No indications beyond those furnished by comparative anatomy help us to
unravel the phylogeny of the Collembola. In most respects, the shortened
abdomen, for example, they are more specialized than the Thysanura, and
most of the features in which they appear to be simple, such as the
absence of a tracheal system and of compound eyes, can be explained as
the result of degradation. In their insunken mouth and their jaws
retracted within the head-capsule, the Collembola resemble the
entotrophous division of the Thysanura (see APTERA), from which they are
probably descended.
From the thysanuroid stock of the Apterygota, the Exopterygota took
their rise. We have undoubted fossil evidence that winged insects lived
in the Devonian and became numerous in the Carboniferous period. These
ancient Exopterygota were synthetic in type, and included insects that
may, with probability, be regarded as ancestral to most of the existing
orders. It is hard to arrange the Exopterygota in a linear series, for
some of the orders that are remarkably primitive in some respects are
rather highly specialized in others. As regards wing-structure, the
Isoptera with the two pairs closely similar are the most primitive of
all winged insects; while in the paired mesodermal genital ducts, the
elongate cerci and the conspicuous maxillulae of their larvae the
Ephemeroptera retain notable ancestral characters. But the vestigial
jaws, numerous Malpighian tubes, and specialized wings of may-flies
forbid us to consider the order as on the whole primitive. So the
Dermaptera, which retain distinct maxillulae and have no ectodermal
genital ducts, have either specialized or aborted wings and a large
number of Malpighian tubes. The Corrodentia retain vestigial maxillulae
and two pairs of Malpighian tubes, but the wings are somewhat
specialized in the Copeognatha and absent in the degraded and parasitic
Mallophaga. The Plecoptera and Orthoptera agree in their numerous
Malpighian tubes and in the development of a folding anal area in the
hind-wing. As shown by the number and variety of species, the Orthoptera
are the most dominant order of this group. Eminently terrestrial in
habit, the differentiation of their fore-wings and hind-wings can be
traced from Carboniferous, isopteroid ancestors through intermediate
Mesozoic forms. The Plecoptera resemble the Ephemeroptera and Odonata in
the aquatic habits of their larvae, and by the occasional presence of
tufted thoracic gills in the imago exhibit an aquatic character unknown
in any other winged insects. The Odonata are in many imaginal and larval
characters highly specialized; yet they probably arose with the
Ephemeroptera as a divergent offshoot of the same primitive isopteroid
stock which developed more directly into the living Isoptera,
Plecoptera, Dermaptera and Orthoptera.
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