of mortality or reproduction must be density-dependent, which means that the extent of reproduction or mortality depends on the density of the population, a concept to which we can return in Chapter 2. It is the main idea on which the concept of population regulation depends. This term has a more restricted meaning than does population limitation, which embraces all the factors that affect numbers, whether or not they vary with density.

      When they are breeding, pairs of birds normally defend some sort of territory, which may include little more than a nest-site in some colonially nesting birds, to a large area also containing food supplies, as in many non-colonial birds. While colonial species, such as the Rook, show clumped distributions, centred on their communal nesting places, other species, such as the Chaffinch, are normally dispersed through suitable habitat, each pair occupying its own territory. Because such birds normally advertise and defend their territories, they are easy to locate and count.

      In the breeding season, some bird populations also contain numbers of non-territorial, non-breeding individuals. This holds for some long-lived species, in which individuals do not breed until they are two or more years old, and in which immatures may make up a large part of the non-breeding contingent. But in many bird populations, some mature individuals may also be found among the non-breeders. This situation mainly arises when more adults are present than the available nesting habitat can hold. Once this habitat is fully occupied, any additional birds must live non-territorially and usually cannot breed. In some such species, such as Carrion Crows or Mute Swans, non-breeders usually occur in flocks, which can be readily seen and counted. Flocks of non-breeding crows sometimes occur on rubbish dumps, while flocks of non-breeding swans often occur on extensive shallow lakes or on rivers near town centres. In other species, including many songbirds and raptors, non-breeders live secretive solitary lives in and around the territories of breeders, or in areas unsuitable for nesting. Such non-breeders are therefore not much in evidence, and in many species they are practically impossible to count.¹ However, two lines of evidence have repeatedly confirmed their existence. Firstly, if territorial birds are removed from their territories, they are often replaced within a few days by other individuals that then proceed to nest (reviews in Newton 1992, 1998). Secondly, in species that are limited by nest-sites, such as many cavity-nesters, the provision of artificial sites (such as nest-boxes) often leads to an immediate rise in breeding density (Chapter 5). The implication is that these incoming birds were previously present in the vicinity but unseen or unrecognised, and would not have nested if opportunities had not been created. In both types of experiment, care is needed to ensure that incoming birds derive from a non-breeding sector, and have not simply moved in from other territories or nesting sites nearby. For a worthwhile experiment, therefore, knowledge is needed of all the breeding pairs of that species present in the surrounding area before any removals of individuals or additions of nest-sites are made.

      Despite the advantages of knowing the numbers of breeders, some bird species are most easily counted in winter. This is the case for many waterfowl and shorebirds which nest in remote or inaccessible places, such as the Arctic, but which migrate to winter at lower latitudes, including Britain and Ireland. The fact that such species often gather in large numbers at traditional sites makes the job of counting that much easier. In some species, adults can be distinguished in the field from young of the year, so that winter counts can reflect both numbers and recent breeding success. In other species, separation of age-groups is not possible except in the hand, so field counts reveal only total numbers. In Britain and Ireland regular counts are made to monitor the numbers of these wintering waterfowl and shorebirds, published in the annual reports of the Wetland Bird Survey (WeBS).

      Other bird species can be readily counted on migration. This is true of some raptors and other soaring birds which migrate by day and in which huge numbers from large areas pass specific concentration points each spring or autumn. There are no established watch-points in the British Isles for raptors, but well-known sites elsewhere include Falsterbo in southern Sweden and Tarifa in southern Spain. Around the British Isles, specific coastal headlands offer good opportunities for counting migratory seabirds. Variations in weather may affect the proportions of migrants that pass specific sites from year to year, so localised counts do not necessarily reflect year-to-year changes in numbers; but when repeated over many years they can reflect long-term trends. Various other constant-effort or observational schemes have been used to detect year-to-year changes in bird numbers, and sometimes it has been possible to compare the results from different monitoring methods on the same populations (Furness & Greenwood 1993).

      POPULATION TRENDS AND FLUCTUATIONS

      If we examine counts of breeding birds in the same area year by year, almost any pattern of fluctuation can be found (Fig. 5). In some species, breeding numbers remain fairly constant through time. Examples of such stability are provided by some birds of prey, such as the Golden Eagle, in which the density of territorial pairs in some large areas of stable land use may vary by no more than 15% on either side of the mean level over several decades (Watson 2010). Most small-bird populations fluctuate somewhat more, perhaps halving or doubling in size from one year to the next, or declining sharply after hard winters, and increasing again in subsequent years. In species which exploit sporadic food suppies, such as finches that rely on tree-seeds or owls that rely on fluctuating rodent populations, the numbers at particular localities can vary by more than 20-fold from year to year, as many pairs settle in years with abundant food, and move elsewhere in poor years. But even these fluctuations are small compared with those in other animals such as certain insects, in which annual changes of 100-fold or more are not uncommon.

