Opetiosaurus Bucchichi. Eine neue fossile Eidechse aus der unteren Kreide von Lesina in Dalmatien



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B. The trunk.

The vertebral column, columna vertebralis, comprises 28 presacral and two sacral vertebrae, probably followed by 100 postsacrals, so that the number of all vertebrae came to about 130.

Of the presacral vertebrae, eight belong to the cervical series of the vertebral column. This decision is based on the presence of hypapophyses, as in many other Sauria (apophyse épineuse inférieure of Cuvier). The following 20 vertebrae can either all be termed dorsal vertebrae, or the last four, do17 through do20, can be termed lumbodorsal vertebrae in the sense of Bergmann30 because of their conspicuously shorter ribs. Lumbar vertebrae, which can be recognized as such by the absence of ribs, are not present in our fossil. The postsacral or caudal vertebrae are unfortunately partly in a bad preservational state, namely in the anterior section that immediately follows the sacrum; even a count of these with the relevant piece of the part has become completely impossible.
Cervical vertebrae.

The first three neck or cervical vertebrae (ce.) have remained attached to the occiput of the skull in the previously mentioned violent separation of the skull from the trunk. Unfortunately, and notwithstanding the careful chemical treatment of the counterpart (Pl. III, Fig. 1), I was unsuccesful in rendering completely clearly what derives from the atlas in particular. Below the pieces of bone that are attached to the occiput, whose several pieces were indicated in the description of the head according to their outlines, one sees, behind the suspensorial apparatus of the quadrate, and namely adjacent to the parotic process, a raised swelling of bone that continues further to the right under the suspensorium and may well be seen as the massa lateralis of the atlas. The second cervical vertebra, like the third, of which only a part is seen to the right on the slab, follow immediately posteriorly. They display their upper surface, namely as one rather broad arch each, to which the zygapophyses are joined laterally and on which a median, crest-like ridge rises. The latter corresponds to the neural spine.

Of the anterior presacral vertebrae, which are preserved on the large part, the first five, namely the 25th until finally the 21st, represent cervical vertebrae, namely the 4th through 8th, in virtue of the hypapophyses that are developed on them and are lacking in dorsal vertebrae. These hypapophyses arise on the underside of the vertebral centrum in the form of a median boss [“Erhabenheit”] that begins narrow, then gradually becomes broader posteriorly, and finally terminates in the usual position beneath the vertebral condyle in an ellipsoidal or nearly ball-like process, whose cap [“Kuppe”], however, appears largely shattered by the pressure of the rock mass. Because the undersurfaces of the vertebral centra are directed dorsally on the main stone slab as a result of the inverted position of our animal, the peculiarity just mentioned can clearly be discerned. The presence of these apophyses does not constitute a diagnostic character of any saurian group, as should [“soll”] be discussed more extensively later, for they are also better developed and in a more peculiar form in other [groups], e.g., in the Pythonomorpha. They also appear in many extant lizards31; Cuvier32 establishes their presence in Monitor niloticus and Calori33 in Monitor terrestris Aegypti (= Varanus arenarius Dum. et Bibr.) and in Lacerta viridis and L. ocellata, where their form is clearly reminiscent of the hypapophyses of our fossil from Lesina.

Otherwise the cervical vertebrae are already robustly developed; in size and form they differ only slightly from the dorsal vertebrae. They are anteriorly broad (18 mm), pass laterally into transverse processes, and narrow posteriorly (12 mm). On their under-surfaces lateral to the hypapophyses described above, their bodies or centra show isolated ridges of bone, predominantly aligned with the longitudinal axis, which were for the attachment of powerful muscles. A deep, concave, semilunate bulge on the anterior margin corresponds to the cotyle on the anterior end of the centrum. It was for the reception of the expanded [“erhabenen”] condyle on the posterior end of the next-most anterior vertebra, with which it articulated. As far as it can be determined from the fairly encursted vertebrae, specifically on the fourth cervical vertebra, the cotyle appears to have had a somewhat transversely expanded form, that is, a more ellispoidal one. The articular processes [“Gelenkfortsätze”] of Joh. Müller, or zygapophyses of Owen, arise laterally on each vertebra as oval expansions and are here, in the cervical series as in other sections of the vertebral column, so arranged that the articular surfaces of the postzygapophyses are directed downward and those of the prezygapophyses upward, so that the articular surface of each postzygapophysis lies atop that of the prezygapophysis of the next-most posterior vertebra and covers it in order to articulate with it. The articular surfaces are somewhat inclined, although differently in the various sections of the vertebral column, hence the name processus obliqui of Soemmering. In the cervical section the anterior surfaces are turned somewhat medially, the posterior ones correspondingly laterally. On our slab, as mentioned, the transverse processes, or processus transversi—diapophyses of Owen—appear laterally on the anterior end of the vertebrae as longitudinally oval, fairly well developed bulges that jut out somewhat obliquely, that is, as ventrally directed bony bulges if the animal is considered in its normal position.

