
Remaining Presacral Vertebrae
The succeeding presacral vertebrae bear a pair of foramina close to the median line on the ventral surface of each centrum (Fig 4 B). Mahendra (1950) states that these openings, serving for the passage of blood vessels, are well developed in Geckonidae and Xantusiidae, and probably represent a primitive feature. Apparently such apertures in presacral vertebrae were recorded for the first time in any animal by Mookerjee and Das (1933), who in investigating Typhlops braminus

found a single opening in each vertebra as far back as the pelvic region. Mookerjee and Das indicate that each ventral aperture passes through the centrum and up to the neural canal. They state that branches of the vertebral artery, instead of passing through the intervertebral spaces along with the spinal nerves, enter the spinal canal through these openings.
Ribs are not present on the atlas or axis, and not normally as in some geckos, on the third vertebra. The fourth, fifth and sixth vertebrae, however, have short bony ribs, somewhat expanded at both ends. Attached to their distal ends, these ribs bear delicate cartilaginous processes which, being bifurcated terminally, have a “fish-tail” appearance (Fig. 4 D) These processes increase in size from before backwards so that the last which are on the sixth vertebra are much longer than the rest. The first of the series, which is on the fourth cervical rib, can usually be seen anteriorly to the suprascapula. The others are covered by the latter. It is difficult in alizarin transparencies to see much muscular detail, but the first process on each side appears to have a muscular connexion with the suprascapula.
In Naultinus, although the anterior two cartilaginous processes on each side are forked, the most posterior one is single, slender and elongated. Calcification of these structures may occur in all species.
Mahendra (1935, 1950) describes similar structures in Hemidactylus occurring on the fourth, fifth, sixth and sometimes seventh cervical vertebrae as being presumably uncinate processes. He duly records them for the first time in a living Lacertilian. Their occurrence in geckos, however, is by no means exceptional, for we have subsequently noted them in alizarin transparencies of Jamaican and Australian genera. Furthermore, the term “uncinate” should not be used for these processes without reservation, and it must be pointed out that these geckos do not possess such structures on the sternal and post-sternal ribs where uncinate processes are normally found; that they would be the only living animals possessing uncinate processes on cervical ribs; that their cervical ribs are in two portions, proximal and distal. The processes do not arise from the proximal portion of the rib as is typically the case with uncinate ribs, but from the distal portion.
The seventh and eighth vertebrae have long, slender, dorsal ribs, each jointed distally with a short, blunt, ventral piece of cartilage which does not reach the sternum. These ventral pieces of cartilage are indistinguishable from the sternal ribs, though they are much shorter. In general, it seems clear that the seventh and eighth vertebrae are in the same series as those posterior to them, rather than with the more anterior vertebrae of the column On this basis it is reasonable to follow Camp (1923) in relegating these two vertebrae to the dorsal series, thus regarding the cervical vertebrae as six in number.
In Hoplodactylus, the ninth and tenth vertebrae are connected directly with the sternum by sternal ribs, which are jointed with the dorsal ribs. The sternal ribs of the eleventh and twelfth vertebrae are connected with the xiphisternum. while the sternal ribs of the thirteenth vertebra almost meet in the mid-ventral line, but do not quite connect with the xiphisternum Sometimes only the twelfth vertebra is connected with the xiphisternum.
In Naultinus, the ninth and tenth vertebrae are connected to the sternum proper, while the eleventh, twelfth and thirteenth are connected with the xiphisternum. In general, the sternal ribs are better developed in Naultinus than in Hoplodactylus and they approach the mid-ventral line more closely and for a long distance posteriorly.
The sternal ribs are always cartilaginous, but may be calcified, even heavily so in the case of Hoplodactylus duvauceli. In the fourteenth to twentieth vertebrae inclusively, the sternal ribs are free ventrally and diminish in size posteriorly. The twenty-first to twenty-fifth vertebrae have their dorsal ribs decreasing in size and have lost almost all trace of the sternal portions. The twenty-sixth vertebra, normally the last presacral vertebra, has no ribs.

