The larger foliose and fruticose lichens are now fairly well known and
described for Europe, and the knowledge of lichens in other continents
is gradually increasing. It is the smaller crustaceous forms that baffle
the investigator. The distribution of all lichens over the surface of the
earth is controlled by two principal factors, climate and substratum; for
although lichens as a rule require only support, they are most of them
restricted to one or another particular substratum, either organic or
inorganic. As organisms which develop slowly, they require an unchanging
substratum, and as sun-plants they avoid deeply shaded woodlands: their
occurrence thus depends to a large extent on the configuration and
general vegetation of the country.
Though so numerous and so widely distributed, lichens have not evolved
that great variety of families and genera characteristic of the allied
fungi and algae. They conform to a few leading types of structure,
and thus the Orders and Families are comparatively few, and more or
less universal. They are most of them undoubtedly very old plants and
were probably wide-spread before continents and climates had attained
their present stability. Arnold[1066] indeed considers that a large
part of the present-day lichens were almost certainly already evolved
at the end of the Tertiary period, and that they originated in a warm
or probably subtropical climate. As proof of this he cites such genera
as _Graphis_, _Thelotrema_ and _Arthonia_[1067] which are numerous in
the tropics though rare in the colder European countries; and he sees
further proof in the fact that many fruticose and gelatinous lichens do
not occur further north than the forest belt, though they are adapted
to cold conditions. Several genera that are abundant in the tropics are
represented outside these regions by only one or few species, as for
instance _Conotrema urceolatum_ and _Bombyliospora incana_.
During the Ice age of the Quaternary period, not many new species can
have arisen, and such forms as were not killed off must have been driven
towards the south. As the ice retreated the valleys were again stocked
with southern forms, and northern species were left behind on mountain
tops all over the globe.
In examining therefore the distribution of lichens, it will be found
that the distinction between different countries is relative, certain
families being more or less abundant in some regions than others, but,
in general, nearly all being represented. Certain species are universal,
where similar conditions prevail. This is especially true of those
species adapted to extreme cold, as that condition, normal in polar
regions, recurs even on the equator if the mountains reach the limit of
perpetual snow; the vertical distribution thus follows on the lines of
the horizontal.
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