Landscape changes and the vertebrate fauna in Sweden
during the last 150 years
by
Kai Curry-Lindahl
Zoological Department, Nordiska Museet and Skansen, Stockholm
The Swedish the of the such that the fauna countryside at beginning nine- It was to a countryside was adapt-
teenth still varied and rich that there ed. what fauna? and the century was so But By large same as to- for both human and animals almost but much richer. What was room beings day, quantitatively now re-
It is mains animal life of secluded everywhere. true that for thousands of years of the certain woods,
had with the forests. First the few and wooded and man struggled advancing surviving glades meadows,
the deciduous forest cleared the in meadows and shallow lakes rich in primaeval was during water vegeta- the normal fauna of Sweden warm period following the last glaciation, and after tion, represents yester- the the close the Swedish and their great climatic change at of day. Human activities variations were not 700 the and in those Man Bronze Age (ca B.C.) struggle was against unexpected abrupt days. as a
in the the encroaching spruce on practically all fronts. The biological being behaved almost same way as
resulted culture birch other animals. Man's influence the struggle in steppes, pine heaths, large on vegeta-
and wooded meadows. All these of land- tion than that of the wild ani- groves types was naturally greater
scape were gradually filled with the fauna adapted mals, but nature's circles were not disturbed to any
cultivation alter- considerable The of to them. Different types of have degree. doings animals, including
and the has the fauna has Scandinavia to the of the nated, as nature changed, man, in up opening nine-
itself. But to the nineteenth these teenth could be fitted into the adapted up century century automatically natural the changes were not so great that they altered the gen- rhythm. Everything went on calmly; eral of the animal world. that could picture changes were not so great nature not im-
Sweden and half herself them. Imagine a century a ago! Spruce mediately adapt to then but less With the of forests dominated as now, they were expansion industry, the far-reaching uniform; they had kept their individuality and char- consequences of the reckless draining of marshes and acter. Wooded meadows were numerous, for the wet ground, the shortsighted, ruthless exploitation of
leaf harvest that created them of the rivers for the regular was great hydro-electric power stations, grow-
the former that value of the coniferous Swedish importance in agrarian economy. In ing forests, nature,
fashioned in way the spruce was kept at bay and the wooded which had been by thousands of years meadows could and there the farmers and for develop freely, a harmonious, most living creatures accept- could collect leaves for winter fodder. The young able way, was transformed violently.
Swedish soil was then still virgin. Thousands of lakes, Let us return to the nineteenth century. A few and the animal world of will marshes fens, the inheritage of the great ice-cap, glimpses.of that century filled the the the show the differences between then and countryside. Through country ran now. Only a silver bands of and brooks. In and half of Sweden glittering streams century a ago great parts were spring they widened and at times the water flooded still wilds, where bears, wolves and lynxes roamed in
meadows. As the the where the of the birds from the the summer approached, low-lying woods, songs rose alluvial fields of horsetail and rush wooded where the still of the sedge, were trans- green meadows, water
the marshes the flocks of birds formed by sun into natural meadows, which were reflected migratory flying
make of that could and where there rivulet mown yearly to use everything over, was hardly a mountain
fodder for the livestock the without serve as during coming flashing trout. winter.
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1. Vegetational map of Sweden.
▬ Southwestern limit of = Picea abies (granen)
▬▬▬▬▬ = Northern limit of Fagus silvatica (boken)
robur ▬ = Northern limit of Quercus (eken)
─ = Northwestern limit of coniferous trees (barrträd).
(These limits are indicated on the map from south to north).
The wolf which and The arctic fox (Canis lupus), had been retreating and common in Blekinge Smâland. northwards the and 1832 during eighteenth century, made a (Alopex lagopus) was a vagrant, in came sally southwards in the nineteenth, and increased in from the mountains right down to southernmost Swe- numbers. It could still be found the middle of the later the in den; nine years same thing happened again. of the arctic fox century in Scania, in southernmost Sweden, where it This rhythm in the frequency curve
the which is was even seen roving over plains. The lynx (Lynx suggests a peak every ninth or tenth year, no
this animal, lynx), however, had left Scania, but was still resident longer observable in now so uncommon
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The most southerly haunts of the bear during the first had practically exterminated the elk (Alces alces) and
half of the nineteenth in Bohuslän and in Sweden. Wolves had century were roe (Capreolus capreolus)
and still Värmland and but their destruction of Dalsland, it was common in done their share, of course,
Dalarna and farther north. In Närke, Vastmanland these two deer could not possibly have led to such
and possibly also northern Uppland, the bear (Ursus disastrous results. It is usual nowadays to explain the
in arctos) was still resident, while Värmland the wol- rapid expansion of elks and roes as a consequence of verine (Gulo gulo) still scampered about. In the the extermination of the wolf. Human hunters still
the had the last of the and if than mountains Lapps liquidated remain, even they are more enlightened
wild Scandinavian reindeer which it be their conti- (Rangifer tarandus), they were a century ago, can hardly
is be found in the absence of wolves that has nowadays to only the Norwegian moun- nence with the gun or
With that the wolf lost its most elks increased in Sweden. There tains. important given and roes scope
source of winter food. must be other contributory factors.
This is the background of the present situation of the carnivores in Sweden. In completely changed The golden oriole(Oriolus oriolus), the hoopoe (Upupa
driven habitat less suitable and the black stork built their surroundings, or into a epops) (Ciconia nigra)
the island of Öland for them, they are compelled to satisfy their biological nests in the groves at Ottenby on
roller flew requirements, of which food and shelter are the most in the Baltic. The (Coracias garrulus) on essential. iridescent wings through the leafy woods of Scania.