      In most bird species, the year-to-year fluctuations in numbers are irregular. But some owls and raptors that depend on cyclically fluctuating rodent populations undergo regular fluctuations of abundance in step with their prey, usually with peaks every 3–4 years. Similarly, some northern gallinaceous birds, such as grouse and ptarmigan, undergo regular cycles of abundance, with a periodicity varying between regions (Chapters 4, 7, 9). The cycle periodicity (length) refers to the regular intervals between peaks, but the size of the peaks often varies greatly from one cycle to the next in the same area (Fig. 5f). The cycle period is not species-specific, and individual species, such as the Red Grouse, have shown cycles of 4–5 years, 6–7 years, 7–8 years and 10 years on different Scottish moors or over different time periods (Watson & Moss 2008). However, different grouse species that live side by side in the same region tend to fluctuate together, so the patterns of fluctuation seem to depend more on local circumstances than on species. Moreover, in some European regions these cycles in gamebird numbers are matched by similar fluctuations in some of their predators, such as the Goshawk and Gyr Falcon, whose numbers typically go up and down in step with their prey (Chapter 4). Yet within suitable habitats, the mean level of abundance of these cyclic species does not necessarily change much over periods of several decades, providing that their habitats remain essentially unchanged. In modern Britain, cyclic fluctuations are evident mainly in extensive stretches of habitat, and much less so in fragmented habitat, such as grouse moors surrounded by grassland or forestry plantations. Perhaps for reasons of habitat fragmentation, cycles are no longer apparent in Black Grouse and Capercaillie in Britain. Also, predator control by gamekeepers is so rife in Britain that it is usually impossible to tell whether specific predator species would fluctuate in parallel with grouse, as they do in the rest of northern Eurasia and in North America.

      FIG 5. Patterns of long-term trends in different bird populations.

      (a) Long-term decline in the numbers of White Storks Ciconia ciconia in Baden-Württemberg, Germany. Based on annual counts of occupied nests, 1950–88. (Redrawn from Bairlein 1996)

      (b) Long-term increase in the numbers of Shags Phalacrocorax aristotelis on the Isle of May, Scotland, showing how numbers increased and then levelled off, but with large annual fluctuations. Based on annual counts of occupied nests, 1944–92. (Redrawn from Harris et al. 1994)

      (c) Irregular annual fluctuations superimposed on a long-term increase in the numbers of Great Tit Parus major pairs in gradually maturing woodland on Texel Island, Netherlands. Based on annual counts of occupied nest-boxes, 1912–43. (From Kluijver 1951)

      (d) Numbers of nesting Grey Herons Ardea cinerea in Britain, showing declines after hard winters (shown by arrows), followed by recoveries. Based on annual nest counts, 1928–92. (Redrawn from Greenwood et al. 1994)

      (e) Weather-driven fluctuations in the numbers of Bearded Tits Panurus biarmicus in southern England. Trends were influenced mainly by the frequency of cold winters (shown by arrows). Based on annual counts of breeding pairs, 1947–92. (Redrawn from Campbell et al. 1996)

      (f) Cyclic fluctuations in the numbers of Red Grouse Lagopus l. scotica, Scotland. Based on numbers shot, 1890–1978. (Redrawn from Hudson 1992)

      Although many bird populations, while fluctuating from year to year, do not normally change much in the long term, such constancy would be expected only in stable environments. Many bird species prefer particular stages in the development of forest or other vegetation. Some species prefer newly planted forest, others thicket stage, and yet others more mature stages. Species of such temporary habitats typically reach high densities in particular localities only for the few years in which their habitat is suitable there, and then decline again as the habitat develops and becomes better suited to other species. But while they decline in one area, they may increase in another, wherever the habitat is reaching a favourable stage. These changes, depending on vegetation succession, can be regarded as natural, although for forest and other habitats, areas at early growth stages have probably become more widespread as a result of human activities. In particular, heather moors, as managed for Red Grouse, represent the first stage in forest succession after a fire, and are artificially maintained in this state by deliberately burning different patches every 10–15 years.

      FIG 6. A Golden Eagle Aquila chrysaetos at winter carrion. In areas of stable land use, where they are not persecuted, Golden Eagles have shown stable nesting densities over long periods of years. (Laurie Campbell)

      For birds of wetlands, the amount of habitat available in some parts of the world varies greatly from year to year, or over longer periods of time, according to rainfall (Chapter 14). This is true for much of Africa, where many of our summer visitors spend their non-breeding season. Usually such populations expand during wetter periods, only to contract again during drier ones, following the trends in their habitats.

