The lands lying within the Polar Circle
are inhabited by an assemblage of animals and plants,
many of which are peculiar to those regions. They
are mostly adapted to the abnormal conditions of life
prevailing in the high latitudes of our globe the
long, dark winters, and the short summers of one long
day. Though the numbers of species and of individuals
are few, there is a keen struggle for existence in
those regions. The prevailing colour of the ground
is white, and since a resemblance in the colour of
an animal to the ground it lives on acts as a protection
to weak ones, and also enables Carnivores to approach
their prey with greater facility, it is not surprising
that we should find the majority of polar animals
coloured white. As I remarked, the polar area
contains a very distinct set of species; most of them,
however, range beyond the confines of the Arctic Circle.
It is therefore scarcely justifiable to raise this
Arctic area into a distinct zoological region equivalent
to the great zoogeographic regions, which have been
established by Sclater and Wallace, though we might,
with Dr. Brauer, look upon it as a sub-region.
There are six typical Polar Land-mammals,
one of which, the Polar Bear, is semi-aquatic.
The Reindeer (Rangifer tarandus) occurs upon
almost all the polar lands, and it has often been
a source of speculation in what manner it has reached
such remote islands as Spitsbergen and Novaya Zemlya the
former of the two being so remote from a continent.
There is no doubt that Reindeer are great wanderers,
owing to the difficulty of finding sufficient food-supply
for the large herds in which they are accustomed to
travel; and for this reason they can cross, and have
been known to cross, distances of from ten to twenty
miles on ice. The Behring Straits, when frozen
over in winter, is frequently traversed by them.
But I quite agree with Dr. Brauer that it
is impossible to account for their presence in Spitsbergen
by an immigration from either Novaya Zemlya, Greenland,
or Scandinavia, under the present geographical conditions.
The seas between the former island and the other land-masses
referred to are rarely entirely frozen over. Even
if this should occur, the distances between Spitsbergen
and Greenland, Novaya Zemlya, or Scandinavia are so
great, that a migration across ice is quite excluded
from the range of possibilities, since Reindeer could
not subsist without food during the time it would take
to travel from one to the other. The manner in
which it did reach Spitsbergen and Greenland will
be discussed more fully below, and I will therefore
proceed to mention the other Arctic mammals.
One of the most important and most
typical species is the Polar Bear (Ursus maritimus),
the greater part of whose life is spent on the ice
and in the sea. The fact that its favourite nourishment
consists of seals proves its excellent and keen faculties
of sight and hearing, and its facility in swimming.
But it is not a dainty feeder, and lives upon almost
all animals which come within its reach; birds, land-mammals,
or fish are not despised in times of scarcity.
Its fur throughout the year is coloured white, though
in old bears it assumes a more yellowish hue.
Another large mammal, perhaps less well known, is the Musk-Ox, which resembles in size the smaller varieties
of Oxen, but in structure and habits is closely allied
to the Sheep. As is implied by the specific name,
it exhales a musky odour; this does not, however,
appear to be due to the secretion of a special gland,
as is the case in other animals with a similar smell.
The skin is covered with long brown thickly-matted
hair, interspersed with white. It is confined
to the most northerly parts of North America and the
American Arctic islands, and to North Greenland.
Though not now living in the Old World, it seems formerly
to have been abundant in Siberia, and, as we shall
learn later on, it was one of the species which took
part in the great Siberian invasion of Europe.
Its remains have been found not only in Germany and
France, but also in the south of England.
The Polar Fox (Canis lagopus)
occurs throughout the Polar Regions, and on islands
where even the Reindeer and the Musk-Ox are unknown.
Beyond the Polar Circle, its range extends into Northern
Asia, to the extreme north of North America, and the
mountains of Scandinavia. Like its congeners,
it had in pleistocene times a more southerly extension,
and fossil remains have been met with in various parts
of continental Europe and in England.
The Stoat (Mustela erminea),
which is known and much valued in commerce under the
name of Ermine, was formerly believed to occur only
in Arctic America and the northern parts of the Old
World, but in more recent years it has been discovered
in a number of the northern islands, such as Saghalien,
in the islands of the Behring Straits, the Aleutian
islands, and also in Greenland and Spitsbergen.
In Europe, it is found as far south as the Arctic
Hare, or perhaps even farther, and it flourishes in
the Alps up to a height of 9000 feet. It offers
a parallel to the Arctic Hare in the fact that in
some countries, such as Ireland, it only rarely turns
white in winter. The Irish form of the Stoat
differs so much from the English, that Messrs. Thomas
and Barrett-Hamilton are of opinion that it is specifically
distinct, as I mentioned in speaking of the divisions
of the British fauna.
The Arctic Hare (Lepus variabilis)
is almost the only one of the typical Arctic mammals
which still inhabits the British Islands, and for
that reason it is to most of us more familiar than
any of the preceding species. Hares have been
described from Greenland by the name of Lepus glacialis,
from the European Alps as Lepus alpinus, and
under other names from Arctic North America; but though
slight differences in the fur and even in the skull
can be pointed out, there is no doubt that all these
are only varieties or races of what, in the British
Islands, is known as the Irish or the Scotch Mountain
Hare, Lepus variabilis. In the Arctic
Regions this Hare remains white throughout the year,
but in Scandinavia and some other parts its fur becomes
brown in the summer, and in Ireland it frequently
remains entirely brown during the whole year, and
never, or only in very rare cases, becomes entirely
white in winter. Besides Scandinavia, Scotland,
and Ireland, it is found in Northern Russia, and also
in the Pyrenees, the Alps, and the Caucasus.
In Asia it occurs not only on the mainland of Siberia,
but it has been obtained on the Akita Mountains in
Japan and on the Mioko San Mountain, and also on the
island of Saghalien. It had in former times a
more extensive range, and its remains have been discovered
in England and in a number of places on the continent
of Europe. The peculiarity of its range, which
will be explained more fully directly, lies in the
fact of the occurrence of isolated colonies in the
mountains of Europe, in Ireland and Scotland, and
in the mountains of Japan. From a distributional
point of view, it is one of the most interesting species
of mammals, and its history throws a flood of light
on the geographical changes which have occurred in
former times.
One more species must be mentioned,
and that is the Banded Lemming (Cuniculus torquatus),
which occurs chiefly in Arctic America, Northern Siberia,
and Greenland. Though frequently mistaken for
the Scandinavian Lemming, there is a striking difference
in the character of the teeth, which has induced zoologists
to put them into distinct genera. The Arctic
Lemming, moreover, is distinguished from the Scandinavian
by the absence of external ears, the densely furred
feet, and by the great length of the two middle claws
in the fore-feet. There are two species of the
true Lemming, namely, the one just referred to, Myodus
lemmus, and Myodus obensis. These may
be looked upon as more or less Arctic species, since
they occur within the Polar Circle, but they are not
so exclusively confined to that region as the Banded
Lemming (Cuniculus torquatus). The remains
of both Cuniculus torquatus and of Myodus
lemmus have been found in British pleistocene
deposits.
Until recently no Lemming remains
had been found to the south of France, but Mr. Barrett-Hamilton
announced to us a short time since that Dr. Gadow
had discovered some skeletons with their skins still
preserved in a cave in Northern Portugal. These
were found to belong to the Scandinavian Lemming (M.
lemmus), and the author incidentally expressed
the opinion that there was some possibility of this
species still inhabiting the mountains of Spain.
The Lemming multiplies with great
rapidity under favourable conditions. In speaking
of his experiences in Siberia Dr. Brehm says:
“All the young of the first litter of the various
Lemming females thrive, and six weeks later at the
most these also multiply. Meanwhile the parents
have brought forth a second and a third litter, and
these in their turn bring forth young. Within
three months the heights and low grounds of the tundra
teem with lemmings, just as our fields do with mice
under similar circumstances. Whichever way we
turn we see the busy little creatures, dozens at a
single glance, thousands in the course of an hour.