At of the nineteenth On the Scanian sand-fields round and Ahus the beginning century man Ljungby
could be heard the bustard mighty great (Otis tarda).
The hollow of the owl cry eagle (Bubo bubo) was
heard all over Sweden, from Scania to Lapland. The
the white-tailed kite (Milvus milvus) and sea eagle
(Haliaeetus albicilla) built their nests at Lake Horn-
borgasjön and other places, where the great snipe
(Gallinago media) was common on all wet ground
round the lake. Not only at the rich bird-paradise
of which later be Hornborgasjön, was to destroyed
by a series of outrages and a period of culpable
neglect, was this now almost extinct snipe to be
found; it also in and was numerous Uppland very
common in Norrland.
The voice of the bittern (Botaurus stellaris) had
been silenced in the reeds, however. For some un-
known reason it disappeared from Sweden as a resi-
dent bird during the nineteenthcentury. Perhaps ex-
contributed ceptionally severe winters in Europe to-
wards its disappearance.
Displaying capercaillie (Tetrao urogallus) were a
the common sight in Swedish woods in spring, and
the black-cock out on ice and in the bogs leking
(Lyrurus tetrix) bubbled their mating call. At that time the leking places attracted gallinacious birds in
numbers than and still greater we can imagine, they
had already begun decreasing in number. Except for
small fluctuations this trend has continued.
Llewellyn Lloyd (1831) wrote at the beginning
of the nineteenth century: 2. Map showing the distribution of the eagle "In many woods and districts where, fifty years owl (Bubo bubo) in Sweden 1943-1948. Each abundance of both and black dot represents a breeding pair. After K. Curry- ago, capercali game be bird the Lindahl 1950. were to found, not a now exists. In spring,
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assemble for the of when the birds purpose pairing, people place themselves in ambush, and shoot with- out distinction cock and hens, by which means the
and birds are frightened and dispersed; afterwards,
and the when the spring is more advanced, young are after hatched, it is certain the old hen will be sought before able shot whole they are to fly; by one a brood of birds thus seven or eight are destroyed..
This refers to the situation in Värmland in the
Thus the decrease in birds years 1827-1828. game
have the Värmland forests seems to begun in during
the latter half of the eighteenth century.
During the nineteenth century it was not only the of and forest former incredibly rich population game birds that began to fail. The passing-bell tolled for
of Sweden's fauna many large representatives during
that far the southern of the century, as as part country
was concerned. The bear and the wolf were driven
northwards. The latter also from central was ejected
Sweden. Owls and birds raptorial were persecuted
with the result that have mercilessly, many species become very rare, or were exterminated completely
Sweden in some provinces. But has not yet wholly lost of owl bird of which a single species or prey we know has bred in Sweden in historical times. Here
in the north man has had such a short time at his
disposal that he has luckily not been able to extermi-
nate a nocturnal or diurnal bird of This, single prey.
however, has occurred in many other European coun-
tries, where the two-legged mammals have had a few
more thousand at their for their work years disposal 3. Distribution of the elk or moose of destruction than in Sweden. (Alces alces) in Sweden 1953. The
estimated number of moose in 1953 Animals have been driven or have otherwise dis-
was about 90.000 animals. Formerly appeared from Sweden in the course of centuries, but the moose was dispersed over all forest others have arrived. Human beings have introduced bearing parts of Sweden, but at the be-
the fallow-deer Dama the brown hare ( dama), (Lepus ginning of the 19th century the species europaeus), the wild rabbit (Oryctolagus cuniculus), had decreased and was limited to a
rather small area in Sweden, where the pheasant (Phasianus colchicus), the rainbow-trout the population of today is most dense. (Salmo gairdneri), and others. Many species, how- Through total protection in 1808-1817 have to Sweden of their accord, main- ever, come own and 1826-1835 respectively, the moose
for it is for them to saved from extinction and began ly birds, naturally, easier occupy was to spread again. During the past hun- and colonize new domains. Many southerly sea-fishes dred years the species has continually (Ze us faber, Brama raji, Mullus surmuletus, Thunnus increased. The staple diagrams reflect
also more in Swedish thynnus) appear frequently the number of elks obtained at two
The harvest The waters. mouse (Micromys minutus) is censuses in 1945 and 1953. curve
shows the harvest, that in 1959 was waiting in Denmark for transport over the Sound, for 32.286 animals. After K. Curry-Lin- it cannot cross under its own power. A flying mam- dahl 1957. mal, the serotine bat (Eptesicus serotinus), has for the
time being halted at the Danish side of the Sound, Perhaps the most remarkable events during the
but later will and settle the of the sooner or it cross over in present century are enormous increase elk,
Sweden. Several animals (Mustela lutreola, Tarsiger the unparallelled expansion of the roe northwards,
cyanurus) can also be expected from the east. and the re-appearance of the lynx as a resident of
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Distribution of the 4. changes mute swan (Cygnus olor) in Sweden from 1850 tot 1955. After K. Curry-Lindahl 1958 B.
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Smâland and some" other southern provinces. There
birds of different are many species (Streptopelia decaocto, Phylloscopus trochiloides, Muscicapa parva,
Serinus canaria, Carpodacus erythrinus) on their way northwards number of them to new ranges. A are quite recent additions to the Swedish avian fauna, and almost birds to Sweden arrive to every year new
Scandinavia. Not infrequently these strangers are pi-
of which at first be oneers a coming expansion may
it until at favour- slow, tentative as were, last, some able spring with suitable climatic conditions, it sud-
and number of denly explodes rapidly occupies a bridgeheads.