      Whatever the type of fluctuation, the degree of variability in any population tends to increase with the span of years over which counts are made. This is partly because the chances of including an unusual year increase with the length of study, and also because short-term fluctuations may be superimposed on a long-term upward or downward trend. In fact, whenever bird species have been studied over periods of several decades, their abundance and distribution patterns are often found to have changed greatly. Some such changes would be expected to occur naturally, perhaps in response to climate or habitat changes, but many can be attributed to human action. In recent decades, many farmland bird species have declined greatly in association with changes in farming practices, as discussed in Chapters 4 and 18.

      Other birds have greatly extended their ranges, becoming common breeders in areas where they were formerly absent. Among land-birds, the most striking example of the twentieth century was the Collared Dove, which until 1930 extended no further west than the Balkans, but then spread rapidly across Europe, reaching Britain about 1955. For many years in newly colonised regions, its numbers increased exponentially (by a constant percentage each year) until habitats became filled and numbers levelled off (Hudson 1972, Hengeveld 1988). The present population of Britain is estimated at around a million pairs.

      Most species that have spread in this way clearly benefit from human activities. Among seabirds, the most spectacular example of range expansion is the Fulmar, which prior to the mid-eighteenth century bred in Europe only in 1–2 colonies in Iceland and on St Kilda off northwest Scotland. A spread started around Iceland at this time, and the Faeroes were colonised in the mid-nineteenth century. The spread continued, and in 1878 Fulmars began to nest on Foula in Shetland. From then on, new colonies appeared successively further south around Britain and Ireland, and the species now also breeds in France, Denmark and Germany. This expansion was linked with the growth of the whaling and fishing industries, which led to much offal and waste being thrown overboard, providing a new food source for Fulmars and other seabirds (Fisher 1966). Initially, overall Fulmar numbers in Britain grew at 13–19% per annum, decreasing to 8% per annum in the mid-twentieth century and to less than 4% towards the end, as they came to exceed 500,000 pairs.

      FIG 7. The Collared Dove Streptopelia decaocto spread westward across Europe in the twentieth century, reaching Britain around 1955, and subsequently increasing to become a common bird of towns, villages and farmsteads. (Frank Snijkers)

      Other conspicuous species have begun to breed here only in recent decades, including the Little Egret and Mediterranean Gull. These are among various bird species that have apparently benefited from climate warming, enabling them to spread northward. As climate continues to warm, we can expect that more bird species will colonise from the south and spread northwards, and perhaps that some other species will retreat and disappear altogether from our islands (Chapter 15). No doubt extreme range changes occurred in other bird species in the past, long before there were ornithologists to record them.

      Within a human lifetime, such spectacular expansions in range and numbers are exceptional, as are precipitous declines to extinction. Yet every species when unchecked has an intrinsic rate of natural increase (r) until it reaches a level (K) at which its numbers become limited by the habitat and resources available locally. If the local environment remains essentially unchanged, the species then tends to remain fairly stable in numbers over the years, fluctuating between limits much narrower than those that would be theoretically possible (Lack 1954).

      The patterns that emerge from studies of bird populations thus depend partly on the timescale over which the studies are made, and partly on the changes in landscapes and food sources that occur in that time. They also depend on the spatial scale. Wide-scale studies of bird populations, involving large numbers of observers, have increasingly enabled the question of spatial synchrony in bird population changes to be addressed. In general, if the factors that influence bird abundance themselves operate over wide areas, such as aspects of weather or human land use, fluctuations on particular study plots can be expected to parallel those occurring over a wider scale. But if the factors are localised, such as some disease outbreaks, then the patterns of fluctuation in bird numbers are likely to vary from place to place across a region.

      ENVIRONMENTAL LIMITING FACTORS

      Bird habitats

      Some bird species are associated with woodland, others with open land of one type or another, and yet others with wetland or sea-coast. In fact, most species are even more specific, being found, for example, in particular stages of forest growth, or in coniferous as opposed to broadleaved woodland, in grassland as opposed to heathland, or in marshland as opposed to open water, and so on. However, some species make use of more than one type of habitat at the same time of year, nesting mainly in one and foraging mainly in another. Species such as the Rook and Woodpigeon nest in trees but feed chiefly on open land, while some upland species, such as the Twite and Golden Plover, nest mainly on moorland but get much of their food from nearby pastures. The requirements of such species are perhaps best described in terms of landscapes or habitat mosaics rather than specific vegetation types, with individuals using different habitat components for different purposes. Seabirds nest on land, mainly on coastal cliffs and offshore islands, but forage widely over the sea, at distances that vary widely between species. Some bird species also use different types of habitat at different times of year, with many waders, for example, nesting inland and wintering on the coast.

