Such an arrangement suggests that in the ancient arthropod type a double segmentation existed, viz. a segmentation of the body, and a segmentation due to the appendages. Undoubtedly, the segments originally corresponded absolutely as in Branchipus, and every appendage was attached to a well-defined separate body-segment. In, however, such an ancient type as Limulus, though the segmentation may be spoken of as twofold, yet the number of segments in the prosomatic and mesosomatic regions are much more clearly marked out by the appendages than by the divisions of the soma; for, in the prosomatic region such a fusion of somatic segments to form the tergal prosomatic carapace has taken place that the segments of which it is composed are visible only in the young condition, while in the mesosomatic region the separate somatic segments, though fused to form the mesosomatic carapace, are still indicated by the entapophysial indentations.
Clearly, then, if the mesosomatic branchial appendages of forms related to Limulus were reduced to the branchial portion of the appendage, and that branchial portion became internal, just as is known to be the case in the scorpion group, we should obtain an animal in which the mesosomatic region would be characterized by a segmentation predominantly branchial, which might be termed, as in vertebrates, the branchiomeric segmentation, but yet would show indications of a corresponding somatic or mesomeric segmentation. The nerve supply to these segments would consist of—
1. The epimeral purely sensory nerves to the somatic surface, equivalent in the vertebrate to the ascending root of the trigeminal.
2. The mixed nerves to the internal branchial segments, equivalent in the vertebrate to the vagus, glossopharyngeal, and facial.
3. The motor nerves to the somatic muscles, equivalent in the vertebrate to the original nerve-supply to the somatic muscles belonging to these segments, i.e. to the muscles derived from van Wijhe's 4th, 5th, and 6th somites.
Further, the centres of origin of these appendage-nerves would form centres in the central nervous system separate from the centres of the motor nerves to the somatic muscles, just as the centres of origin of the motor parts of the facial, vagus, and glossopharyngeal nerves form groups of cells quite distinct from the centres for the hypoglossal, abducens, trochlear, and oculomotor nerves.
In fact, if the vertebrate branchial nerves are looked upon as the descendants of nerves which originally supplied branchial appendages, then every question connected with the branchial segmentation, with the origin and distribution of these nerves, receives a simple and adequate solution—a solution in exact agreement with the conclusion that the vertebrate arose from a palæostracan ancestor.
It would, therefore, be natural to expect that the earliest fishes breathed by means of branchial appendages situated internally, and that the evidence for such appendages would be much stronger in them than in more recent fishes.
Although we know nothing of the nature of the respiratory apparatus in the extinct fishes of Silurian times, we have still living, in the shape of Ammocœtes, a possible representative of such types. If, then, we find, as is the case, that the respiratory apparatus of Ammocœtes differs markedly from that of the rest of the fishes, and, indeed, from that of the adult form or Petromyzon, and that that very difference consists in a greater resemblance to internal branchial appendages in the case of Ammocœtes, then we may feel that the proof of the origin of the branchial apparatus of the vertebrate from the internal branchial appendages of the invertebrate has gained enormously.
The Respiratory Chamber of Ammocœtes.