But the countless and still increasing numbers prove
their own destruction. Soon the lean tundra ceases
to afford employment enough for their greedy teeth.
Famine threatens, perhaps actually sets in. The
anxious animals crowd together and begin their march,
hundreds join with hundreds, thousands with other
thousands, the troops become swarms, the swarms armies.
They travel in a definite direction, at first following
old tracks, but soon striking out new ones; in unending
files defying all computation they
hasten onwards; over the cliffs they plunge into the
water. Thousands fall victims to want and hunger;
the army behind streams on over their corpses; hundreds
of thousands are drowned in the water or are shattered
at the foot of the cliffs; the remainder speed on;
other hundreds and thousands fall victims to the voracity
of Arctic and red foxes, wolves and gluttons, rough-legged
buzzards and ravens, owls and skuas which have followed
them; the survivors pay no heed. Where these
go, how they end, none can say; but certain it is,
that the tundra behind them is as if dead, that a
number of years pass ere the few who have remained
behind and have managed to survive slowly multiply
and visibly re-people their native fields.”
This eloquent passage reminds us of the manner in
which migrations of all kinds of animals have taken
place in former times, and are still taking place.
It is principally want of food which compels them
to search for new homes.
One
of these, the Snow Bunting (Plectrophenax nivalis),
is a typically Arctic species. In summer it is
widely distributed, and is found in Spitsbergen, Novaya
Zemlya, Siberia, and the Arctic Regions generally.
In winter it migrates down into North America, into
Japan, Northern China, Turkestan, Southern Russia,
and occasionally even across Europe into North Africa.
Very characteristic Arctic birds are the Eider Ducks
belonging to the genus Somateria. Three
species have visited the British Islands. The
common Eider Duck (S. mollissima), which is
of such high commercial value, is abundant in Norway
and northward, throughout the Polar Regions.
The appearance of the King Eider (S. spectabilis)
on our coasts is an extremely rare occurrence, and
even in Norway it is only known as a visitor, but
on Novaya Zemlya and along the Arctic shores of Siberia,
in Greenland and Arctic North America, it is known
to breed. The third species, Steller’s Eider
(S. Stelleri), seems to be still rarer, and only in the Aleutian islands
and in the north of Alaska can it be said to be at all abundant. It is probable
that the famous Great Auk also was a typical Arctic species.
Its range extended to both sides of the Atlantic.
In Newfoundland and on the coast of Iceland it is
known to have been met with in considerable numbers
within historic times; and no doubt, like all Arctic
species, it extended farther southwards at a more remote
period.
The members of the genus Lagopus,
including the various species of Grouse, are likewise
of northern origin. The British Red Grouse (L.
scoticus), which may be looked upon as a form of
the Scandinavian Willow Grouse (L. albus), constitutes in some respects a curious case
of parallelism with the Arctic Hare, since the latter,
in its more southern station, generally retains the
summer fur throughout the year. The allied Ptarmigan
(L. mutus) inhabits Scandinavia, the Ural Mountains,
and some of the Asiatic mountain ranges. It is
also found in the European Alps and in the Pyrenees.
The North European range of the Ptarmigan suggests
that we are dealing with an ancient species which
came south from the Arctic Regions at about the same
time as the Arctic Hare; but it is more probable,
that this species has entered Europe more recently
with the Siberian migrants from Central Asia, where
indeed the genus had its original home. The Black
Cock (Tetrao tetrix) and the Capercaillie (Tetrao
urogallus) have also come to us from the east,
and have even penetrated into Ireland. They are
therefore some of the few instances of members of
the Siberian invasion having become temporarily established
there.
Reptiles and amphibia are altogether
unknown in the Polar Regions, but a large number of
fish, chiefly marine, have taken their origin there.
The Salmon family are of Arctic origin, as also are
the Sticklebacks and the Perches, many of the Cod
family, the Herrings, and several of the Flat fish.
It would lead me too far to refer
to the invertebrate fauna of the Polar Regions, but
a few remarks on the Arctic plants may not be out of
place.
The principal Arctic genera are Salix,
Ranunculus, Draba, Pedicularis, Potentilla, Saxifraga,
Carex, Juncus, Luzula, Eriophorum, and others.
Among the most characteristic Arctic
plants may be mentioned Dryas octopetala, to
which I have already referred as occurring in the west
of Ireland; Saxifraga oppositifolia, another
British species, occurs in the higher mountains of
Scotland, Ireland, and Wales; Braya alpina,
Papaver nudicaule, Lychnis apetala, Diapensia
lapponica, and Lobelia Dortmanna, which
is found in the lakes of Scotland and Ireland.
The dwarf birch (Betula nana) also, which still occurs in Scotland and
the North of England, and which had formerly a wider range in the British
Islands, should be included among these; but there are other plants probably of
Arctic origin, though not now occurring in the Arctic Regions, and to these may
be classed the so-called American species of plants which are found on the
northern and western coasts of Ireland, in the Hebrides, in Scotland, and in
North America. These are no doubt the relics of an Arctic flora which flourished
in high latitudes in past times when the climate there was more temperate.
As none of them occur in Siberia,
they must either have found their way to North America
and to Europe from the Arctic Regions, or have travelled
from North America across the latter to Europe.
In any case a former land-connection between the two
continents must have existed. This becomes the
more evident when we examine the remarkable results
obtained by the late Professor Heer, who first described
the Tertiary plant-beds in North Greenland. No
less than 282 species of plants have been described
by this eminent botanist from these deposits.
A large number of the plants found were trees belonging
to the genus Sequoia, Thujopsis, and
Salisburia, besides beeches, oaks, planes, poplars,
limes, and magnolias. That they grew on the spot
is proved by the fruits, which have been obtained
from these beds in various stages of growth.
From a similar deposit in Spitsbergen
a large number of fossil plants have also been brought
to light, many of which are identical with those found
in Greenland; and some of the Greenland forms (such
as Taxodium distichum and Sequoia Langsdorfii)
have been found too in Alaska, showing that there
was probably a continuity of land between Spitsbergen
and North America by way of Greenland. Two species
of Sequoias, namely, S. sempervirens
and S. gigantea, the well-known Californian
giant trees, are very closely allied to the Greenland
forms discovered by Professor Heer.
Heer assigned the Arctic plant-bearing
beds to the Miocene epoch, but doubts have been recently
thrown upon this opinion by Mr. Starkie Gardner, who
brought forward arguments in support of his theory
of their being of the Eocene age. Professor Heer,
however, was able to meet these criticisms, and he
is ably supported in his views by Professor Engler
and other eminent continental botanists.
It is evident that under the present
conditions of temperature none of those plants could
have flourished in Greenland. The climate must
have been much milder than it is at present.
Professor Heer estimated from the general aspect of
the fossil flora that the mean annual temperature
of North Greenland was at least nine degrees centigrade,
and that the mean winter temperature was not below
zero.
It will hardly be necessary for me
to review here the various theories which have been
advanced by geologists and botanists to account for
this remarkably high temperature in such northern
latitudes. Any one who has read the writings
of the late Dr. Croll cannot help being struck by the
facts he adduces to show the importance of ocean currents
in relation to the distribution of heat over the globe,
and it seems to me that the view which attributes
the mild climate prevailing in former times in Greenland
to warm ocean currents reaching the Polar Circle is
the one least open to serious objections. If
we suppose that the North Atlantic Ocean was bridged
by a land-connection between Scandinavia and Greenland
by way of Spitsbergen, and between Greenland and North
America, the Polar Ocean would be practically a closed
sea. If, then, a wide passage existed somewhere
about Behring Straits to allow a warm current to enter
and circulate within the Arctic Seas, we should have
the southern shores of Greenland washed by the warm
Atlantic current and the northern shores by a warm
Pacific current, which combination would undoubtedly
produce the effect of raising the temperature throughout
the Polar Regions very considerably; and especially
would that be the case with regard to Greenland and
the neighbouring islands.