      In asking what limits the distributions and numbers of particular bird species, the most obvious answer is that it depends on the amount and quality of habitat available. Recent declines in the distributions and numbers of many bird species in Britain and Ireland can be attributed to the destruction and degradation of their habitats, and consequent reduction of living space. They include, for example, many species of marshland and wet meadows, which now have far fewer places to live than in the past. Conversely, other species have expanded as new habitat has been created. They include birds of conifer forest, such as the Common Crossbill and Siskin, which spread greatly during the last century following the widespread afforestation of former open land, and various waterfowl which expanded in association with the construction of reservoirs and gravel-pits. Other species have increased through targeted conservation actions, the Bittern for example benefiting in recent years from a concerted programme of reedbed creation (Brown et al. 2012).

      FIG 8. The Siskin Carduelis spinus is a small seed-eater which nests in conifer forests. It increased and spread widely in Britain and Ireland following widespread afforestation. (Laurie Campbell)

      Although the habitat tells us where to find particular species, it gives no indication of the densities at which those species are likely to occur, or even whether they will be present at all. These densities may be influenced by the resources (such as food and nest-sites) or natural enemies (such as competitors and predators) that occur there. It is with these limiting factors – which determine the abundance of species within the habitats they use – that this book is largely concerned. As a result of the effects of different limiting factors, areas of essentially the same habitat can hold markedly different densities of birds. The richest conifer forests in Britain can support more than 1,000 songbird pairs per km², but the poorest ones hold fewer than 100. Similarly, although farmland in Britain and Ireland looks on a casual glance the same today as it did in 1960, it now supports far fewer birds, chiefly as a result of changes in management.

      Limiting factors

      To understand how bird populations are limited within the habitats they occupy, it is helpful to distinguish between the external (environmental) factors that influence populations and the intrinsic (demographic) features that these factors affect. External limiting factors include resources (notably food) and natural enemies, including competing species, predators and parasites (some of which cause obvious disease). Any one of these factors can be considered limiting if it prevents a population from increasing or causes it to decline. Particular populations may be affected by more than one, perhaps all, of these different factors, but often one factor emerges as overriding at a given time. These different biotic factors also interact with one another, and with non-biotic factors such as weather. Intrinsic (demographic) limiting factors include the rates of births, deaths and movements, the net effects of which determine local population trends. Both extrinsic and intrinsic factors can be considered as ‘causing’ population changes, the former as ultimate environmental factors and the latter as proximate mechanisms. Thus, within suitable habitat, a population might be said to decline because of food shortage (the ultimate cause) or because of the resulting mortality (the proximate cause).

      It is not just a matter of finical semantics to distinguish between these two types of explanation for population changes. If we are to understand what determines the average level of populations, and why this level varies from year to year or from place to place, we must study the external factors. In our attempts to manage bird populations, it is these external factors that must be altered before any desired change in population level can be achieved. All the external limiting factors mentioned above can have immediate direct effects on bird breeding, mortality or movements, and hence on local numbers.

      Food and other resources

      To be limited by food, a species need not be up against the food limit all the time. Shortages may occur only occasionally, under specific weather conditions, or every few years. Some resident bird species are limited by winter conditions, and may be cut back so severely by lack of food in hard winters that several years of recovery are necessary until numbers reach their former level (for Grey Heron see Fig. 5d). Similarly, many species of arid regions, including British breeding birds that winter in the Sahel zone of Africa, tend to decline from food shortage during periods of drought, only to increase again as food becomes more available during the intervening wet periods. The season when limitation occurs can also change over time as feeding conditions alter, with shortages falling in winter in some years (affecting mortality) and in the breeding season (affecting reproduction) in other years, as in some seabirds (Chapter 4).

      The evidence that certain bird species can be limited by food is mostly circumstantial: (1) long-term or sudden changes in bird numbers often accompany long-term or sudden changes in food supplies; (2) numbers are higher in years when food is abundant than when it is scarce; or (3) numbers are higher in areas where food is abundant than where it is scarce. Additionally, in a few species, experimental manipulations of food supplies have been followed by appropriate changes in numbers, giving firmer evidence for the role of food, as explained in Chapter 4.

      FIG 9. Grey Heron Ardea cinerea, whose nests have been regularly counted in Britain since 1928. The species suffers marked declines in hard winters when fresh waters are frozen, denying the birds access to fish. (Frank Snijkers)

      The relationship between birds and their food is not always straightforward. With certain types of food (such as tree-seeds), the amount eaten in one year may have little or no influence on the amount available the next. But with other types of food (such as many invertebrates), the amount eaten in one year can greatly influence the amount available the next. In this way, prey and predator can interact in various ways, with both short-term and long-term consequences to the population levels of both. Secondly, it is not only the quantity and availability of food that are important, but also its quality, especially for herbivores, whose food can vary greatly through the year in digestibility and nutrient content (Chapter 3).

