spiracle into a pre-spiracular portion—the main portion of which passes to the mucous membrane of the palate as the palatine (pnVII.)—and a postspiracular portion, the hyomandibular (hy.) trunk which supplies the muscles of the hyoid arch and also sends a few sensory fibres to the lining of the spiracle, the floor of mouth and pharynx and the skin of the lower jaw. Combined with the main trunk of the facial are branches belonging to the lateralis system.
Lateralis Group of Nerves.—The lateralis group of nerves are charged with the innervation of the system of cutaneous sense organs and are all connected with the same central region in the medulla. A special sensory area of the ectoderm becomes involuted below the surface to form the otocyst, and the nerve fibres belonging to this form the auditory nerve (VIII.). Other portions of the lateralis group become mixed up with various other cranial nerves as follows:
(a) Facial portion.
(1) Ophthalmicus superficialis (op.s.VII.): passes to lining of nose or to the lateral line organs of the dorsal part of snout.
(2) Buccal (bucc.VII): lies close to maxillary division of V. and passes to the sensory canals of the lower side of the snout.
(3) External mandibular (md.ex.): lies in close association with the mandibular division of V., supplies the sensory canals of the lower jaw and hyoid region.
Lateralis vagi (l.n.X.) becomes closely associated with the vagus. It supplies the lateral line organs of the trunk.
In the lamprey and in Dipnoans the lateralis vagi loses its superficial position in the adult and comes into close relation with the notochord.
In Actinopterygians and at least some Selachians a lateralis set of fibres is associated with IX., and in the former fishes a conspicuous trunk of lateralis fibres passes to some or all (Gadus) of the fins. This has been called the lateralis accessorius and is apparently connected with V., VII., IX., X. and certain spinal nerves.[1]
Vagus Group (IX., X., XI.).—The glossopharyngeus (IX.) forks over the first branchial cleft (pretrematic and post-trematic branches) and also gives off a palatine branch (pn.IX.). In some cases (various Selachians, Ganoids and Teleosts) it would seem that IX. includes a few fibres of the lateralis group.
Vagus (X.) is shown by its multiple roots arising from the medulla and also by the character of its peripheral distribution to be a compound structure formed by the fusion of a number of originally distinct nerves. It consists of (1) a number of branchial branches (X.¹ X.² &c.), one of which forks over each gill cleft behind the hyobranchial and which may (Selachians) arise by separate roots from the medulla; (2) an intestinal branch (v.n.X.) arising behind the last branchial and innervating the wall of the oesophagus and stomach and it may be even the intestine throughout the greater part of its length (Myxine).
The accessorius (XI.) is not in fishes separated as a distinct nerve from the vagus.
With increased development of the brain its hinder portion, giving rise to the vagus system, has apparently come to encroach on the anterior portion of the spinal cord, with the result that a number of spinal nerves have become reduced to a less or more vestigial condition. The dorsal roots of these nerves disappear entirely in the adult, but the ventral roots persist and are to be seen arising ventrally to the vagus roots. They supply certain muscles of the pectoral fins and of the visceral arches and are known as spino-occipital nerves.[2]
These nerves are divisible into an anterior more ancient set—the occipital nerves—and a posterior set of more recent origin—(occipito-spinal nerves). In Selachians 1-5 pairs of occipital nerves alone are recognizable: in Dipnoans 2-3 pairs of occipital and 2-3 pairs of occipito-spinal: in Ganoids 1-2 pairs occipital and 1-5 pairs occipito-spinal; in Teleosts finally the occipital nerves have entirely disappeared while there are 2 pairs of occipito-spinal. In Cyclostomes no special spino-occipital nerves have been described.
The fibres corresponding with those of the Hypoglossus (XII.) of higher vertebrates spring from the anterior spinal nerves, which are here, as indeed in Amphibia, still free from the cranium.
Sympathetic.—The sympathetic portion of the nervous system does not in fishes attain the same degree of differentiation as in the higher groups. In Cyclostomes it is apparently represented by a fine plexus with small ganglia found in the neighbourhood of the dorsal aorta and on the surface of the heart and receiving branches from the spinal nerves. In Selachians also a plexus occurs in the neighbourhood of the cardinal veins and extends over the viscera: it receives visceral branches from the anterior spinal nerves. In Teleosts the plexus has become condensed to form a definite sympathetic trunk on each side, extending forwards into the head and communicating with the ganglia of certain of the cranial nerves.
- (J. G. K.)
V. Distribution in Time and Space
The origin of Vertebrates, and how far back in time they extend, is unknown. The earliest fishes were in all probability devoid of hard parts and traces of their existence can scarcely be expected to be found. The hypothesis that they may be derived from the early Crustaceans, or Arachnids, is chiefly based on the somewhat striking resemblance which the mailed fishes of the Silurian period (Ostracodermi) bear to the Arthropods of that remote time, a resemblance, however, very superficial and regarded by most morphologists as an interesting example of mimetic resemblance—whatever this term may be taken to mean. The minute denticles known as conodonts, which first appear in the Ordovician, were once looked upon as teeth of Cyclostomes, but their histological structure does not afford any support to the identification and they are now generally dismissed altogether from the Vertebrates. As a compensation the Lower Silurian of Russia has yielded small teeth or spines which seem to have really belonged to fishes, although their exact affinities are not known (Palaeodus and Archodus of J. V. Rohon).
It is not until we reach the Upper Silurian that satisfactory remains of unquestionable fishes are found, and here they suddenly appear in a considerable variety of forms, very unlike modern fishes in every respect, but so highly developed as to convince us that we have to search in much earlier formations for their ancestors. These Upper Silurian fishes are the Coelolepidae, the Ateleaspidae, the Birkeniidae, the Pteraspidae, the Tremataspidae and the Cephalaspidae, all referred to the Ostracophori. The three last types persist in the Devonian, in the middle of which period the Osteolepid Crossopterygii, the Dipneusti and the Arthrodira suddenly appear. The most primitive Selachian (Cladoselache), the Acanthodian Selachians (Diplacanthidae), the Chimaerids (Ptyctodus), and the Palaeoniscid ganoids (Chirolepis) appear in the Upper Devonian, along with the problematic Palaeospondylus.
In the Carboniferous period, the Ostracophori and Arthrodira have disappeared, the Crossopterygii and Dipneusti are still abundant, and the Selachians (Pleuracanthus, Acanthodians, truesharks) and Chondrostean ganoids (Palaeoniscidae and Platysomidae) are predominant. In the Upper Permian the Holostean ganoids (Acanthophorus) make their appearance, and the group becomes dominant in the Jurassic and the Lower Cretaceous. In the Trias, the Crossopterygii and Dipneusti dwindle in variety and the Ceratodontidae appear; the Chondrostean and Holostean ganoids are about equally represented, and are supplemented in the Jurassic by the first, annectant representatives of the Teleostei (Pholidophoridae, Leptolepidae). In the latter period, the Holostean ganoids are predominant, and with them we find numerous Cestraciont sharks, some primitive skates (Squatinidae and Rhinobatidae), Chimaerids and numerous Coelacanthid crossopterygians.
The fish-fauna of the Lower Cretaceous is similar to that of the Jurassic, whilst that of the Chalk and other Upper Cretaceous formations is quite modern in aspect, with only a slight admixture of Coelacanthid crossopterygians and Holostean ganoids, the Teleosteans being abundantly represented by Elopidae, Albulidae, Halosauridae, Scopelidae and Berycidae,