It might be urged that the constant
darkness during winter must have had an injurious
action upon the flora, but it is found that in countries
such as Northern Russia, where southern plants are
housed during winter in greenhouses, the light being
almost entirely excluded by a covering of straw, no
serious damage is done thereby to the plants.
It seems probable that a similar gradual
refrigeration of climate in northern latitudes has
taken place after Miocene times as has been proved
to have occurred in Europe.
Some years ago Dr. Haacke propounded
the hypothesis that the centre of creation of all
the larger groups of animals was situated in the region
of the North Pole, and that the newly originated groups
must always push the older ones farther and farther
south into the most remote corners of the earth.
As instances of the correctness of his view he quotes
the fact that the more ancient mammals, such as Monotremes,
Marsupials, Lemurs, Edentates, and Insectivores, all
inhabit the more southerly parts of the world.
The Apteryx, Moa, Rhea, and the Ostrich, as well as
AEpyornis, which is only recently extinct, are found
in the same regions. But we have no palaeontological
evidence in favour of these extravagant views.
Fossil Edentates and Marsupials are almost entirely
confined to the Southern Hemisphere, and the supposition
that because these primitive mammals inhabit the extreme
south of our great continental land-masses, they therefore
came from the north, cannot be said to be an argument.
Nevertheless, I am quite with Dr. Haacke in considering
that the North Pole, or, we might say, the lands within
the Arctic Circle, have been the place of origin of
some of our European mammals, and there can be no
doubt that certain species in other groups, among
invertebrates and also plants, have originated in the
Polar Regions. The facts of geographical distribution
teach us that in these regions there has been a centre
of origin within comparatively recent geological times.
I have on a previous occasion drawn attention to the
range of the Reindeer: that it lives almost throughout
the Polar lands, and that it spreads into North America,
Northern Europe, and Northern Asia. We have,
again, fossil proof that its range extended down to
the Pyrenees in Europe in pleistocene times.
But there is not a scrap of evidence that it ever
during any time occurred farther south, either in Europe,
Asia, or North America. Its original home must
therefore have been in the Polar Regions, for if it
had originated either in Central Europe, Asia, or
America, there is no reason why it should not, in the
natural course of events, have extended its range
to the south as well as to the north.
The Arctic Hare presents us with a
very similar case of distribution. Like the Reindeer,
it inhabits, as we have learned, the Polar Regions
and the northerly parts of the Old World and the New;
but while we have only fossil evidence of the former,
more southerly, extension of the range of the Reindeer,
the Arctic Hare furnishes us with a still stronger
proof of its past southward range in the survival of
small isolated colonies in some of the southern mountain
ranges of Europe and Asia. It is generally believed
that the occurrence of the Arctic Hare in these southern
mountains is a standing testimony to the severity of
the climate at the time when it commenced its southerly
increase of range, but I have already shown that the
climate of Europe at that time was not necessarily
colder than it is at present, but that it may have
been somewhat milder. I think that a vast increase of ice in the Polar Regions
has taken place only at a comparatively recent date, and that both the Reindeer
and the Arctic Hare originated there during a much more temperate climate than
obtains at present. A great sensation was produced among European zoologists and
anthropologists when the discovery was first announced that the remains of the
Reindeer had been found in the Pyrenees, and it naturally gave rise to many
speculations as to the nature of the climate at the time when its range extended
so far south. The greater number of our best authorities
are still of opinion that the existence of the Reindeer
in Southern Europe points to the prevalence of an
arctic climate in that region. It is generally
overlooked, however, that the Reindeer-remains occur
in company with many typically southern animals, which,
if they had been found alone, would have been held
to be a certain indication of a warm climate.
The French geologist Professor Lartet, indeed, was
of opinion that the temperature during the time when
the Reindeer lived in the Pyrenees must have been
rather milder than it is at present. Similarly, Mr. Harle argues,
that the extremely cold climate probably did not extend
to South-western France, since that area only received
occasional visits from some of the representatives
of the Arctic fauna.
Long ago North American zoologists
recognised the existence in their country of two well-marked
races of the Reindeer (Caribou) a smaller
one with rounded antlers, and a larger one
in which the antlers are more or less flattened out. Two somewhat similar races can also
be traced in the fossil remains of the Reindeer in
Europe. It was, I think, Gervais who first pointed
out that the Reindeer remains from the north of France
differed from those found in the south; and Lartet
referred to the fact that the southern remains were
more like what, in America, is called the Barren-ground
Caribou, while those from Central European deposits
all belonged to the Siberian variety, which is more
like the Woodland Caribou of North America. In
Ireland, Professor Leith Adams also drew attention
to the curious fact that all the Irish Reindeer remains
resemble the Norwegian variety rather than the Siberian;
and Mr. Murray was so much struck by the close resemblance
between the Spitsbergen and Greenland forms with the
Barren-ground Caribou, that he based some speculations
on a former land-connection between these countries
on this circumstance.
We have, therefore, records of the
present or the former existence of a Reindeer resembling
the North American Barren-ground form in Greenland,
Spitsbergen, Scandinavia, Ireland, and the South of
France. In England the remains of the two forms
occur mixed, but I do not know in how far either the
one or the other predominates. The Barren-ground
Reindeer is in Europe altogether confined to the west;
the most easterly locality that I am acquainted with
being Rixdorf, near Berlin. The majority of the
European remains of the Reindeer seem to belong to
the Siberian or Woodland variety, and it would appear
as if some intercrossing between the two forms had
occurred in Lapland, since it is stated that in that
country the Reindeer is somewhat intermediate between
the two. All the Asiatic remains also resemble
the Woodland variety.
As far as I know, no explanation has
been attempted to account for this peculiar range
in Europe of the two forms of Reindeer. But if
we look more closely into the mode of occurrence of
the Reindeer remains, we find that the Barren-ground
form, seems to have existed in Western Europe long
before the other variety made its appearance there.
It was pointed out by Struckmann that the Reindeer
in Southern Europe occurs in older deposits than in
the north. In speaking of the northern ones, he
had of course chiefly the German deposits in view.
It is in one of the oldest pleistocene deposits
in Germany that the isolated instance, referred to
above, of the occurrence of the Barren-ground Reindeer,
near Berlin, has been noted.
There is still a further point which
illustrates the supposition that the Barren-ground
Reindeer was a more ancient inhabitant of Europe than
the Woodland one. The latter in all Central European
stations (in fact almost wherever it occurs fossil)
is associated with the remains of the typical inhabitants
of Siberia, such as the Glutton, Sousliks, Lemmings,
and others; but in the deposits in which the Barren-ground
Reindeer have been found in South-western France,
no other Arctic mammal finds a place. Again,
in Irish deposits none of the Siberian migrants are
found. The only explanation of this remarkable
fact is that the two varieties of the Reindeer have
come to Europe by different routes. We have learned
already from the observations of Mr. Murray that there
are evidences of the existence of a former land-connection
between North America, Greenland, and Spitsbergen.
Professor Petersen tells us that, according to recent
surveys, a high submarine plateau with a sharp fall
of 1000 fathoms towards the Atlantic Ocean begins
from Northern Norway and is continued as far as Spitsbergen.
Several islands, such as Bear Island, King Charles
Land, and others, arise from this plateau, and these
must be looked upon as the remains of a sunken land.