Among the birds that have spread rapidly during the present century are the starling ( Sturnus vulgaris), the fieldfare (Turdus pilaris), the marsh harrier (Circus
the haliaëtus), the her- aeruginosas), osprey (Pandion
6. The distribution of the coot (Fulica
atra) in Sweden corresponds to the occur-
rence of eutrophic lakes. After K. Curry-
Lindahl 1960 B.
the on (Ardea cinerea), the mute swan (Cygnus olor),
tufted duck (Aythya fuligula). the great-crested grebe
(Podiceps cristatus), the Slavonian grebe (P. auritus),
the wood pigeon (Columba palumbus), the common
curlew (Numenius arquata), the black-headed gull
(Larus ridibundus), the herring gull (L. argentatus).
the water rail (Rallus aquaticus), the moorhen (Galli-
nula chloropus) and the coot (Fulica cristata). It will
be noticed that many of the birds on their way north-
birds of lakes rich wards are water typical in nutri-
that this be ment. It is possible expansion can ex-
the of lakes and plained by progressive drying up
marshes in the southeastern steppe regions, at the northern has same time as the temperature in Europe These climatic fluctuations risen. are probably a
repetition of similar earlier phenomena. We are no
doubt concerned with long, rhythmical, secular cycles 5. Distribution ranges in Sweden of the the animal whimbrel (Numenius phaeopus) (in the which affect birds most noticeably in northwest) and of the curlew (N. arquata) world.
central and north- are only overlapping in of Reptiles and amphibians, in spite a rising mean western Sweden, but the latter species is be temperature in the twentieth century, seem to re- expanding following the cultivated val-
somewhat. These creatures to leys in the northern part of the country. treating belong groups After that life latitudes. K. Curry-Lindahl 1955. are mainly adapted to in warmer
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The northern occurrence of certain species in Europe
must be viewed against the background of the warm, post-glacial period, when they immigrated and reach- then, with the ed their present ranges. Since excep-
of few to have been tion a species, they seem re- treating steadily southwards.
Shoals of tunas are becoming more and more com-
Swedish off the mon in waters west coast, as are
those of other fishes with a southern distribution.
the half of the twentieth the During first century
population of Sweden increased by nearly two million
in people. In 1950 there were twice as many people
had been earlier. the country as there a century But there million seventy years ago were a more people
living in rural areas than there are now. Further,
enormous technical developments, which have trans-
formed both the and the of life, have countryside way
taken the of time. This place during same space proc-
ess has in its turn also caused an increase in the num-
bers of several species of animals that have been able
to adapt themselves to, and have been favoured by,
the new conditions. The fox (Vulpes vulpes), the
blackbird hooded crow (Corvus corone), the (Turdus
and the few of these merula) herring gull are a spe-
Others have been less but cies. adaptable, a sur-
able prisingly large number of animal species seem to
side side with Sweden—if live by man in presentday
allowed do It often man they were only to so. is most
who prevents such co-existence.
distribution animals The present of is, as implied
due historical but it is not above, largely to reasons,
the habitat that alterations only changes in causes in
the distribution in one or another direction. The
of the its its adaptability species, power to compete,
dynamics, that is to say the characteristics of the spe- 7. Distribution of Sweden’s three species of limits also of The snakes, showing the northern of ranges. cies, are great importance. continually
— Coronella austriaca (= hasselsnok) is a favour changing ecological conditions some species, southern element, which probably invaded but handicap others. An example of this in ovir days Sweden from the south via Denmark, from
is the of the the and records known; the last increasing frequency elk, roe, where only eight are
1914. several birds. On the other hand we have the retreat was made in Except in the Åland archi-
—it does not occur in Finland. Natrix of the wolf and the decline of the crested lark (Gale- pelago natrix (= snok) is thermally intermediate be- rida cristata), the white stork (Ciconia ciconia) and tween the smooth snake and the adder, which the fire-bellied toad (Bombina bombina). —is also shown by its distribution area. Vipera rôle of berus The man in nature is of great significance (= huggorm) is less sensitive to cold and northern well in and factor. Man occurs in areas as as birch as an ecological Zoogeographie has, forest region. This species may have arrived in- more than any other species, either or directly south well from the as as from the east, which transformed directly, consciously or unconsciously, is not excluded for the ringed snake either.
of his in short time. K. Curry-Lindahl, 1958 A. great areas surroundings a very After In Sweden this transformation has covered practically
the whole of the country; only parts of the mountain It is not only by extensive changes of habitats in ranges and the seas have hitherto remained un- the form of cultivation, draining, changes of water
changed. level for hydro-electric developments and the like
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8. The decrease of the white stork (Ciconia ciconia) in Sweden in 1900-1954. Dots indicate breeding pairs. The two
nestlings of the only Swedish nest in 1954 died halfgrown. No breeding in 1955 and later. After K. Curry-Lindahl 1957.
rather that man has altered the possibilities of existence for however, the temperature curve has been er- animals. also the of the Small fluctuations have occurred He interferes directly in size ratic. in a largely
one-sided and climate. Thus the between about 1860 populations by hunting, continuous warmer period
subduction of oil in the and 1920 had maritime trend with mild winters and predators, pollution seas, a
cool After that climate became which at one blow may exterminate whole popula- summers. the more
the introduction of continentalwith cold and but tions of wintering aquatic birds, winters warm summers,
the of insecticides and the like. the be the direction foreign species, use now tendency a,gain seems to in
All these start an almost endless chain reaction in the of a maritime climate. These small variations seem
affected, un- considerable in the animal habitats and the consequences may be to have caused changes
and unfortunate. could be such mobile as expected Many examples world, particularly among species
given. birds, sea-fishes and winged insects. A new advance
towards the north has been observed among southerly
In the of the thousands course of years that have species, while northerly species are moving in the the last climate has fluc- elapsed since glaciation, the same direction, but this tendency is not general.
