In Leptolinae the embryonic development culminates in a polyp, which
is usually formed by fixation of a planula (parenchymula), rarely by
fixation of an actinula. The planula may fix itself (1) by one end,
and then becomes the hydrocaulus and hydranth, while the hydrorhiza
grows out from the base; or (2) partly by one side and then gives rise
to the hydrorhiza as well as to the other parts of the polyp; or (3)
entirely by its side, and then forms a recumbent hydrorhiza from which
a polyp appears to be budded as an upgrowth.
In Trachylinae the development produces always a medusa, and there is
no polyp-stage. The medusa arises direct from the actinula-stage and
there is no entocodon formed, as in the budding described above.
_Life-cycles of the Hydromedusae._--The life-cycle of the Leptolinae
consists of an alternation of generations in which non-sexual
individuals, polyps, produce by budding sexual individuals, medusae,
which give rise by the sexual process to the non-sexual polyps again,
so completing the cycle. Hence the alternation is of the type termed
metagenesis. The Leptolinae are chiefly forms belonging to the inshore
fauna. The Trachylinae, on the other hand, are above all oceanic
forms, and have no polyp-stage, and hence there is typically no
alternation in their life-cycle. It is commonly assumed that the
Trachylinae are forms which have lost the alternation of generations
possessed by them ancestrally, through secondary simplification of the
life-cycle. Hence the Trachylinae are termed "hypogenetic" medusae to
contrast them with the metagenetic Leptolinae. The whole question has,
however, been argued at length by W. K. Brooks [4], who adduces strong
evidence for a contrary view, that is to say, for regarding the direct
type of development seen in Trachylinae as more primitive, and the
metagenesis seen in Leptolinae as a secondary complication introduced
into the life-cycle by the acquisition of _larval budding_. The polyp
is regarded, on this view, as a form phylogenetically older than the
medusa, in short, as nothing more than a sessile actinula. In
Trachylinae the polyp-stage is passed over, and is represented only by
the actinula as a transitory embryonic stage. In Leptolinae the
actinula becomes the sessile polyp which has acquired the power of
budding and producing individuals either of its own or of a higher
rank; it represents a persistent larval stage and remains in a
sexually immature condition as a neutral individual, sex being an
attribute only of the final stage in the development, namely the
medusa. The polyp of the Leptolinae has reached the limit of its
individual development and is incapable of becoming itself a medusa,
but only produces medusa-buds; hence a true alternation of generations
is produced. In Trachylinae also the beginnings of a similar
metagenesis can be found. Thus in _Cunina octonaria_, the ovum
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