From Arctic America, thinks Professor
Schulz, we probably have had an uninterrupted
migration during the greater part of later Tertiary
times up to the commencement of the Pliocene epoch partly
over a direct land-connection between Greenland, Iceland,
and the Faroes, and also between Arctic America, Spitsbergen,
Franz Josef Land, etc. There was also a
connection between Asia and Alaska.
The distribution of the Barren-ground
Reindeer in Europe seems to warrant the belief that,
at the time it began its southward wanderings from
the Polar area, Northern Norway must have been connected
with Greenland in the manner just indicated, but,
as I shall explain later on, Russian Lapland and part
of Northern Russia, or the land between the White
Sea and the Baltic, must at that time have been submerged
by the sea. The greater part of Denmark and the
lowlands of Sweden were likewise submerged, but Scandinavia
extended south as far as Scotland, while Scotland
was connected with Ireland, and the latter with England
and France. The Reindeer migrating south into
Scandinavia could only reach the continent of Europe
by way of the British Islands. It appeared there
in the west and gradually extended its range east,
where, as I mentioned above, it has occurred in a
few isolated localities.
The advent of the Woodland form of
the Reindeer in Europe took place at a much later
stage. It came, as I indicated, with the hordes
of Siberian migrants which invaded Europe during what
is known as the Inter-glacial phase of the Glacial
period. Scandinavia, not being then directly
connected with continental Europe, was not accessible
to it; neither was Ireland, which had by that time
become disconnected from Great Britain. None
of the Siberian migrants seem to have been able to
cross the River Garonne, and we therefore find neither
the Woodland Reindeer nor any of the typical Siberian
species represented in the Pyrenean deposits.
The Woodland Reindeer persisted in
continental Europe until comparatively recent times,
and it has since made its way into Scandinavia across
Northern Russia, and probably mingled with the older
stock of the Barren-ground form. In the same way,
it may have come about that in the English pleistocene
deposits the remains of the two races occur.
In a recent contribution to our knowledge
of the deer tribe, Mr. Lydekker
suggests that the former division of the Reindeer
races into the two forms of Woodland and Barren-ground
Caribou, no longer holds good. He now recognises
no less than six races, as follows:
1. Rangifer tarandus typicu. " " spitzbergensi. " " caribo. " " terrae-nova. " " groenlandicu. " " arcticus.
I hardly think these can be considered
of equal value; indeed, though there may be differences
between R. groenlandicus, typicus, arcticus,
and spitzbergensis, the antlers exhibit a certain
much closer relationship among one another than to
R. terrae-novae and caribou. But
the whole subject is by no means as well known as could
be wished, and a very careful comparative study of
recent and fossil remains of the Reindeer from various
parts of the Old and New Worlds is much needed to
put our views on a firmer basis.
The presence of the Arctic Hare in
Ireland and the absence of the common European Hare
(Lepus europaeus) can be explained in a somewhat
similar manner. The Arctic Hare is the older
of the two species corresponding with the
Barren-ground Reindeer and the European
Hare the newer one, associating, like the Woodland
Reindeer, in its westward migration with Siberian
animals, though probably of Oriental origin.
Its discontinuous
range and its isolated position in the Alps, Pyrenees,
and the Japanese mountains, all tend to show that
it is an ancient species. Moreover, its presence
in Ireland in the plain as well as in the mountains,
clearly points to the fact that, in the British Islands
at any rate, the Arctic Hare was the first comer,
and that subsequently the European Hare invaded these
countries. It probably found Ireland then no longer
accessible, having since become separated from England.
Again and again do we find the statement repeated,
that the presence of the Arctic Hare in Europe is
a clear proof of the former prevalence in our continent
of an Arctic climate. But if so, why should this
Hare at present live and thrive in Ireland, which
has a particularly mild climate in winter, and be
absent from so many continental stations where the
temperature more resembles that of its native home?
If we suppose that the European Hare migrated to Europe
from the east, after the Arctic Hare had become established
in Western Europe, and drove the latter into the mountains
or northward whenever the two came into contact, we
should have, it seems to me, a better explanation
of the range presented by the two species. I
was formerly of opinion that the European Hare had
come with the Siberian animals from Siberia, but it
appears to me more likely now, that it reached our
continent with the Oriental migrants, and only then
joined the Siberians in Eastern Europe.
The evidence in favour of a former
land-connection between Scandinavia and Greenland,
rests on many other facts besides those already brought
forward. That some form of land-connection formerly
existed between Europe and Greenland is now indeed
almost universally accepted. That it was situated
more to the south between Scotland and Greenland is
a supposition which has been actively supported by
many leading authorities, but it seems to me that
if such a land-bridge existed, it must have been in
very early Tertiary times, whilst the northern one,
such as I have indicated, may have originated later
and persisted until a recent geological date.
The distribution of few groups of
animals is now better known than that of the larger
butterflies and moths (Macro-lepidoptera); even
those of Siberia have been fairly well investigated.
The interesting facts obtainable from their distribution
are therefore of special value. No less than
243 species of Lepidoptera are mentioned by
Moeschler as being common to North America and Europe.
It is extremely probable that a fair number of these
have either migrated direct from America to Europe
or vice versa, though many may be of Asiatic
origin, and have wandered east and west from their
original home. The following twelve species are
mentioned by Petersen as occurring in Arctic
Europe and also in Arctic North America, but not in
Asia: Colias nastes, Colias hecla,
Syrichthus centaureae, Pachnobia carnea,
Plusia parilis, Anarta Richardsoni,
Anarta Schoenherri, Anarta lapponica,
Anarta Zetterstedti, Cidaria frigidaria,
Cidaria polata, Eupithecia hyperboreata;
and these, as he remarks, point to the possibility
of a former direct land-connection between Europe and
North America.
Mr. Petersen believes that the chief immigration into the
Arctic area of Europe is post-glacial and took place from Siberia, since the
majority of the species are still to be found in that country at the present
day. He also draws particular
attention to a fact, which I shall discuss
more fully in the next chapter, namely,
that the most characteristically Arctic forms of Northern
Europe, which also partly occur in the Alps, are entirely
absent from the Caucasus.
Adopting the glacial views of some
of our leading geologists, Petersen comes to the logical
conclusion that Central Europe could not have possessed
any butterflies during the height of the Glacial period,
but since all evidences seem to point to the chief
migration from Siberia having taken place after the
Glacial period, he concludes that they must have survived
the severe cold of that time in Central Asia.
He leaves us, however, to imagine under what possible
geographical conditions the climate in Europe could
be too severe for a lepidopterous fauna, while at
the same time Central Asia could maintain an abundant
one.
In a suggestive note on the origin
of European and North American Ants, Professor Emery
states that a great number of North American
ants are specifically identical with European ones;
whilst Dr. Hamilton tells us, as an instance,
that specimens of the beetle Loricera coerulescens
from Lake Superior and from Scotland do not seem to
vary to the extent of a hair on the antennae.
He enumerates 487 species of Coleoptera as
being common to North America, Northern Asia, and
Europe, many of which no doubt have migrated by the
Americo-European land-connection.
Arctic Scandinavia or Lapland, according
to Sir Joseph Hooker, contains three-fourths of the
entire number of species of plants known from the
whole circumpolar area. His view, that the Greenland
flora is almost exclusively Lapponian, having
only an extremely slight admixture of American or
Asiatic types, again points to a former
more intimate connection between North America and
Arctic Europe, and indeed he remarks, “It
is inconceivable to me that so many Scandinavian plants
should, under existing conditions of sea, land, and
temperature, have not only found their way to Greenland
by migration across the Atlantic, but should have
stopped short on its western coast and not crossed
to America.”