Text-fig. 4.—Vertebrae of Naultinus elegans. A, Second and third cervical vertebrae, lateral view. B, Eighteenth presacral vertebra, ventral view. C, Eighteenth presacral vertebra, lateral view. D, Fourth vertebra, anterior view, buf, foramina for blood vessels; cn, centrum; cp, distal cartilaginous process of rib, cpf, capitular facet; cpt, capitulum of rib; cv, cervical rib; hyp, hypapophysis; ic, intercentrum; od, odontoid process; poz, postzygapophysis; prz, prezygapophysis; r, rib; tb, tuberculum of rib.

Noble (1921) was surprised to find in dorsal ribs of Sphaerodactylus a cartilaginous or fibro-cartilaginous band which extended from the angular portion of the head of the rib to the middle of the neural arch of the vertebra. He suggested that this structure, which was distinct from the capitular head of the rib was a last vestige of the tuberculum found in the double headed ribs of primitive reptiles but believed to be lost in Lacertilia. A similar band of tissue representing the tuberculum occurs in the New Zealand geckos. When sections of dorsal vertebrae and ribs of Hoplodactylus pacificus were examined, it was noted that the tuberculum was distinctly bony. (Fig. 4C.)
Camp (1923) did not find parasternal ribs, (abdominal ribs or gastralia) in any of the geckonid genera at his disposal. On the other hand, Mahendra (1950) notes that alizarin preparations of Hemidactylus flaviviridis show the presence of a few delicate parasternal ribs in the superficial parts of the rectus muscle just behind the sternum. No traces of such gastralia have been observed in any of the New Zealand geckos.
The sacrum is found normally from the twenty-seventh and twenty-eighth vertebrae, the “sacral processes” of which are widely expanded and connected with the ilia. A great deal of variation occurs in this region, and more posterior vertebrae may take part in the formation of the sacrum if one of the anterior processes fails to develop. In this connexion there may be asymmetry on right and left sides.
The first vertebrae of the caudal series have transverse processes diminishing in size from before backwards, and may show traces of caudal ribs as short, blunt, much reduced structures. These caudal ribs are calcified and are particularly well seen in Naultinus, where some four or more occur on each side. El-Toubi and Khalil (1950), in a comparative study of the osteology of some Egyptian geckos, drew attention to the presence of caudal ribs, which as far as they were aware had not previously been recorded in living Lacertilia. These small caudal ribs, articulating with the transverse processes of the first four or so caudal vertebrae were observed by El-Toubi and Khalil on one side only in alizarin transparencies. In the New Zealand geckos they are clearly seen on both sides.
It has been noted that the position of the sacrum in the New Zealand geckos is not fixed and that a caudal vertebra may function as a sacral vertebra. In such a case, both the caudal transverse process and its distal rib are expanded for articulation with the ilium. This variation is of interest considering the uncertainty that has existed for some time regarding the nature of sacral processes in lizards. Moodie (1907), investigating the sacrum of the Lacertilia, claimed from embryological evidence that “It can thus be very definitely stated that the Lacertilia occupy an isolated place among all other known reptiles in not having any sacral ribs whatever.” On the other hand, Kamel (1951), in studying the development of ribs in the sacral region of Chalcides ocellatus found that the sacral ribs are quite distinct. They chondrify and become united with the transverse processes.
Typically the processes of normal sacral vertebrae in New Zealand geckos are much expanded, and each bears distally a wide, articulating flange of uncalcified cartilage. This fuses very often with the corresponding structure on the neighbouring sacral process.
In the caudal region, Y-shaped chevron bones are attached to the ventral region of the intervertebral joints, beginning usually between the third and fourth or fourth and fifth caudal vertebrae.
Autotomy occurs very commonly in the fifth caudal vertebra. The first four vertebrae of the series are usually pygals and lack the transverse unossified septum which passes through the body of the centrum of the succeeding caudal vertebrae. In the autotomous region of the tail the vertebrae become extremely elongated.