      Some birds can store large amounts of food as reserves within their bodies, so that (at least in larger species) the food acquired at one time of year can affect their breeding or survival at another (Chapter 3). Other species store substantial amounts of food externally in caches, and can thus survive periods when food would otherwise be difficult to obtain. Examples include those species of tits that store beechmast and other seeds for later use (Chapter 3). While food is the main resource limiting bird numbers, some species can at times be limited by other resources, including water and other components of habitat, such as safe nest-sites and roost-sites (Chapters 5 and 11).

      Interspecific competition

      Most bird species overlap in their resource needs with other species, both closely related and more distant, including other kinds of animals. The seeds eaten by finches, for example, are also consumed by many other creatures, including other birds, mammals and insects. Such food sharing creates the potential for competition, because some of the food removed by one species might otherwise have been available for a second. If either of such species is limited by food, then it follows that the numbers of the one could influence the numbers of the other. Similarly, many species require special nest-sites, such as tree-cavities. If such cavities are in short supply, their use by one species can exclude another, restricting the number that can breed. In any pair of competing species, one species is usually more successful than the other, but the winner in one type of situation may be the loser in another (Chapter 10).

      Predators and parasites

      With or without competitors, food and other resources can provide a ceiling on bird numbers. In some species, however, breeding numbers may be held well below that potential resource-ceiling by natural enemies (Chapters 8 and 10). In some areas, for example, the numbers of Grey Partridges are held at a low level by predators, and when predators are removed, Partridges can increase to achieve much higher breeding densities (Chapter 8; Potts 2012). Similarly, the numbers of some ducks and gamebirds are sometimes greatly reduced by parasites or pathogens, so that in some years their numbers sink well below the level that might otherwise be expected. Because of spatial variations in impact, a predator or parasite can have devastating effects in a small area for a short time, but averaged over a larger area the impact might be small.

      Relationships between resources and natural enemies

      It is often hard to tell whether bird breeding numbers are limited primarily by resources or by natural enemies, without a field experiment in which one or other is manipulated against an appropriate control. Studying the causes of mortality will not necessarily help to identify the limiting factor. Imagine that the density of a territorial bird species is limited by habitat quality or food supply, so that some individuals, unable to obtain a territory, are forced into unsuitable habitat, where they were eaten by predators (the ‘doomed surplus’ idea of Errington 1946). From a study of mortality, one would conclude that predators limited numbers, because virtually all the deaths occurred through predation; however, the underlying limiting factor was habitat quality or food supply, which influenced the density of territories in which survival was possible. To change density in the long term would entail a change of habitat, not of predators. The key point is that the factors that cause most mortality in bird populations are not necessarily those that ultimately determine their population levels. Experiments are often needed to reveal the true limiting factor.

      In any case, assessing the causes of mortality in a bird population is not always straightforward. A bird weakened by food shortage may succumb to disease, but just before death it may fall victim to a predator. For this bird, food shortage is the underlying cause of its death, disease appears irrelevant but potentially fatal, while predation is the actual immediate cause of death. In reality, no single causal factor is likely to account wholly for a given population level. Reproduction and survival are seldom influenced by one factor alone, but by several, which may act independently or in combination. In situations such as these, the main limiting factor can be considered as the one that, when removed, will permit the biggest rise in numbers. Again, this factor can best be revealed by appropriate field experiments.

      Different limiting factors can interact in complex ways. For example, if food were abundant, a bird with many gut parasites might be able to keep itself just as well-nourished as one without parasites, simply by eating more. But if food were scarce, a parasitised bird – through having part of its daily intake absorbed by the parasites – might die of starvation, while one without parasites might survive. In such cases, parasites accentuate the effects of food shortage. Similarly, if food were plentiful, a bird might be able to obtain its needs in only a small part of each day, keeping a constant watch for predators and foraging only in safe sites, thereby reducing its risk of being caught and killed. But if food were scarce, the same bird might have to expose itself for longer each day, reduce its vigilance, and feed in less safe places, thereby increasing its risk of predation. In cases like these, alterations in either food supplies or predator or parasite numbers could have marked effects on bird numbers. Our understanding of the interactions between different limiting factors is advancing year by year, and is discussed in Chapters 11, 12 and 13.

      Weather impacts

      Extremes of weather clearly affect the numbers of birds and other animals. At high latitudes, where life is governed primarily by temperature, the numbers of many resident bird species become lower than average after unusually cold winters, and higher after mild ones. At lower latitudes, where life in arid regions may be limited by water, the numbers of many birds decline during extreme drought years, and recover again in more normal ones. For the most part, weather patterns influence birds indirectly, by affecting their habitats and food supplies. Some such weather effects can be delayed, as when rain stimulates plant growth which only later will provide food for birds, while other weather effects can be immediate, as when a snowfall suddenly makes food on the ground unavailable. Sometimes, however, extreme weather can reduce bird numbers directly, as when individuals die of hypothermia or heat stress or, in some parts of the world, are battered by hailstorms or hurricanes (Chapter 14). Such extremes can sometimes cause sudden catastrophic declines in bird numbers, after which several years are needed before their numbers fully recover.