Hooker’s view, that the Scandinavian
flora is of great antiquity, that, at the advent of
the Glacial period, it was everywhere driven southwards,
and that during the succeeding warm epoch the surviving
species returned north, has been adopted by the great
majority of naturalists.
The natural corollary of this theory
is that there must have been, between the beginning
of the Glacial period and the present time, either
two independent land-connections between the Polar
Regions and Northern Europe at different epochs to
enable animals and plants to travel southwards and
once more to regain their former northern home, or,
that during the whole of the Glacial period the Polar
Regions were uninterruptedly connected with Northern
Europe, until the fauna and flora had once more reached
their northern goal, after the Polar lands had been
desolated by the supposed rigours of that period.
In following the history of the Arctic
migration to Europe, it is of great importance to
determine the nature and the time of duration of these
land-connections. The Greenland flora is a very
instructive one in helping us to understand many of
the problems connected with the origin of the European
plants and animals. To judge from the remarks
of Professor James Geikie and Mr. Clement Reid, no
flowering plants could have existed in the British
Islands during the height of the Glacial period, and
one would suppose that the cold in Greenland at that
time must have been far more intense than in England.
If no flowering plants could exist in the latter country,
then very surely none could in Greenland, where the
climate was of necessity by far more rigorous.
It will be a surprise, therefore, to those who are
acquainted only with Professor Geikie’s views
of the nature of the Glacial period, that two of the
most eminent Swedish botanists, who have made a special
study of the flora of Greenland, have come to the
conclusion that a survival of flowering plants has
taken place in Greenland itself from pre-glacial times.
According to Professor Nathorst, only a few
plants could have survived the Glacial period in Greenland.
The species now peculiar to that country may perhaps,
he thinks, be the remnants of those which existed
in pre-glacial times. Mr. Warming, on the other
hand, is of opinion that the main mass of Greenland’s
present flora survived the Glacial period there, and that the remainder was carried from Europe
and North America by occasional means of distribution
of the nature indicated by Darwin.
Very similar views on the origin of
the present Polar flora are expressed by Colonel Feilden,
who says, “To my mind it seems indisputable
that several plants now confined to the Polar area
must have originated there and have outlived the period
of greatest ice-development in that region”. No land-connection at all
need be supposed to have existed in recent geological
times, that is to say, during the Glacial period or
after, if Mr. Warming’s and Colonel Feilden’s
views be adopted. A pre-glacial connection would
be sufficient to explain the general features of distribution.
An admission is thus obtained from these two independent
authorities that the climate during the Glacial period
must have been vastly less severe in the Polar Regions
than is generally conceded. I am of opinion that
not only the whole of the present flora, but also
the fauna of Greenland survived the Glacial period
in that country.
If we suppose that an extensive centre
of origin existed in the Polar area, or we may say
in Greenland, both animals and plants would have been
able to spread from it into Northern Europe and North
America by means of the land-connections which are
generally supposed to have existed in pliocene
times, that is to say, just before the commencement
of the Glacial period. There must have been at
this time a connection too between Scotland and Scandinavia,
which will be dealt with more fully presently.
The important point is to consider what light the
Greenland flora and fauna will throw upon the problem
of the continuity of the aforesaid land-connection
during the Glacial period. We have seen that
the Barren-ground Reindeer, a typically Polar species,
penetrated as far south as the Pyrenees, the Arctic
Hare went as far, while a number of other species
of Polar animals and also of plants occur in the Alps.
Of these it remains to be seen how many have come direct
by way of Northern Europe or from the Polar Regions
by way of Asia. At any rate, as the origin of
the Alpine animals and plants will be discussed in
another chapter, there is no need to dwell on this
subject at present.
From the nature of the distribution
in Ireland of Arctic plants and animals, which occur
mostly on the north and west coasts, it would seem
that a stream of migration entered from Scotland, and
I have no doubt that that same migration came into
Scotland directly from Scandinavia by a route over
which now roll the waves of the North Sea. There
is, moreover, a very
interesting so-called American element in the north-western
European flora, that is to say, plants now found in
North-west Europe and North America without occurring
in Greenland or any of the islands which might have
formed the former highway between the Old World and
the New. These are probably some of the more
ancient Polar plants which have become extinct in the
Arctic Regions and survive in isolated patches in favourable
localities. We find seven species of these American
plants in Ireland, almost entirely confined to the
north and west coasts. These are Spiranthes
Romanzoviana, Sisyrinchium anceps, Naias
flexilis, Eriocaulon septangulare, Juncus
tenuis, and Polygonum sagittifolium.
To them must be added another plant recently discovered
by the Rev. Mr. Marshall in the south of Ireland,
namely Sisyrinchium californicum. As I
have mentioned in former writings, there are three
species of North American freshwater-sponges in Ireland
which have not hitherto been discovered elsewhere
in Europe or in Asia. These, namely Ephydatia
crateriformis, Heteromeyenia Ryderi, and
Tubella pennsylvanica, all occur in some of
the lakes near the western coast of Ireland.
There are in all groups of animals
instances of species which are confined to Europe
and North America, while unknown from the Asiatic
continent, but none, as far as is known, have such
a very discontinuous range as that of the animals
and plants just referred to. In some cases the
species still occur in Greenland, and in this way make
it still clearer that their migration in former times
took place from one continent to the other by way
of that country. As an interesting instance of
such distribution may be mentioned the Common Stickleback
(Gasterosteus aculeatus), which is found in
Greenland, North America, and Europe, but is quite
absent from Asia. Then again, the Nine-spined
Stickleback (Gasterosteus pungitius) is confined
to Western Europe and North America, though an allied
species, Gasterosteus sinensis, lives in China
and has probably penetrated there from the New World
across the old Behring Straits land-connection.
The Coleoptera Diachila arctica,
Elaphrus lapponicus, and Blethisa multipunctata
are good instances of species which have come to us
from North America by way of Greenland. I have
already referred to the Lepidoptera, but might add
that eleven species of Anarta occur in Scandinavia,
eight of which reappear again in Labrador, none of
them, however, being met with in Siberia. Then
again, take the interesting Crustacean Lepidurus
(Apus) glacialis. It is found in Greenland,
Spitsbergen, Lapland, and Norway; and formerly, as
we know from fossil evidence, it ranged into Scotland.
Another Phyllopod, viz., Branchinecta paludosa,
inhabits Greenland, Lapland, and Norway. Mr.
Kennard suggests that the freshwater Snail Planorbis
glaber might also belong to the same migration.
And there are no doubt large numbers of others.
Professor Emery mentions that Northern
Europe possesses one peculiar genus of Ant, viz.,
Anergates. This is closely allied to Epoccus,
another genus confined to North America. It seems
probable, therefore, that both of these have sprung
from an Arctic genus which sent two branches southward
into the two continents without there being any migration
through Asia.
The general range of the Arctic plants
and animals gives no reason to suppose that the Greenland
fauna and flora of the present day were exterminated
by the Glacial period and then reintroduced into that
country. Nor have we any evidence that such a
fauna and flora migrated across the British Islands
northward. The Greenland animals and plants too
are altogether much more like the Lapland ones than
those of Scotland. It will also become evident
to the reader of this work that no very extensive
migrations could have taken place during the post-glacial
period, and that almost everything points to a survival
of both fauna and flora in northern latitudes throughout
the Glacial period.
If we take into consideration the
palaeontological evidence of the two races of Reindeer
in Europe, one of which came to us from the north,
and that the Arctic Hare and one of the races of the
Stoat entered our continent from the same direction when
we, moreover, carefully review the numerous other
instances quoted of plants and animals which could
only have reached us from the north, the irresistible
conclusion is forced upon us that a land-connection
existed at no very distant period between Northern
Europe and the Arctic Regions of North America.