tuated in birds winter in their mighty, many-thousand-year cycles. During More and more breeding quar-
times it historic has been possible to register the minor ters, and for other species the winter quarters extend
fluctuations in Sweden. The twelfth, thirteenth and farther north.
fourteenth have been climate far from Scandinavia centuries seem to relatively Changes in may cause
the 1400 north of birds mild; period to 1700, on the other hand, an immigration into Europe compelled
colder. At the seek habitats for because of an considerably present temperature to new reproduction
shows trend. the hundred in their a rising During past years, increasing aridity original range.
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it Cause Habitat A landscape is dependent on climate, and may Species Category
climatic therefore be said that faunal changes due to Canis lupus — Ha F + A N factors also caused the habitat. Vulpes vulpes Ha G St are by changes in +
Alopex lagopus — Ha A St In an analysis of the variations in the frequency of
Ursus arctos — Ha F + A St of the land- animal species, therefore, the changes Meles meles + E+Cl F St
due to man, or to causes F scape originating directly Gulo gulo — Ha A + N
F from the actions of must be from — Ha N man, distinguished Lynx lynx
Phoca vitulina — Ha C St those caused by climatic ecological factors in which
P. hispida — Ha C St man has played no part.
— St Halichoerus grypus Ha C
Capreolus capreolus + Ha + Cl + E F+ P St
In the following discussion, consideration has been Cervus elaphus — Ha F St
F St paid only to those vertebrate species which, during Alces alces + Ha + E Mi — Ha A of have shown either Rangifer tarandus a long period observation, a clear but increase or decrease, not only in frequency, Podiceps cristatus + E W Mi above all in distribution. In this geographical con- P. griseigena + E W Mi introduced nection animals deliberately by man are P. auritus + E W Mi
that took Ardea cinerea + E W Mi ignored completely, as are changes place in Ciconia ciconia E Hs + P I W Mi prehistoric times. Aythya fuligula E W Mi During recent centuries, ninety-one species of A. ferina E W Mi Swedish vertebrates have shown fluctua- — significant Clangula hyemalis E+ Ha? W Mi tions in frequency. Of these forty-five have increased, Somateria mollissima + Ha C Mi Tadorna tadorna E C Mi while the other forty-six have decreased (Tablés I
— ? C W Mi Anser anser + in in and II). Birds are the majority both groups, A. erythropus — Ha A Mi with and decreas- thirty-five species increasing thirty A. f abalis — Ha + E? F+ Ma Mi
— Mi ing. The equivalent figures for mammals are five and Branta bernicla E C E Ha? W St thirteen. Cygnus olor + +
— Ha F Ma Mi C. cygnus + — Aquila chrysaëtos Ha F+A St+Mi Table I. Variations in distribution and frequency N Accipiter gentilis — Ha F Symbols Milvus milvus — ? F +W Mi
+ = Increase Haliaeetus albicilla — Ha W+C St+Mi
= Decrease J- Circus aeruginosus Ha? +E W Mi
Partition haliaëtus + E+ Ha? W Mi Abbreviations for causes
Falco peregrinus — Ha M Mi CI = Climatic factors
F. rusticolus — Ha A St+Mi E = Ecological factors except man and climate — Lyrurus tetrix E + Ha? F+ Ma St Ha = Human activities
Tetrao urogallus — E + Ha? F St
Abbreviations for habitats Rallus aquaticus + E+Cl W Mi
= Arctic and subarctic P Mi A regions Crex crex Ha
C = Coasts Gallinula chloropus + E+Cl W Mi+St
- Mi F Forests Fulica atra + E+ CI W St+
= Hs Human settlements Otis tarda — Ha+E? P Mi + St
Mi Ma = Marshes Haematopus ostralegus + E C +W
M = Mountains Vanellus vanellus E W + P Mi
P = Plains Charadrius dubius + Ha C + W+Hs Mi — S = Sea Gallinago gallinago Ha W+A Mi
U = Übiquitous species G. media — ? W +A Mi
~ Mi W Watercourses (lakes, rivers) Numenius arquata + E W+P
Philomachus pugnax — Ha W+A Mi Abbreviations for categories St Larus argentatus + Ha + E+ Cl C+W Mi + N+ Mi Migratory L. ridibundus E W+ C Mi + St
N = Nomadic Chlidonias nigra — ? W Mi
St = Stationary Fratercula arctica — ? C Mi
Mi Columba oenas Ha? F+C Species Cause Habitat Category C. palumbus + E F+P+Hs Mi+St
F St decaocto + E P St Lepus timidus Ha Streptopelia
— St Castor fiber Ha W St Bubo bubo Ha F + C + A
St Coracias — Cl F Mi Rattus rattus E Hs garrulus
cristata — St R. norvegicus + E Hs St Galerida E Hs + P
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Species Cause Habitat Category Table III. Probable causes of increase or decrease
— St Corvus corax Ha F+C Changes in land- Climatic and
C. corone cornix + Ha G Mi + St scape due to man other ecological
C. monedula E P+F-LHs St or man's direct changes (except
Garrulus glandarius + E F St actions man's rôle)
Turdus pilaris + E F +A N +Mi
T. musicus E A + F M Mammals increase 3 2
merula E F Hs T. + + Mi+St decrease 12 1
Phoenicurus ochruros E Hs Mi Birds increase 4 27
E Cl? F Hs Mi Luscinia luscinia + + + decrease 12 7
Locustella naevia E W Mi + Fishes increase — 5
E W Mi Acrocephalus scirpaceus + decrease 2 —
Sylvia borin + E G Mi
Prunella modularis E F+A Mi Total increase 33 Anthus campestris — E C + P Mi decrease 26} 1}«
Sturnus vulgaris + E + Ha? G Mi
Carduelis carduelis — ? FJ-P N + Mi
Emberiza calandra — E P St of There are several species of different vertebrates
Bombina bombina — ? P St which it is to whether at present impossible say — Salmo salar Ha W Mi human other factors for the or are responsible