      Human impacts

      In addition to various natural limiting factors, human activities can impose great losses on bird populations, sufficient to cause widespread population declines. The most obvious impact is through habitat destruction, as forests are felled and marshes drained and converted to farmland or other human use. At the same time, modern agricultural practices are continually reducing the habitats and foods available to birds on farmland. Because the overall numbers of any species depend on the amount of habitat available, destruction of habitat results in reductions in both local and overall population levels. It is not only the overall loss of habitat that is important, but also its fragmentation, as a result of which the distributions of species become increasingly patchy. With only small numbers in each patch, and little or no immigration from elsewhere, it is easy for populations to die out in habitat fragments, reducing their overall numbers below the level expected in the amount of habitat remaining (Newton 1998).

      Other losses imposed by human activity are more direct. Examples include the heavy mortalities inflicted on certain species by excessive hunting or unlawful killing, wind-turbines, overhead wires, fences and fishing gear (Chapter 17), and on other species by various pesticides and pollutants (Chapters 18 and 19). Some pollutants have affected birds and other animal populations over wide areas by influencing the physical or chemical nature of their habitats, the acidification of some lakes and rivers by atmospheric pollutants being an obvious example. Of all these various human impacts on the natural world, ongoing climate change is the one likely to affect the greatest range of species, and to have the biggest long-term influence on plant and animal distribution patterns (Chapter 15).

      DEMOGRAPHIC FACTORS

      Measuring breeding and mortality rates

      Counts of bird numbers, accumulated over many years, are important because they tell us about the fluctuations and trend of a population, which can then be related to changes in environmental variables such as weather. But to understand the internal workings of bird populations, the reproductive and mortality rates must also be studied. Reproductive rates can be assessed either by finding the nests and recording the numbers of young produced by individual pairs, or by measuring the overall ratio of young to adults at the end of the breeding season. Annual survival rates are usually measured from ringing, which enables individuals to be identified in subsequent years. From survival rates, mortality rates can be readily calculated, for when expressed on a percentage scale, both together add to 100% (for example, 60% survival means 40% mortality).

      Some studies in particular areas involve checking to find what proportion of birds present in one year are still there the next. This method does not tell us where or when particular individuals die, nor does it separate deaths from permanent emigration, but it does give a measure of year-to-year persistence in an area. This measure can be close to annual survival in species in which individuals normally use the same nesting places in successive years, and rarely change sites.

      An alternative method of estimating mortality depends on comparing the numbers of individuals ringed as chicks that are subsequently found at different ages by members of the public. From the ratios of birds reported in their first to second year, second to third year, third to fourth year and so on, the mortality in different age classes can be estimated. This method requires that many young birds have been ringed, but does not usually suffer from the emigration problem because it does not depend on recoveries from a single researcher operating in a limited study area. It provides some information on the seasons of death, although there are potential biases in estimates. It also depends on people reporting a sufficient number of the ringed birds that they find, whether dead or alive. Such data have indicated that for most species the main period of natural mortality of full-grown birds is in winter, while for others it is during the breeding season when the birds are most active. Examples of species in which ring recoveries indicate that adults die mainly in winter include Grey Heron and Oystercatcher, and in the breeding season Blackbird and Marsh Tit.

      Ringing has also shown another important point: namely, free-living birds are not like captive birds, or like people in modern societies, where most individuals can expect to reach old age. In wild birds, deaths are frequent across all age-groups, and very few individuals achieve what would be old age for their species. Most do not even survive to halfway through their potential lifespan. Hence, for wild birds, knowledge of maximum longevity is much less meaningful than knowledge of the annual mortality rate, the average age at death, or the average expectation of further life, all of which can be calculated from ring recoveries. Some early estimates from bird-ringing, which revealed annual mortality rates in small birds well in excess of 50%, were met with scepticism, because people knew that in captivity the same species could live for many years. However, the estimates from wild birds turned out to be correct.

      Once estimates of reproductive and mortality rates are available for a population, they can be used to calculate the trend of the population: whether it is likely to increase or decrease, and at what rate. It is unwise to predict far into the future, however, because reproductive and mortality rates often change as a population grows or contracts, so constant re-assessment of demographic rates is necessary. Demographic data gathered nationwide by the BTO have proved useful for studying population processes at regional or national scales where the effects of dispersal can be safely ignored, but less so at local scales where changes in population levels can be greatly influenced by immigration/emigration rates, as well as by reproduction and mortality.

