_Origin of Metamorphosis._--In connexion with the question whether
metamorphosis has been gradually acquired, we have to consider two
aspects, viz. the bionomic nature of metamorphosis, and to what extent
it existed in primitive insects. Bionomically, metamorphosis may be
defined as the sum of adaptations that have gradually fitted the larva
(caterpillar or maggot) for one kind of life, the fly for another. So
that we may conclude that the factors of evolution would favour its
development. With regard to its occurrence in primitive insects, our
knowledge of the geological record is most imperfect, but so far as it
goes it supports the conclusion that holometabolism (i.e. extreme
metamorphosis) is a comparatively recent phenomenon of insect life. None
of the groups of existing Endopterygota have been traced with certainty
farther back than the Mesozoic epoch, and all the numerous Palaeozoic
insect-fossils seem to belong to forms that possessed only imperfect
metamorphosis. The only doubt arises from the existence of insect
remains, referred to the order Coleoptera, in the Silesian Culm of
Steinkunzendorf near Reichenbach. The oldest larva known, _Mormolucoides
articulatus_, is from the New Red Sandstone of Connecticut; it belongs
to the _Sialidae_, one of the lowest forms of Holometabola. It is now,
in fact, generally admitted that metamorphosis has been acquired
comparatively recently, and Scudder in his review of the earliest fossil
insects states that "their metamorphoses were simple and incomplete, the
young leaving the egg with the form of the parent, but without wings,
the assumption of which required no quiescent stage before maturity."
It has been previously remarked that the phenomena of holometabolism are
connected with the development of wings inside the body (except in the
case of the fleas, where there are no wings in the perfect insect). Of
existing insects 90% belong to the Endopterygota. At the same time we
have no evidence that any Endopterygota existed amongst Palaeozoic
insects, so that the phenomena of endopterygotism are comparatively
recent, and we are led to infer that the Endopterygota owe their origin
to the older Exopterygota. In Endopterygota the wings commence their
development as invaginations of the hypodermis, while in Exopterygota
the wings begin--and always remain--as external folds or evaginations.
The two modes of growth are directly opposed, and at first sight it
appears that this fact negatives the view that Endopterygota have been
derived from Exopterygota.
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