S. trutta — Ha W Mi changes now going on or already completed. One Lucioperca sandra E W+ C St+N of these is the bustard, now extinct in Sweden. Mullus surmuletus E S St great Table number within Thunnus thynnus E S Mi IV shows how the of species
Scopthalmus maximus E S Mi the different classes of animals have shared the in-
Pleuronectes platessa + E S Mi + St whether crease or decrease in various habitats, and
nomadic. are resident, or Many of the for the de- they migratory spe- An analysis reasons increase or than of the bi- cies of animals belong to more one the different shows crease in species (Table III) that, the wood which is otopes mentioned, e.g. pigeon, as a result of changes in the landscape caused by found the on the round in woods, in groves plains, man, or other factors directly traceable to man, seven This human settlements, and now even in towns. species (three mammals and four birds) have been bird is increasing in all these habitats. Other animals, while fewer than favoured, no twenty-six species the advan- e.g. pike-perch (Lucioperca sandra), are (twelve mammals, twelve birds, and two fishes) have in one of the habitats, the coast, and found cing chiefly only their habitats and ranges greatly reduced; one the table under that are thus represented in only type species has been exterminated (the wild reindeer) and of the environment. another has been re-introduced after extermination number of Only three habitats have an increased (the beaver, Castor fiber). watercourses, and covered The species, viz. sea, areas, consequences of other ecological factors, chief- by ubiquitous species, which include habitats found ly climatic, have been quite different, for they have all over the favoured fewer than country. no thirty-four species (two mam- The distinction between the human and other eco- mals, twenty-seven birds, and five fishes), while only the fauna logical factors causing changes in is an eight species, (one mammal, and seven birds) have artificial account of the fact that one, not only on declined in frequency, and their geographical distri- man himself is a creature included in the bution has become restricted. biological in the animals, but processes of nature same way as
in affects the also because he, many ways, directly Table II. Variations in distribution and frequency size of animal populations. When man, with mis-
Increase Decrease guided zeal, reduces the number of predators, for
he example, may cause positive or negative conse-
is for other If a extermi- Mammals 5 species 13 species quences species. predator nated reduced numbers in Birds 35 species 30 species or greatly in one district,
Reptiles — species — species for time favour other but the next it may a species,
Batrachians — 1 species species for food between the step is frequently competition Fishes 5 species 2 species wished individuals of the species that humans to help,
another to in- Total 45 species 46 species or predator is given an opportunity the eliminationof crease unhindered (e.g. the fox after
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Sweden has a reliable population of beavers. In some
places, however, the beaver has again been reduced
which as a consequence of water regulation, against
all the constructional skill and industry of the four-
footed engineers are in vain, particularly as short-
period regulation, for hydro-electric purposes, usually
takes place during the winter, when the dams made
their by the beavers to protect dwellings are damaged washed and their stores of food away.
than the wolf to be Rather more 100 years ago was
found all over Sweden. During the period 1827-
wolves killed 1839, 6,790 were in Sweden, eighty-two
of them in Scania and 271 around Stockholm. As
late there wolves north of as 1883 were in Uppland,
showed the Swedish capital. In 1945 a State census
that the number of wolves in the country did not
exceed low twenty-five, a figure, but, however, prob-
ably much too optimistic in view of the wolf's low
The wolf Sweden's reproduction figures. is now most
and of the uncommon carnivore, is obviously verge
of of the 9. Extension range badger
(Meles meles) in Sweden from
1850 to 1955. After K. Curry-
Lindahl 1960 A.
inter- the wolf and the lynx). Thus practically every
ference by man that changes the character of a bi-
ocoenosis begins a chain reaction of inverted habitat
conditions for animals and plants. An extremely good
example of this is what happened in Europe after the
extermination of the rabbit by myxomatosis.
detail the Let us study in greater thirty-three species of vertebrates whose of life has been way changed by
the activities of man. We will begin with those that have been unfavourably affected.
The mountain hare (Lepus timidus) was driven
from the southern plains of Sweden by the brown
introduced and it is south of hare, by man, now rare
the northern boundary of Scania.
Formerly the beaver was to be found by many of
all the watercourses of Sweden, probably over the 10. Distribution of the brown bear exterminated the last known country. It was by man, (Ursus arctos) in Sweden. Arrows being shot in Jämtland in 1871. In 1922 specimen indicate dispersal trends during the into the first beavers were reintroduced the same K. last years. After Curry-Lin- followed and province. They were by others, now dahl 1960 A.