      Movements

      Despite national monitoring programmes, most detailed studies of bird populations have been concerned with the local processes that affect local numbers. Yet almost all species have geographical ranges larger than an observer’s study plot, and individual birds continually move in and out, on local or longer journeys. Such movements can produce much more rapid and pronounced rises in local density than any increase in fecundity or survival. They can also cause rapid abandonment of areas if conditions suddenly become unsuitable. Moreover, as natural habitats become ever more fragmented by human activities, movements are likely to play an increasingly crucial role in maintaining local populations and the genetic interchange between them.

      Movements are important, therefore, because they facilitate rapid changes in local densities in response to changes in local conditions. They enable birds to leave areas where survival or reproductive prospects are poor and to find other areas where conditions are better. In any widespread species, we can expect that in some areas (called source areas) production of young is more than enough to offset mortality (net exports), whereas in other areas (called sink areas) the reverse could hold, so that densities there can be maintained only by continued immigration (net imports). In such sink areas, movements thus become crucial in the maintenance of local densities. Among many bird species, dispersal occurs chiefly in late summer, in the period just after the young become independent of their parents, but in some species also at other times in the non-breeding season.

      Migration is the most spectacular of bird movements, occurring in response to seasonal changes in food supplies, themselves governed by seasonality in climate. Typically, many birds travel over hundreds or thousands of kilometres each autumn to lower latitudes where food suppies remain more available in winter. They then move back to their higher-latitude breeding areas in spring to reproduce on the summer flush of food that occurs there. The species that leave Britain and Ireland completely for the winter include aerial insectivores, such as swifts and swallows, and warblers that eat mainly aphids and other insects from fresh leaves. Birds that come from higher latitudes to winter in Britain and Ireland include species whose food supplies in their breeding areas become inaccessible under snow or ice, such as waterfowl and waders, and passerines that eat seeds and berries, such as some finches and thrushes. These large-scale seasonal movements of birds illustrate how food supplies influence bird densities and distributions through long-distance movement patterns.

      While most migratory bird species move for the winter to lower latitudes in the same hemisphere, some travel longer distances to the opposite hemisphere. Because the seasons are reversed between the northern and southern hemispheres, with the northern winter coinciding with the southern summer, such species gain the advantage of summer conditions year round. They include such impressive migrants as the Swallow and Arctic Tern which breed in the northern hemisphere and winter in the southern, and the Sooty and Great Shearwaters which breed in the southern hemisphere and winter in the northern. As a result of migration, some tropical areas, already rich in resident birds, may receive migrants from the northern hemisphere for half the year (the northern winter) and from the southern hemisphere for the other half (the southern winter). Migration formed the subject of an earlier New Naturalist volume (Newton 2010), so I shall not discuss it further here.

      Use of information on demographic rates

      While knowledge of the external limiting factors is required to understand what determines the level and trend of any bird population, studies of reproductive and mortality rates allow us to understand the mechanics of population change. Remember, though, that two populations may have identical rates of births, deaths and movements, and yet persist indefinitely at quite different densities, if the resource levels on which they depend differ between areas (see below). Conversely, populations at the same density may have different rates of births, deaths and movements. Providing that in each population the inputs (from births and immigration) equal the losses (from deaths and emigration), numbers will remain constant through time, regardless of density. For continuing stability, a change in one parameter must be offset by a corresponding change in another. For example, each year Blue Tits reared up to three times as many young per pair in an area in southern France as they did in another area in Corsica. However, the breeding populations in both areas remained approximately stable, evidently because after leaving the nest the juveniles survived better in Corsica than in France and began breeding, on average, at an earlier age (Blondel et al. 1992). Similarly, Blackbirds in southern England had better breeding success in suburban London than in rural areas (2.0 versus 1.7 young per year), but this difference was offset by differential mortality, so that both populations were balanced in the longer term (Batten 1973).

      FIG 10. The Lapwing Vanellus vanellus has declined greatly in Britain over the past 50 years, associated with a decline in breeding success. (Alan Martin)

      Where reproductive and survival rates have been studied in the same population during periods of increase (or stability) and decline, the comparison has pointed to where the cause of the decline might lie. If reproduction declined while survival stayed the same, the problem would probably lie in the breeding areas, but if survival had decreased the problem could lie in the breeding or wintering areas, depending on when the extra deaths occurred. Long-term decline in the numbers of Lapwings in Britain was associated with a decline in reproduction but no change in survival (Peach et al. 1994), while changes in a Puffin population were attributed to changes in survival rather than reproduction (Harris & Wanless 1991).

