If the bud, however, is destined to give rise not to a free medusa,
but to a gonophore, the development is similar but becomes arrested at
various points, according to the degree to which the gonophore is
degenerate. The entocodon is usually formed, proving the medusoid
nature of the bud, but in sporosacs the entocodon may be rudimentary
or absent altogether. The process of budding as above described may be
varied or complicated in various ways; thus a secondary, amnion-like,
ectodermal covering or ectotheca (fig. 45, C, ect.) may be formed over
all, as in _Garveia_, &c.; or the entocodon may remain solid and
without cavity until after the formation of the manubrium, or may
never acquire a cavity at all, as described above for the gonophores.
_Phylogenetic Significance of the Entocodon._--It is seen from the
foregoing account of medusa-budding that the entocodon is a very
important constituent of the bud, furnishing some of the most
essential portions of the medusa; its cavity becomes the sub-umbral
cavity, and its lining furnishes the ectodermal epithelium of the
manubrium and of the sub-umbral cavity as far as the edge of the
velum. Hence the entocodon represents a precocious formation of the
sub-umbral surface, equivalent to the peristome of the polyp,
differentiated in the bud prior to other portions of the organism
which must be regarded as antecedent to it in phylogeny.
If the three principal organ-systems of the medusa, namely mouth,
tentacles and umbrella, be considered in the light of phylogeny, it is
evident that the manubrium bearing the mouth must be the oldest, as
representing a common property of all the Coelentera, even of the
gastrula embryo of all Enterozoa. Next in order come the tentacles,
common to all Cnidaria. The special property of the medusa is the
umbrella, distinguishing the medusa at once from other morphological
types among the Coelentera. If, therefore, the formation of these
three systems of organs took place according to a strictly
phylogenetic sequence, we should expect them to appear in the order
set forth above (fig. 46, Ia, b, c). The nearest approach to the
phylogenetic sequence is seen in the budding of _Cunina_, where the
manubrium and mouth appear first, but the umbrella is formed before
the tentacles (fig. 46, IIa, b, c). In the indirect or coenogenetic
method of budding, the first two members of the sequence exhibited by
Cunina change places, and the umbrella is formed first, the manubrium
next, and then the tentacles; the actual mouth-perforation being
delayed to the very last (fig. 46, IVa, b, c). Hence the budding of
medusae exemplifies very clearly a common phenomenon in development, a
phylogenetic series of events completely dislocated in the ontogenetic
time-sequence.
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