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Table IV. Resident, migratory and nomadic species of increasing or decreasing vertebrates,
distributed according to habitat
belts and lakes General Human Sea Coasts Bogs Forests Plains Migratory Nomadic distribution settlements Mountains Arctic subarctic Watercourses & Stationary
Mammals
increase 1 1 0 0 3 1 0 5 0 0
decrease 1 4 3 6 0 1 9 1 3
Birds
increase 2 2 0 3 5 0 8 4 19 10 31 2
decrease 2 1 4 6 3 11 7 10 6 24 1
Batrachians
increase 0 0
decrease 1 1
Fishes
increase 4 1 0 1 3
decrease 0 0 2 0 2
Total
increase 3 3 0 3 4 6 0 11 5 19 15 34 2
decrease 3 1 8 0 9 3 17 8 13 16 27 4
of If for which had been absent for half This extinction. it were not a continuous im- they a century.
from the would no be a caused of migration east, it longer movement is not by a pressure over-
member of the Swedish fauna. population in the mountains.
Thus the wolf direct is a species that, owing to During this movement the bear has penetrated as
has had the the the Niemisel and Boden persecution by man, to retreat to most near coast as districts,
inaccessible of the which other that it is not the parts country. shows, among things,
The arctic fox was formerly to be found everywhere cultivation of the countryside but direct persecution the and in mountain range. Shooting poisoning were by man that has restricted the geographical distribu-
the of the reduction of bear Sweden. the Sarek most probably main causes tion of the in Nowadays the of of National where species. In spite a long period protection it Park, hunting is prohibited (except has been unable to regain its former frequency. for Lapps), with the surrounding forest land, is the
During the past three centuries bears have been best bear country in Sweden.
withdraw from the of their compelled to greater part Despite bitter persecution, the wolverine (glutton) former Swedish reduced has succeeded well former range. They were in num- remarkably in keeping its bers and the driven the distribution the survivors were to mountains. area in mountains, although a con-
In 1925 the bear was protected on Crown lands. This siderable reduction has occurred in the forest land.
of lasted till and In it is period protection 1942, seems to this case, too, direct persecution by man that have led stabilization of the the been decisive. The wolverine fact to a population in has is in primarily
the af the for it in the forests mountain areas, in valley Vindel, ex- a mountain species, although occurs ample, and parts of the Lule Lappmark, where the too. populations were good. In 1942 hunting was permit- ted for two months annually, and in some mountain districts the bears reduced number. The has suffered form were greatly in lynx, too, persecution by man.
At the end of the 1940s and the 1950s bears forest and reduction has been during It is a dweller, its rapid
hunted down have wandered down from the mountains to the in Sweden since the species was merci-
the decade coniferous forests, and thus returned to places from lessly during the nineteenth century. In
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the whole of the southern and central parts of Swe-
Dalarna den except for the northernmost parts of and the wooded districts of south-western Västergötland
the borders of where on Halland, it disappeared
about 1906. At the same time the northern limit had
moved northwards, but not far enough to compensate
the animal for its lost territory in the south. In 1925
the permanent distribution area was at a minimum,
and almost in In 1928 the was wholly Jämtland. lynx
the whole of Sweden. was protected in There were
then possibly a few left in Jämtland, but as Einab
have been Lönnbebg wrote in 1930, they must very
few. The same writer stated at the same time that
the lynx had been practically exterminated in Väster-
bottens län.
Unless these low estimate of the reports give a too frequency of the lynx in Norrland in 1925, the protec-
of the the nick of time. tion species came in During
fifteen years of immunity from hunters, the lynx in- creased and the limits of moved slowly, its range were
northwards and eastwards. No movement southwards,
which would have been natural for this more species,
seems to have occurred, however. But unexpected
made Dalsland finds of lynx were in during the 1920s
and 1930s. It is uncertain whether these isolated oc-
currences were recruited from the few lynxes in Nor-
whether the has been way, or lynx never really ex- 11. Regression of distribution of terminated in Dalsland. the lynx (Lynx lynx) in Sweden
from 1850 to 1900 and increase In 1943 lynx-hunting was allowed for two months
from 1925 to 1951. Circles = con- in Sweden. At about the same time finds of lynx centration of records during 1940- from were reported along a passage through Uppland, 1954. After K. Curry-Lindahl Södermanland, and and in 1958 A. Östergötland Smâland,
south-western Sweden it was met with in southern
Dalsland, Västergötland and western Smâland, where
it is still resident.
1827 2.506 shot the In of the fact that the has increased to 1836, lynxes were in country spite lynx its distribution but from 1916 to 1925 only 117. area in Sweden, and thereby demon-
the first half of the nineteenth the strated that live side side and his During century it can by with man
occurred far south Smâland and domestic the still the of lynx as as Blekinge, animals, species is on verge due while its limit in the north ran through northern extermination, solely to persecution, though local-
isolated nomadic the of rather Jämtland and Angermanland. Only ly population Jämtland is at present at a animals visited southern Lapland and Västerbotten. satisfactory level.