      In many other bird populations that have been studied in recent decades, long-term change in breeding numbers has been associated with change in annual survival, while in other populations it has been associated with change in reproductive success (Newton 1998, Saether et al. 2004, Saether & Engen 2010). The latter include several ground-nesting waders and gamebirds which in recent decades have experienced poor breeding, caused mainly by predation (Roodbergen et al. 2012). In other species, long-term population decline was associated with reduction in both breeding and survival. For example, the White Stork has suffered from poor reproduction in its European breeding areas, caused by food shortage resulting from land drainage and pesticide use, and also from poor survival in its African wintering areas, caused by food shortage resulting from drought and pesticide use (Dallinga & Schoenmakers 1989, Kanyamibwa et al. 1993, Bairlein 1996).

      It is not only the long-term trends, but also the year-to-year fluctuations that can often be explained in terms of either breeding or mortality. In some bird populations, breeding numbers increased in years that followed good breeding and decreased in years that followed poor breeding. This was true of Pied Flycatchers, for example, and also of Capercaillie and Black Grouse in different areas (Fig. 11; Virolainen 1984, Summers et al. 2010). It seemed that, in these populations, spring/summer conditions in breeding areas had most influence on subsequent year-to-year changes in breeding numbers, and that such populations were therefore below the level that winter habitat would support. In other bird populations, such as the Great Tit and Swallow, breeding numbers varied from year to year according to previous winter conditions, implying that such populations were close to the level that winter habitat would support (Box 1). However, over several years, the same population could change from one state to another, as its status with respect to available resources and other limiting factors changed. In yet other bird populations, changes in breeding numbers from year to year were associated with changes in both breeding and mortality, which together caused an increase or decrease in numbers. They included species that ate the same type of food year-round in the same area (such as some owls dependent on voles, Chapter 4) and other species in which changes in breeding numbers were affected by conditions in both breeding and wintering areas (such as the White Stork, mentioned above).

      FIG 11. Demography and population change. (a) Relationship between the annual survival of adult Swallows Hirundo rustica and annual change in breeding density (from Møller 1989). (b) Relationship between the annual breeding output of Pied Flycatchers Ficedula hypoleuca and annual change in breeding density (from Virolainen 1984). Significance of relationships: Swallow, b = 3.44, r² = 0.93, p < 0.001; Pied Flycatcher, b = 0.15, r² = 0.74, p < 0. 001.

      Sometimes a long-term upward or downward trend in a bird population can be caused by different factors to those that affect the year-to-year fluctuations about the trend. For example, over the latter half of the twentieth century, the Dark-bellied Brent Geese wintering in western Europe (including Britain) increased in numbers by more than tenfold, in recovery from over-hunting in the past. Nevertheless, winter numbers still fluctuated substantially from year to year, as a result of annual variations in breeding success in the Siberian nesting areas, caused mainly by fluctuating levels of predation on the eggs and chicks (Chapter 7). Hence, in this population the long-term trend could be attributed to conditions in wintering areas, and the short-term fluctuations to conditions in breeding areas.

      Without information on trends in breeding numbers, one cannot judge the effect on a population of a decline in either reproduction alone or survival alone. Decline in the production of young does not necessarily lead to population decline, because it may be offset by improved survival. Decline in survival does not necessarily lead to population decline, because it may be offset by improved reproduction. One species that increased during a period of declining adult survival was the Kittiwake (Coulson 2011), and some that increased during periods of declining reproduction were the Wigeon (Mitchell et al. 2008) and the wild geese discussed in Chapter 2.

      Studies of population limitation are especially difficult in migrants that divide each year between breeding and wintering areas far apart, because conditions in either or both areas can affect their breeding numbers, as in the Brent Geese just discussed. In recent years, many species of migratory birds that breed in Britain and Ireland have declined. These declines seem to have been both more prevalent and more marked among species that winter in Africa than in those that winter in Europe (Sanderson et al. 2006). Europe-wide trends in breeding numbers between 1970 and 1990 were analysed in 30 pairs of closely related species. In each pair, one species wintered in Africa and the other in Europe (as in Tree Pipit and Meadow Pipit, Whinchat and Stonechat). Significantly more negative trends emerged in the African-wintering species, regardless of breeding habitat. This is perhaps not surprising, considering the rate at which habitat in Africa is degrading under human influence, but more study is needed to understand the details. European breeding birds that winter in the Sahel zone of Africa declined to low levels during the period 1969–76, when rainfall there was low, increasing afterwards. However, from about 1987 to the present, as these species recovered, other species that winter in more humid habitats further south in Africa began to decline, with the causes yet to be established (Thaxter et al. 2010). Nevertheless, it seems that both groups were at the time limited primarily by conditions in their wintering areas.

      LIFE-HISTORY FEATURES

      Birds span a wide range of life-history types, from short-lived, mostly small-bodied species to long-lived, mostly large-bodied

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