The thickness of the thallus varies considerably according to the
species. In some _Pertusariae_ with a stoutish irregular crust there is
a narrow amorphous cortical layer of almost obliterated cells, a thin
gonidial zone about 35 µ in width and a massive rather dense medulla
of colourless hyphae. Darbishire[336] has described and figured in
_Varicellaria microsticta_, one of the Pertusariaceae, single hyphae
that extend like beams across the wide medulla and connect the two
cortices. In some _Lecanorae_ and _Lecideae_ there is, on the contrary,
an extremely thin thallus consisting of groups of algae and loose fungal
filaments, which grow over and between the dead cork cells of the
outer bark. On palings, there is often a fairly substantial granular
crust present, with a gonidial zone up to about 80 µ thick, while the
underlying or medullary hyphae burrow among the dead wood fibres.
_b._ HYPOPHLOEODAL LICHENS. These immersed lichens are comparable
with the endolithic species of the rock formations, as their thallus
is almost entirely developed under the outer bark of the tree. They
are recognizable, even in the absence of any fructification, by the
somewhat shining brownish, white or olive-green patches that indicate
the underlying lichen. This type of thallus occurs in widely separated
families and genera, _Lecidea_, _Lecanora_, etc., but it is most constant
in Graphideae and in those Pyrenolichens of which the algal symbiont
belongs to the genus _Trentepohlia_. The development of these lichens is
of peculiar interest as it has been proved that though both symbionts
are embedded in the corky tissues, the hyphae arrive there first, and,
at some later stage, are followed by the gonidia. There is therefore no
question of the alga being a “captured slave” or “unwilling mate.”
Frank[337] made a thorough study of several subcortical forms. He found
that in _Arthonia radiata_, the first outwardly visible indication of the
presence of the lichen on ash bark was a greenish spot quite distinct
from the normal dull-grey colour of the periderm. Usually the spots are
round in outline, but they tend to become ellipsoid in a horizontal
direction, being influenced by the growth in thickness of the tree.
At this early stage only hyphae are present; Bornet[338] as well as
Frank described the outer periderm cells as penetrated and crammed with
the colourless slender filaments. Lindau[339], in a more recent work,
disputes that statement: he found that the hyphae invariably grew between
the dead cork cells, splitting them up and disintegrating the bark, but
never piercing the membranes. The purely prothallic condition, as a weft
of closely entangled hyphae, may last, Frank considers, for a long period
in an almost quiescent condition—possibly for several years—before the
gonidia arrive.
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