lizards), H. Gadow (cloaca of crocodile), J. F. v. Bemmelen (Sphenodon and others), W. H. Gaskell and H. Gadow (heart of tortoise).
Sense Organs.
1. Tegumentary Organs of some Tactile or other Sense.—Reptiles possess apparently no traces of those tegumentary sense organs which, belonging to the domains of the trigeminal and vagus nerves, have spread far over the body in fishes and batrachia. They were developed by those classes in correlation with their essentially aquatic life. This does not apply to the reptiles which, as a class, are of absolutely terrestrial origin. Nevertheless all recent reptiles possess numerous low sense-organs, “tactile bodies,” in most parts of the skin, connected with the regional, spinal nerves. They are most obvious in snakes, appearing as one or more little colourless spots near the apex of each scale on the back. The spot is formed by a little cluster of epidermal cells, connected with a sensory nerve. Their lowest stage they show in Sphenodon and in lizards, whilst in crocodiles they have reached a higher stage, at the bottom of the pit, since the tactile bodies, mostly several together, have sunk into the cutis, below the epiderm, forming a little pit, mostly near to the anterior margin of the flat scutes. They are most obvious on the belly of crocodiles, whilst in the American alligator such pits are scarcer, not because the organs are absent, but because these have sunk still farther into the skin. The last stage is that met with in tortoises, which possess such tactile bodies in considerable numbers in the softer subepidermal layers, beneath the large horny shields which themselves show no traces of them.
2. Taste.—The respective organs do not seem to have been investigated. That they exist is amply proved by the careful predilection for certain kinds of food which is shown especially by vegetarian tortoises and lizards, independent of smell. Many lizards are, for instance, very fond of sugar.
3. Nose.—The sense of smell is well developed in all reptiles. In none is the olfactory organ degraded; that the nasal passages, the nose itself, are never degraded is explained by the fact that all reptiles invariably breathe through the nose, except snakes during the act of swallowing their prey. The nostrils, always paired, are frequently provided with valves, to shut out the water, or sand. In some water tortoises, e.g. Trionyx, Cholys, the nostrils are prolonged into a soft, unpaired proboscis. Double tubes exist in the snake Herpeton (see Snakes, Opisthoglypha). The nostril leads into an antrum or vestibulum, this again into the nasal cavity proper, at the dorsal farther end enters the olfactory nerve, whilst ventrally it leads into the nasolaryngeal duct, with its posterior narial opening, or choana. The ducts are short in snakes and lizards, the choanae lying in the front part of the palate, but in tortoises and crocodiles they are placed far backwards, as has been described under Skull above. Into the nasal cavity projects, from the septum, a concha, least developed in tortoises, most in lizards and snakes. Crocodiles show a beginning of separation into several conchae as in birds and mammals. A large nasal gland lies against the lateral, or ventral, side of the outer wall of the nasal cavity, into which also opens the naso-lacrymal duct. Jacobson's organ, of uncertain function, is present in most reptiles. It is paired. In tortoises it is still placed within its nasal cavity, against the median wall, and is still nothing but a recess of the same and its mucous lining. In lizards and snakes the organ has become completely separated from the nasal cavity, lying below it and opening, each by a separate passage, into the palate mouth, close to or still within the choanae. In snakes it is mushroom-shaped, with a very short stalk. It lies immediately below the floor of the nasal capsule, and the membranous wall of the cavity on which it lies is covered and protected by a bone, commonly called the turbinal, which extends out from the median nasal system to the maxilla. In crocodiles these organs are vestigial and soon disappear.
4. Ear.—In crocodiles the outer ear lies in a recess, dorsally overhung by the lateral edge of the bony squamoso-frontal bridge; it carries a flap of skin, provided with muscles, to close the ear tightly. In lizards the outer ear is quite unprotected, and when the meatus is very short and wide, the drum is quite exposed. No reptiles possess cartilages comparable to the mammalian outer ear. Sphenodon, chameleons, snakes have no outer ear, the skin passing over the region. So also in tortoises, but in some of the aquatic kinds its position is well indicated by softer land thinner skin; in others, for instance marine turtles, a thick leathery plug, or a bigger scale marks the former-position. In various lizards, chiefly burrowing in sand, the ear passage is very narrow, or closed. The middle ear or tympanic cavity is quite obliterated in snakes, Amphisbaenas and some other snake-shaped lizards. In Anguis may exist individual traces. The cavity communicates with the mouth. In lizards the communication is a wide recess, lined with black pigment, so that in these creatures the whole auditory chain can easily be inspected from the opened mouth. In tortoises the recesses are contracted into the Eustachian tubes, each of which opens by a separate aperture into the roof of the mouth. In the crocodiles part of the cavities is transformed into an intricate system of canals and passages. The two Eustachian tubes open together in the mid-lines protected by a valve, between the basioccipital and basisphenoid; thence arises a median passage which with lateral arms and loops extends upward through the occiput into the cranial roof, communicating with the tympanic cavity, and further continued through the quad rates and beyond into the mandibles, by the siphonium.
In spite of the obliterated tympanic cavity of snakes, and the closed up outer ear passage and absence of a tympanic membrane in snakes and tortoises, these creatures can hear very well. The same applies to Sphenodon, but it seems doubtful whether chameleons can hear.
Through the whole middle ear, from the fenestra ovalis to the drum-membrane, stretches the chain of auditory ossicles or cartilages, partly attached to the posterior wall by the common lining membrane. The arrangement appears simplest in snakes, in chameleons and in tortoises, not because it is primitive but because it is so much reduced, partly in correlation with the abolition of the outer ear. In these creatures the columella goes as a bony, slender rod straight to the middle of the quadrate, against which it leans, or with which it articulates by a short piece of cartilage, the extra-columella. Here the whole chain ends. It looks like a proof that columella = stapes, extracolumella = incus, and quadrate= malleus; or, with the usual ignoring of the little extra-columellar piece, that quadrate = incus, Gegenbaur's favourite impossibility. In those lizards which have a tympanic membrane conditions are far less reduced. The extra-columellar piece sends out three distal processes; one leans on to the middle of the tympanic membrane, the second usually is fastened to the bony dorsal rim of the meatus, the third is directed downwards and is continued as a thin ligament towards the inner angle of the articular of the mandible, but before reaching this it comes to grief, being squeezed in between the quadrate and the posterior end of the pterygoid. The hyoid proper is of no account in snakes and tortoises, since it is reduced to Very short distal pieces attached to the base of the tongue; but in lizards it remains in its original length, or it even lengthens, and shows many vagaries in its position and attachments. In embryos of Sphenodon and lizards it arises from near the junction of the columella with the extra-columella. It becomes very long, too long for the available space (perhaps correlated with lingual functions), and it forms a high loop, thereby causing the peculiar loop of the chorda tympani; the upward bend of the hyoid becomes connected with the parotic process of the cranium. Next aborts the portion between this connexion and the original proximal end of the hyoid, near the columellar mass. The upper end of the hyoid either remains attached to the parotic process (various lizards and Sphenodon) whence the lingual apparatus remains suspended, or the hyoid, having broken loose, leaves a little cartilage, Versluy's cartilage, behind, at the end of the parotic process, and the hyoid horn remains free, in the majority of lizards. In Sphenodon, whilst