This is not a new hypothesis. Many geologists
are of opinion that a land-passage did exist within
comparatively recent times, uniting Europe, Greenland,
and North America. But the position of this old
land-bridge, as I have mentioned, has been generally
placed somewhat farther south than I should feel inclined
to put it.
The fact that very extensive glaciers formerly covered the
mountains of Scandinavia on the eastern side, whilst they scarcely reached the
sea on the west, seems to favour the view of a warm current having washed the
western shores. As I shall attempt to show later
on, the Arctic Ocean extended across Northern
Russia at that time from the White Sea to the Baltic that
is to say, to the eastern shores of Scandinavia, which
country was then joined to the north of Scotland.
The predisposing agents to a copious snowfall existed
in Scandinavia, viz., an excessive evaporation
of the warm Atlantic waters and unusual precipitation
in the form of snow owing to the cold given off by
the Arctic waters on the east side of the mountains.
It is therefore probable that the land-connection
which united Europe and North America was farther
north than has been supposed.
If we sail straight across from Northern
Scandinavia to Greenland, we traverse an exceedingly
deep marine basin; but if we examine the sub-marine
bank which runs all along the coast of the former country
from south to north, we find that it does not end when
the extreme north of the land is reached. The
bank extends much farther north, and is continued
as far as Spitsbergen. As I have said before,
the latter, as well as Bear Island, must be looked
upon as the remains of a large mass of sunken land the
ancient Scandinavia stretching far into the Arctic
Circle. Professor Nathorst speaks of Spitsbergen
as a northern continuation of Europe, not only geographically,
but also botanically and geologically. However,
this northern land must have stretched even farther not
perhaps farther north, but farther west. Here
lay the old land-connection between Scandinavia, Greenland,
and North America. One of the highest
authorities on the geographical distribution of plants,
Professor Engler, maintains that the arguments in favour
of this Arctic connection of America with Europe are
more weighty than those for a land-bridge between
Greenland, Iceland, the Faroes, and Great Britain.
Moreover, he is of opinion that a certain number of
species of plants belonging to the Alpine flora of
Arctic Siberia have travelled from Scandinavia via Greenland and North
America to Eastern Asia, and not direct from Scandinavia to Siberia.
That this ancient Arctic land-connection
existed almost throughout the Glacial period appears
to me probable. It has often been suggested that
such a land-barrier was one of the principal causes
of the production of the glacial phenomena in Europe,
and as such it must have existed intact certainly
during the earlier stages of the Glacial period.
The barrier must then have gradually subsided in one
or two places; and once a breach was formed, the complete
union between the Atlantic and the Arctic Oceans could
not have been long delayed.
The terrestrial fauna and flora, as
we have seen, lend strong support to the view of the
former connection between Scandinavia and Greenland,
but many other facts point in the same direction.
It was Edward Forbes who first drew attention to the
presence of a number of species of littoral molluscs
on the coast of Finmark which also occur on the coast
of Greenland, and he expressed the firm conviction
that they indicated by their existence on both sides
of the Atlantic some ancient continuity of the coast-line.
He held that the line of migration of these mollusca
was probably from west to east, and that it must have
taken place during physical conditions entirely different
from those prevailing at present. If Forbes’s
view is correct, a current must have existed from the
north coast of North America along the northern shore
of the ancient land which stretched east as far as
Europe. We have also some palaeontological evidence
bearing on the existence of such a current.
As we shall learn presently, the early
stages of the Glacial period were accompanied by a
marine transgression over Northern Russia and Germany an
overflow, as it were, of the waters of the Arctic Ocean
covering a great part of Northern Europe, with the
exception of Norway. One continuous ocean ultimately
extended from the east coast of England across Holland,
Northern Germany, and Russia to the White Sea. The south of England being at that
time joined to France, and Scotland to Scandinavia,
there was no direct communication between this large
North European Sea and the Atlantic. The glaciers
which took their origin in the Scandinavian Mountains
discharged icebergs into this sea, and many of them
no doubt were stranded on the east coast of England.
The boulders of Scandinavian origin which have been
discovered in recent geological deposits on that coast
have generally been traced to the action of land-ice,
but the supposition that they have been carried by
icebergs the older theory appears
to me the more probable one. Such boulders begin
to make their first appearance in the Red Crag, a deposit
which is now looked upon as belonging to the newer
pliocene series. But whether we call
it pliocene or pleistocene really matters
little. The important fact is, that glacial phenomena,
consisting of the appearance of boulders foreign to
the country together with an invasion of Arctic shells,
are now ushered in upon a coast which shortly before
teemed with the southern life of a Mediterranean character.
Among the new arrivals in these English crags there
are no less than eighteen species of North American
marine mollusca. Since the German Ocean had then
no direct communication with the Atlantic, these mollusca
could only have come from the White Sea, and Forbes’s
Arctic current would offer an explanation of
the manner in which they were enabled to migrate there
from their original home.
It might be urged that we have no
grounds for the supposition that the German Ocean
was practically a closed basin; and that these American
species probably inhabited at that time the whole of
the North Atlantic Ocean. But if such had been
the case, we ought to have evidence of the occurrence
of some of these species in the newer Tertiary deposits
along the west coasts of the British Islands.
Such beds exist; there is, however, not a trace in
any of them of any American mollusca. In examining
the marine deposits of St. Erth, on the coast of Cornwall,
which are believed to be of about the same age as the
newer crags, Messrs. Kendall and Bell were much struck
by the absence of the species characteristic of the
latter. The St. Erth fauna led them to believe
that the Arctic Ocean could not then have opened into
the Atlantic, but that a land-communication had existed
between Europe and North America, so as to form a
barrier of separation between the two oceans.
This again perfectly harmonises with the views I have
expressed, and supports them.
Let us now look a little more closely
at the history and the fauna of the Baltic and the
adjoining lakes, in order to gain additional information
as to the geographical changes which have had such
lasting influence on the peninsula of Scandinavia.
The Baltic is a shallow sea covering an area of 184,496
square miles, and its waters are decidedly brackish.
The fauna is a poor one, being too salt for the purely
freshwater species and not salt enough for the typical
marine forms. The absence of some animals which
we should expect to find there is one of the remarkable
features about the Baltic, but, on the other hand,
some species occur which are altogether strangers
to the fauna. And these, moreover, are confined
to the extreme northern end of the sea. I need
only refer to the Arctic Seal (Phoca annelata),
which is confined to the Gulf of Bothnia, and to the
four-horned sting-fish (Cottus quadricornis), neither of which occur on the west
coast of Scandinavia. But there are others which
point in an equally unmistakable manner to the former
existence of a marine connection between the Baltic
and the southward prolongation of the Arctic Ocean known
as the White Sea. It is generally admitted now
that such a union between these two seas, viz.,
the Baltic and the White Sea, occurred in recent geological
times, but opinions differ as to the duration of this
connection. I adhere to the view expressed by
Murchison and others, that the boulder-clay is a marine
deposit. I am also convinced that the Arctic
Ocean, as I have already mentioned, transgressed over
the lowlands of Northern Russia at about the time when
the newer crags were being deposited on the east coast
of England; that the same large sea also covered Northern
Germany, Denmark, Holland, and the lowlands of Sweden,
and laid down the lower continental boulder-clay which
is spread over such vast tracts of land in those countries.
I shall have occasion to refer to this again more
fully in the next chapter; meanwhile, it should be
remembered that this stage was followed by a partial
retreat of the northern sea, though Scandinavia did
not become joined to the Continent. The date
of this retreat of the sea, corresponds probably to what is known as the
inter-glacial phase of the Glacial period, and I think it must have been during
this time that the Forest-Bed on the coast of Norfolk was laid down.
