So close is this similarity, from the comparative point of view, between
the dorsal head-shield of the Osteostraci and the dorsal cephalic region of
Ammocoetes that it justifies us in taking Ammocoetes as the nearest living
representative of such types; it is justifiable, therefore, to interpret by
means of Ammocoetes the position of other organs in these forms. First and
foremost is the hard plate {339}known as the post-orbital plate, so
invariably found. In Fig. 134, C, I have inserted (_cr._) the position of
the membranous cranium of Ammocoetes, and it is immediately evident that
the primordial cranium of the Osteostraci must occupy the exact position
indicated by this median hard plate. For this very reason this median plate
would be harder than the rest in order to afford a better protection to the
brain underneath. This plate, because of its position, may well receive the
same name as the similar plate in the trilobite and various palæostracans
and be called the glabellum.
EVIDENCE OF SEGMENTATION IN THE HEAD-SHIELD--FORMATION OF CRANIUM.
We may thus conceive the position of the nose, lateral eyes, median eyes,
and cranium in these old fishes. In addition, other indications of a
segmentation in this head-region have been found. The most striking of all
the specimens hitherto discovered are some of _Thyestes verrucosus_,
discovered by Rohon, in which the dorsal shield has been removed, and so we
are able to see what that dorsal shield covered.
In Fig. 136, I reproduce his drawing of one of his specimens from the
dorsal and lateral aspects. These drawings show that the frontal part of
the shield covered a markedly segmented part of the animal; five distinct
segments are visible apart from the median most anterior region. This
segmented region is entirely confined to the prosomatic region, _i.e._ to
the region innervated by the trigeminal nerve. An indication of similar
markings is given in Lankester's figure of _Eukeraspis pustuliferus_ (see
Fig. 127, B), and, indeed, evidence of a segmentation under the
antero-lateral border of the head-shield is recognized at the present time,
not only in the Cephalaspidæ, but also in the Pteraspidæ, as was pointed
out to me by Smith Woodward in the specimens at the British Museum. Also,
in _Cyathaspis_, Jaekel has drawn attention to markings of a similar
segmental nature (Fig. 137).
There seems, then, little doubt but that these primitive fishes possessed
something in this region which was of a segmental character, and indicated
at least five segments, probably more.
Rohon entitles his discovery 'the segmentation of the primordial cranium.'
It would, I think, be better to call it the segmentation of {340}the
anterior region of the head, for that is in reality what his figures show,
not the segmentation of the primordial cranium, which, to judge from
Ammocoetes, was confined to the region of the glabellum.
What is the interpretation of this appearance?
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