This distribution is interesting, for it shows that until
the hunting of the lynx with modern, devastating On the basis of the bounty paid for shooting seals, the
the habitat of the three resident in Sweden—the seal weapons began, main species was species common southern and central Sweden. Nowadays the main (Phoca vitulina), the ringed seal (P. hispida) and the
is seal geographical distribution of the lynx in the north; grey (Halichoerus grypus)—are clearly declining
the that marked the northern limit of numbers. The of this be other area formerly in cause can hardly
the animal north of the southern than them their is now only slightly man who, by disturbing at breeding
limit of its continuous As how- and them in the and range. early as 1850, places persecuting archipelagoes
there was in its habitat formed the coast districts, reduces the The maritime ever, a gap by populations.
plains of Västergötland and Östergötland. landscape is changed as man takes possession of islets
About the year 1900 the lynx had retreated from and rocks from the seals. The fact that the ringed
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12. Distribution of the lynx (Lynx lynx) in Europe about 1800. After K. Curry-Lindahl. 1951.
seal was able to survive the long warm period in the The red deer (Cervus elaphus), which was still to
Baltic indicates and that, in spite of its Arctic origin, it is be found in Östergötland in the seventeenth for of climate not reasons that it is retreating from the eighteenth centuries, is now restricted by man to
of Baltic sea. Scania, where the population probably consists
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13. The distribution of range of the lynx (Lynx lynx) in Europe in 1950 shows quite another picture than 150 years
earlier. After K. Curry-Lindahl 1951.
An about 150 animals. attempt to introduce it into A few centuries ago wild reindeer were found in
has been which shows that all the districts Sweden Västergötland successful, mountain of and in large parts
the be introduced suitable forest of the Norrland Norrbotten down species can into forests, in right to
tracts. the coast. It was exterminated little by little by the
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Lapps, who shot the last specimens about 1880. The may seem today, were on the verge of extinction.
wild reindeer still exists This critical situation followed in Norway. was gradually by a
spectacular expansion which is without parallel in
The reduction and local extermination of many birds northern Europe. Under the protection of the law, been decimation have the results of purposeful or the elk re-occupied practically the whole of Sweden,
unconscious interference the sites. To and the is well the on breeding roe on way.
this of birds the lesser white-fronted The little category belong ringed plover (Charadrius dubius) is a
goose (Anser erythrops), the whooper-swan (Cygnus wader, which in the inland has occupied different
habitats Rubbish of cygnus), the golden eagle (Aquila chrysaëtos ), the created by man. tips, the banks
artificial and colo- goshawk (Accipiter gentilis), the white-tailed sea ponds, bathing beaches have been
eagle, the peregrine falcon (Falco peregrinus), the nized by this species. The herring gull is clearly
gyr-falcon (F. rusticolus), the corncrake (Crex crex), favoured by rubbish tips, from which it is able to
the owl and the demand for food. eagle raven (Corvus corax). It is satisfy its It is only during recent
that the of these of that this of food has been utilized. typical majority species are birds years new source
and owls. The lesser white-fronted has It is that other factors have prey goose probable ecological con-
victim increase for years been the of ruthless slaughter in its tributed to the enormous of this species. The
hidden, isolated moulting localities during the period eider (Somateria mollissima) has increased greatly in
when it In for numbers recent which to be cannot fly. 1940, example, the skulls during years, seems
of 270 birds, all of which had obviously been killed directly due to the protection the bird has enjoyed.
at the same time, were found at Virihaure in Swedish
Such Lapland. massacres set their mark on the fre-
of the lesser quency white-fronted goose for a long
for from the time, geese great areas gather at moult- ing places.
Another group of birds has been driven from its dwelling places by drainage and similar actions.
Several of these species have found similar habitats in southern and central Sweden in exchange, for marshes are born, live, and die in rapid succession in a post-glacial, young country with land elevation, such Sweden. as Some species, however, have de- creased both and dis- visibly numerically as regards tribution. Among these are the snipe (Gallinago gallinago), and, above all, the ruff (Philomachus
both which pugnax), , of are now mainly to be found irj Norrland.
The catastrophic decrease in the frequency of salm- on (Salmo salar) and trout (S. trutta) in Swedish waters is due solely to man.
Unlike those mentioned above, there are other ani- mals that have been favoured clearly by man and his transformationof the landscape. Directly encouraged by the general cultivation of the land and by man's extermination of predators are the fox and the hooded
two that know crow, species how, in the absence of
make the best of all enemies, to practically types of habitat and edible that human everything beings 14. Retreat of the roe deer (Capre- leave from 1750 to 1830 was fol- behind them. The advance of the roe and the olus)
lowed elk by a miraculous spread is due, naturally, to a number of contributory northwards Sweden. The over factors, of which man is not the least important. At expansion was particularly rapid the end of the eighteenth century and the beginning between 1940-1955. After K. of the nineteenth, the two species, curious though it CURRY-LINDAHL 1958A.
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distance from them. It is difficult at present to determine how much the often at some but reduction increase in number and of The is cultivation, its ex- or range many starling helped by
animals due to other northwards has another is partly to man or wholly eco- pansion probably background,
logical factors. The Swedish populations of the long- although it has been facilitated by human activities. for- tailed duck (Clangula hyemalis), for example, have Olavi Kalela (1950) assumes that the starling limit which for of been declining rapidly. It is known that the species merly had a northern reasons
is often the victim of terrible oildischarge catastro- climate moved southwards during the seventeenth
known whether and and that its phes in the Baltic sea, but it is not eighteenth centuries, present expan-
affected the inhabitants of of its old habitat. they have particularly sion is a re-occupation but Swedish mountain waters. The bean goose (Anser Many animals are hampered greatly by man,
fabalis) has declined greatly in numbers both on its it is doubtful whether the species in question are
Swedish breeding grounds and in its European winter- decreasing in numbers. The hedgehog (Erinaceus
where it is but in this for is than other ani- ing places, hunted, case, too, europaeus), example, more any
it has been to show causal mal the victim of the motorcar. During impossible any connection. motoring
or in southern and central Sweden, numbers If the reduction disappearance of the greylag goose trips large been (A. anser) from its former nesting sites is due to hunt- of dead hedgehogs, which have run over during roads. This factor ing and interference is impossible to determine with the night, can be seen on the new The decimation the influence the of the certainty, but it seems not unlikely. must, in long run, frequency
of the black-grouse and caper-caillie in comparison to species. and difficult for bats that of former centuries, depends probably in the It is getting more more to
find suitable accommodation and first place on ecological factors caused by the trans- in trees buildings.