None of the Siberian mammals apparently
entered Scandinavia at the time when they invaded
Central Europe and penetrated as far west as England
and Western France. Nor did the great Oriental
mammals, like the Mammoth and others, reach Scandinavia;
and Professor Pohlig argued, on the strength of these
facts, that the latter country was either for a very
short time only free from ice, or that it had defective
land-communication with the Continent during inter-glacial
times. This seems to me scarcely to explain the
facts of distribution and account satisfactorily for
the absentees. Nor does it, of course, harmonise
with the views that I have announced above. Professor
Engler’s remark, that Scandinavia probably projected above the glacial sea
as an island, is more in accordance with these views, though the term island is
scarcely applicable to that country, since it was always, as I said, indirectly
joined to the Continent. The fauna of Scandinavia,
both fossil and recent, points to a direct isolation
of that country from the continent of Europe during
a considerable period.
Another proof that Northern Russia
and the lowlands of Sweden were covered by the sea
comes to us from a study of the fauna of the relict
lakes the “Reliktenseen” of
Leuckart. This name was first applied by Leuckart
to lakes containing marine organisms, which are supposed
to have been flooded by, or to have been in close
communication with the sea at some former period,
like the lakes Ladoga and Onega in Russia. His
views have been worked out subsequently in greater
detail by Loven and O. Peschel, who gave them their
strong adherence. Many leading zoologists, such
as Professor Sars and others, have since adopted them,
and though discredited by Professor Credner, the theory
still offers the best explanation for the origin of
marine animals in freshwater lakes.
Professor Credner’s contention,
that marine mollusca are always absent from these
relict lakes, seems at first sight a stumbling-block
to the theory. But the explanation is really
simple enough. It is to Dr. Sollas that
we owe a very ingenious explanation of the origin of
freshwater faunas. He showed that all freshwater
organisms in their early stages of development are
provided either with some process enabling them to
attach themselves to a foreign object, or that they
pass this period within the body of the parent.
This is a provision of nature to prevent freshwater
organisms from being floated out to sea, where they
would perish, until they reach maturity and can cope
with floods and currents. Had Professor Credner
been aware of Dr. Sollas’s views, no doubt he
would have modified his criticisms, for, as most marine
mollusca have free-swimming larvae, they would have
little chance of becoming permanent residents of lakes.
During their larval stage, marine molluscs are quite
a prey to the currents of the sea. They have practically
no swimming organs, and only move by lashing to and
fro the tender cilia with which they are provided.
This disposes, therefore, of Professor
Credner’s main criticisms. As for the fauna
of the relict lakes, we are now only concerned with
those of Northern Russia, Finland, and Sweden.
In the lakes Wetter and Wener in the latter country
occurs the four-horned sting-fish (Cottus quadricornis), which, as we have learned, also inhabits
the northern part of the Baltic, and, as was suggested,
migrated there at a time when the latter was connected
with the White Sea. The principal food of this
little fish consists in a marine Crustacean called
Idotea entomon, an animal allied to our common
woodlouse. This is a typical marine species,
but it occurs also in the relict lakes of the countries
mentioned above, as well as in the Baltic and the Caspian.
Perhaps the best known form with a similar range is
the Schizopod crustacean ysis relicta, which is clearly a descendant of the Arctic marine
Mysis oculata, of which it was formerly considered
a mere variety. The two Amphipods ammaracanthus
relictus and Pontoporeia affinis and the
Copepod Limnocalanus macrurus, ar, ar, are three additional well-known Arctic
crustaceans whose range differs but little from those above-mentioned.
These facts all go to prove that the sea formerly covered the
lowlands of Sweden, Finland, and Northern Russia. The fauna of Scandinavia, as
we have seen, indicates that during the greater part of the Glacial period the
country was not directly connected with continental Europe as it is now. It
seems that the barrier of separation probably consisted of a broad expanse of
ocean on which floated numerous icebergs, which originated from the Scandinavian
glaciers as they reached the sea. This was a cold sea, whilst Western
Scandinavia was washed by the Gulf Stream. We might look upon
the boulder-clay which covers such vast tracts of
country in Northern Germany, Russia, and Holland as
deposits formed by this sea rather than the ground-moraine
of a huge Scandinavian glacier. I shall refer
to this subject again in the next chapter; meanwhile
it may be remembered that the boulder-clay of Northern
Europe exactly resembles in all important particulars
the similar accumulations met with in the British
Islands. They resemble one another also in the
occasional occurrence of sea-shells, the frequent
appearance of bedded deposits, and the often inexplicable
course taken by boulders from their source of origin.
There occurs often a singular mixture and an apparent
crossing of the paths of boulders in the boulder-clay.
Professor Bonney remarks that these are less
difficult to explain on the hypothesis of distribution
by floating ice than on that of transport by land-ice,
because, in the former case, though the drift of winds
and currents would be generally in one direction,
both might be varied at particular seasons. So
far as concerns the distribution and thickness of
the glacial deposits, he says there is not much to
choose between either hypothesis; but on that of land-ice
it is extremely difficult to explain the intercalation
of perfectly stratified sands and gravels and of boulder-clay,
as well as the not infrequent signs of bedding in
the latter. Two divisions are generally recognisable
in the continental boulder-clay a lower
and an upper. An inter-glacial phase characterised
by a less severe climate is assumed to have intervened
between the deposition of the two. In Russia
no such division can as a rule be made out, and sea-shells
are either entirely absent or extremely scarce.
It has been pointed out by Professor J. Geikie that
the erratics a name applied to boulders
in boulder-clay in the upper division have
travelled in a different direction from those contained
in the lower. Taking for granted that the boulder-clay
is a marine deposit, this phenomenon seems to indicate
that the current which prevailed during the early
part of the Glacial period in this North European
ocean was different from the prevailing current during
the latter part. I have attempted to explain this
circumstance by the supposition that during the early
part of the Glacial period the Northern Sea had a
connection with the Ponto-Caspian Sea a
sea formed by the junction of the Black Sea and the
Caspian. There is geological
evidence, as will be explained in the following chapter,
that the area of these two seas was considerably larger
in glacial times than it is now, and that they were
joined across the valley of the Manytch. After
the inter-glacial phase of the Glacial period, the
North European Ocean became connected with the Atlantic
Ocean across the north of England,
the junction between the former and the Ponto-Caspian
having meanwhile become dry land.
A fresh current, now flowing westward, was set up in
the North European Ocean, which accounts for the fact
just cited that the erratics in the upper continental
boulder-clay have travelled in a different direction
from those in the lower. The boulder-clay laid
down by the sea on the midland and northern counties
of England, just as was the case with the similar
deposit on the Continent, is generally accredited
to the action of land-ice. It is by most geologists
looked upon as the ground-moraine, partly of the huge
Scandinavian glacier which is supposed to have impinged
upon the English coast, partly of local British glaciers.
But renewed geological investigations
on this point throw doubts upon these theories.
Thus Mr. Harmer remarks in a recent contribution to
glacial literature, that “it is difficult
to see how the Baltic glacier could have reached East
Anglia, though ice-floes with Scandinavian boulders
might easily have done so, while had the Norwegian
ice filled the North Sea and overflowed the county
of Norfolk, some evidence of its presence ought to
be found in the glacial beds of Holland.”