formation of forests and heaths into cultivated land, The golden plover (Charadrius apricarius) is being
but in this other factors unconnected with driven from marshes in southern but case, too, many Sweden,
im- remains intact. man may be present. Human beings played an the mountain population
rôle the the The reader find Table I portant in extermination of great bustard, probably expected to in but that other factors also contributed. other birds that Such birds the it is possible vary in frequency. as
The decline of the stock dove (Columba oenas) is bittern, the black-tailed godwit (Limosa limosa), the
its the razorbill usually attributed to man, who has destroyed nest- avocet (Recurvirostra avocetta), (Alca
but the versatile choice the and the ing trees, species is so in its torda). wren (Troglodytes troglodytes)
of that this be the redstart nesting places can hardly only (Phoenicurus phoenicurus) (increasing in
the stock dove does show in such cause. In not Norrland) a any case, by any changes frequency during short the first half of the means seem to be in uninterrupted retreat. period as present century,
The the numbers of the and of climatic factors such increase in mute swan probably on account as severe
the marsh harrier during the past quarter of a century or mild winters (the bittern, the razorbill, the wren),
coincides with their of but the favourable climatic the periods protection, at or circumstances during spring
number of birds and waders with colonization same time a aquatic migration consequent (the bittern,
that are show similar the black-tailed the the not wholly protected a expan- godwit, avocet, redstart).
sion. Man was the vertebrate These therefore, formerly only enemy variations, have nothing to do with
the mute and its has the transformation of the Short-term fluc- of swan, protection, therefore, landscape.
probably contributed greatly to the augmentation of tuations do not give sufficient evidence for a long-
its geographical distribution and its increased fre- term forecast.
quency.
The which is not It is in this to discuss all the osprey, dependent on eutrophic impossible paper spe- and also Table lakes for its fishing, which was protect- cies which, in I, are assumed to increase on ed much later than the and the marsh of climatic factors. mute swan account ecological, mainly
harrier, is another species that has increased greatly, Sweden is a sparsely populated country in com- particularly during the past fifteen to twenty years. parison with other European countries, and this is,
Its be contribution to this tenden- and has been, factor in 'the preservation may a always a positive possi-
but it be the bilities of animals to survive in of trans- cy, can hardly principal cause. Nearly spite a steady
birds the is able formationof the alone of all the of prey, osprey to countryside. Hardest pressed are the
the number of lakes and coastal waters feared and of draw on great large carnivores, which are by man, for its food, and its nesting places are not necessarily which man's opinions are prejudiced. in the vicinity of its fishing domains, but are quite Several species of animals possess a remarkable
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ability to adapt themselves to continually changing species of vertebrates in Sweden will increase rather conditions. this added that the number than of the fact that To must be decrease, in spite man is busily of be the in the habitats of the animals. species seems to on increase by immigra- engaged reducing Just
Sweden's the of the the is in that as bio- tion. situation on periphery geo- at present tendency direction, a animals that of the graphical distribution of many implies logical consequence improved climate. But the
in direction often result in climatic in the direc- changes a northerly new pendulum may swing opposite additions the Swedish fauna. It therefore be tion, and then the fauna will abandon the northern to may assumed, rather paradoxically, that the number of countries first.
REFERENCES
Kalela, O., 1950: Zur säkularen Rhytmik der Arealver- Cuhhy-Lindaiil, K., 1951: Lons (Lynx lynx) historia och
nuvarande förekomst änderungen europäischer Vögel und Säugetiere, mit be- i Sverige och övriga Europa.
42: sonderer Berücksichtigung der Überwinterungsverhält- Sverigcx Natur, yearbook. pp. 122-162.
nisse als Kausalfaktor. Ornis Fennica, XXVII: pp. 1-30. Curry-Lindahl, K., 1955: Djuren och människan i svensk
Field the North natur. Stockholm. Lloyd, L., 1831: Sports of of Europe 463 pp. residence comprised in u personal narrative of a in K., 1957: Nâgra djurarters utbredning.
and in the 1827-1828. 2nd ed. Atlas över 45-46: 1-8. Sweden Norway years Sverige. pp.
760 1958: pp. London. Curry-Lindahl, K., A. Djurgeografi, populations- Lönnbehg, E., 1930: Lodjurets förekomst i Sverige de dynamik och nutida faunaförändringar. Ymer. 78:
5-57. sista hundra âren. Kungl. Svenska Vetenskapsakade- pp.
miens Skrifter i Naturskyddsärenden. 14: pp. 1-36. Curry-Lindahl, K., 1958: B. Svanarna pâ Stockholms
ström. Sveriges Natur, yearbook. 49: pp. 111-119.
1960: - The maps are reproduced from the following literature: Curry-Lindahl, K., A. Djuren i färg däggdjur,
1950: Bubo bubo 199 Curry-Lindahl, K., Berguvens, (L.), kräldjur, groddjur. 2nd ed. pp. Stockholm.
förekomst i Sverige jämte nâgot om dess biologi. Vir Curry-Lindahl, K., 1960: B. Vâra Fâglar i Norden. Part
9: 113-165. 559-1022. Fàgelmrld. pp. II, 2nd ed., pp. Stockholm.
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