All the phenomena of distribution
of the British fauna and flora are, as we have seen,
much more easily explained by the supposition of a
damp, temperate climate, such as might have been produced
by the proximity of a cold sea on one side and of
a warm one at the other, than by invoking an arctic
climate with enormous glaciers. Most of the living
animals and plants would have been exterminated under
the latter conditions. Palaeontological evidence
in Great Britain clearly indicates that southern species
migrated first to these islands, that Arctic species
were then driven south from their native lands, probably
owing insufficient food-supply and climatic changes
in the north, that finally eastern species
invaded the country all this without the
annual temperature of Europe being apparently much
affected. For we find in the British pleistocene
deposits and Mr. Lydekker draws particular
attention to this remarkable fact a curious
intermingling of southern and northern mammals, which
undoubtedly lived side by side. Everybody knows
that northern and Arctic species can live perfectly
well in a temperate climate, but that it is almost
impossible to acclimatise southern animals in an Arctic
or even temperate one. We have in this circumstance
almost a proof, therefore, that the climate cannot
have been very cold. Though a cold sea bathed
the shores of Eastern England, and even eventually
invaded a portion of Northern England, the warm ocean
on the west must have effectually prevented any great
lowering of temperature.
At the time when the North European
Sea flooded a portion of England, Scandinavia was
still connected with Scotland, and the latter with
Ireland. There is no doubt that
the food-supply in the Arctic Regions was decreasing
with an increase of snowfall and with the gradual
lowering of the land, which reduced also the habitable
area. Arctic species therefore were driven south
in search of fresh pastures. But it need not
be supposed that anything like a vast destruction of
the fauna of the Arctic Regions took place. Only
fewer mammals were able to find food in a given space
than heretofore. This southward migration may
have commenced, in the case of plants and the invertebrates,
at a much earlier time, during the Miocene
or Pliocene Epochs, but it is doubtful
whether the mammals and birds which we find in our
pleistocene and recent deposits began to travel
south much before the commencement of the Glacial
period. The beginning of the Glacial period in
England, I think, is indicated by the deposition of
the Red Crag, though the latter is generally regarded
as belonging to the pliocene series.
Much of the northward migration from the British Islands
of Lusitanian and other forms had then ceased, but
we have in Scandinavia, just as in these islands,
a southern relict fauna and flora, plants and animals
which had wandered across what is now the German Ocean
from Scotland to Scandinavia, and have never become
extinct in that country to the present day. I
need only mention the Red Deer, the Badger, and Slugs
of the genus Arion.
Professor Blytt directs attention
to some such southern relict species of plants now
only found in the extreme south-west of Scandinavia,
such as Asplenium marinum, Hymenophyllum
Wilsoni, Carex binervis, Scilla verna,
Erica cinerea, Conopodium denudatum,
Meum athamanticum, and Rosa involuta.
The Arctic fauna and flora in Scandinavia that
is to say, the descendants of those species which
migrated direct from Greenland and Spitsbergen, as
we have seen, are numerous. They of course persisted
throughout the Glacial period in the country, and are
now in many localities being exterminated partly by
change of climate, partly by a keen competition with
more vigorous rivals which have come to Scandinavia
from the east. It is a curious circumstance, as
pointed out by Professor Blytt, that the Arctic plants
in the Botanic Gardens at Christiania are able to stand almost any amount
of sunshine, but are very liable to be injured by the frost, and have to be
covered in the winter. A similar observation has been made in the case of the
Alpine plants at Kew Gardens, which have to be wintered in frames, though their
homes are either in the high Alps among the everlasting snows or in the
intensely cold climate of Greenland. Many of the Scandinavian plants exhibit
instances of discontinuous distribution, thus showing their ancient origin; and
there is altogether nothing in the fauna and flora of that country which might
lead us to believe that these were exterminated during the Glacial period and
reintroduced subsequently. The climate during that period in Scandinavia was
probably more equable and moister, with a greater snowfall in winter and with
less sun to melt the snow during summer, so that the development of glaciers
took more formidable dimensions, chiefly on the east side. The lowlands of
Sweden were covered by the sea, whilst many of the valleys were choked with
great glaciers, which cast off portions of ice as they reached the sea, just as
the Greenland and other northern glaciers do.
A country which at the present day probably somewhat
resembles the former Scandinavia climatically is Tierra
del Fuego, in the extreme south of South
America. Though there is an abundant snowfall,
so that glaciers reach the sea in many parts of the
country, the flora has been described by travellers
as luxuriant; and it appears that the fauna also is
richer than might be expected from the cheerless climate.
Towards the latter part of the Glacial
period the land-connection between Scandinavia, Spitsbergen,
and Greenland broke down, and the waters of the Arctic
and Atlantic Oceans joined. Whether it was at
this time or later that the other land-connection
between Scandinavia and Scotland collapsed is difficult
to determine; but it is certain, I think, that Scotland
was still united with Ireland even after these two
great land-bridges ceased to exist.
SUMMARY OF CHAPTER IV.
The faunThe fauna of the Arctic Regions is
much poorer than that of the other regions which are
dealt with in this work. In some groups, such
as Reptiles and Amphibia, there are no representatives
at all, but no doubt a larger number of species existed
there in earlier Tertiary times. At least we
have fossil evidence that during the Miocene Epoch
plants of many families flourished in Greenland of
which no vestige is now left in the Polar area.
Climatic conditions must therefore have changed, as
in Europe. A gradual refrigeration took place,
owing probably to the slow withdrawal of the current
which supplied the Arctic Sea with warmth. Greenland
and Europe were then connected, and the Arctic Ocean
was separated from the Atlantic. This land-connection
is supposed to have lain far north between Scandinavia,
Spitsbergen, and Greenland, and must have persisted
until towards the end of the Glacial period.
As the temperature decreased and the
land-area available in the north diminished, the surplus
population, consisting of animals and plants, and
possibly also of human beings, moved southward.
We have traces in Europe, and especially in the British
Islands, of a very early migration from the north
in the so-called American plants and in the freshwater
sponges. The geographical distribution of some
of the Arctic species of mammals is referred to in
greater detail, to show how the relative age of their
entry into Europe can be determined. Two forms
of Reindeer, resembling the Barren-ground and Woodland
varieties, have been met with in European deposits,
but only the former occurs in Ireland and the south
of France, whilst eastward the other becomes more common,
and finally is the only one found. It is believed
that the Barren-ground is the older form as far as
Europe is concerned, and that it came to us with the
Arctic migration, and that the other Reindeer reached
Europe much later from Siberia, when Ireland had already
become detached from England. The range of the
Arctic Hare is equally instructive. It must have
been a native of Europe since early glacial or pre-glacial
times before the common English Hare had
made its appearance in Central Europe. Along
with other Arctic forms, it entered Northern Europe
directly from the Arctic Regions, by means of the former
land-connection which joined, as I remarked, Lapland
with Spitsbergen, Greenland, and North America.
There need not have been a post-glacial connection
between Europe and Greenland; the present flora of
that country may have survived the Glacial period
in the Arctic Regions, as has been maintained by some
botanists and other authorities. Professor Forbes
argued from the occurrence of the same species of shore
mollusca on the coast of Finmark and Greenland that
these two countries were not long ago joined, so that
a slow migration from west to east along an ancient
coast-line could have taken place. That such a
migration actually occurred is further made probable,
judging from the presence of American mollusca in
the Crag deposits on the east coast of England.
These came into the North Sea in the first place direct
from the Arctic Ocean at a time when the two oceans
freely communicated with one another across the lowlands
of Northern Russia, Northern Germany, and Holland.
Arctic shells are also found below the boulder-clay
on the Baltic coast, and a free communication such
as indicated is generally held to have taken place
at no very distant date. The so-called “relict
species” marine animals left in freshwater
lakes in districts formerly covered by this sea lend
some support to this view. But the view that the
continental boulder-clay is a marine deposit is not
now held except by a few, though I here bring it forward
again, as it seems to me to fit in so much better
with the known facts of distribution. The sea
just referred to probably existed throughout the greater
part of the Glacial period; and icebergs, which originated
from the Scandinavian glaciers, would have brought
detritus and boulders to the lowlands. Scandinavia
was then connected with Scotland, and England with
France.