Somewhat less well-developed on the fourth and fifth cervical vertebrae, they increase, on the following ones, in eben dem Masse more conspicuously in size than the ribs that attach to them, which is naturally the case as well in the dorsal vertebrae. For the attachment of the ribs, the distal end of each diapophysis possesses a simple, weakly expanded, ellipsoidal little capitulum for articulation with a complementarily weakly impressed articular surface on the somewhat thickened proximal end of the corresponding rib. This feature is also repeated on the dorsal vertebrae.

The neurapophyses and neural spines, that is, the upper vertebral arches and their spines, the processus spinosi, are not visible on account of the aforementioned inverted position of our animal.

As a result of the enormous pressure exerted by the surrounding masses that was once borne by the neck of the lizard, and the resultant tear between the third and fourth cervical vertebrae in the vertebral column, the other vertebrae were also dislocated at their joints, each one more or less shifted from its normal position; the entire cervical section of the column was strongly bent on itself and twisted to the left as far as back as the third sternal rib [“Brustrippe”]. Possibly a spasmodic contraction of the abundant neck musculature during the death throes at the violent end of the animal also contributed to effecting this so significant and conspicuous bending of the neck.
Dorsal vertebrae.

These forces appear also to have acted upon the dorsal segment of the vertebral column. Here as well, an abnormal, if also slight, bending of the same [the column] toward the right side of the animal may be assumed, for in the natural position it would extend more horizontally.

The number of dorsal vertebrae, as the rib-bearing but hypapophysis-free vertebrae can be seen, come to twenty. Lumbar vertebrae are lacking, and there follow immediately after the dorsal vertebrae two sacral vertebrae, a characteristic that appears in the Varanidae among others.

The dorsal vertebrae (do.) generally have the form of the cervical vertebbrae, as a mere glance at our slab suffices to show. Their size relationships show only slight differences. The length of their bodies, or centra of Owen, vary only a little from the measurement of 2 cm [“von dem Masse zu 2 cm”]. Longest are those in the middle of the dorsal section of the vertebral column: they grow somewhat shorter toward the sacrum and likewise toward the last cervical vertebra in gradual transition. The total length of the dorsal series comes to 38 cm.

The dorsal section of the vertebral column also shared in the injuries that our animal had suffered before and just after the time of its deposition and embedding from the pressure of the waters and of the mud supported within them [“des darin erhaltenen Schlammes”]. Furthermore, the actions in the quarry during the excavation of the petrefaction also contributed their own [injuries]. The left forelimb and the remains of the shoulder that attach to it proximally were displaced to the right from their natural position, and the right limb was likewise displaced laterally. The former thereby came to lie transversely on the vertebral column, whose respective part was covered, so that the vertebrae found there become [“werden”] hidden. As one is able to conclude from the dimensions of the preceeding and succeeding vertebrae, there are three vertebrae in this place, of which the anterior one, that is, the fourth dorsal vertebra, is only exposed anteriorly, the middle (or fifth) one not at all, and the following sixth dorsal vertebra only posteriorly. The other dorsal vertebrae are as a whole more or less well preserved and with their undersurfaces in view at the level of the stone slab.

The centrum of these vertebrae is thick and robust and has a relatively flat ventral surface (from anterior to posterior) with a median, channel-like depression that is somewhat shallower toward the ends of the vertebrae and whose raised margins, like the ridges mentioned on the cervical vertebrae, may have served for the attachment of muscles or fibrous ligaments. From right to left the surface is convex, and laterally softly rounded. The transverse processes are strong and longitudinally round and constitute uninterrupted [“continuierlich”] lateral, anterolaterally directed extensions of the vertebra, whose under-surface expands onto them. The zygapophyses are rather wide and similar in form to those of the cervical vertebrae. The attitude of their articular surfaces, however, appears to be less inclined. Their morphology is difficult to discern on our fossil and is best apprehended on the vertebrae of the posterior half of the dorsal section of the vertebral column, from about the 14th vertebra to the 20th. In this region the columna vertebralis shows a weak rotation about its axis toward the left side. Where no calcite crust adheres to the vertebra, nor any damage to the bony part has taken place during the excavation of the skeleton, ones sees to the right of the posterior end of the vertebra, anterior to the transverse process of the succeeding vertebra, roundish protuberances that correspond to the articulated pre- and postzygapophyses of the two abutting vertebrae.

Neurapophyses and neural spines—covered by the vertebral centra, as in the cervical vertebrae—are not visible.
Ribs.

To the transverse processes, as mentioned in the cervical series, are attached the ribs, or costae, or pleurophyses of Owen, which were maintined by connective tissue in a simple joint. Their proximal (upper), simple, undivided capitular end comprises the ellipsoidally depressed, nearly vertical articular concavity [“Gelenkpfanne”] for the reception of the correspondingly convex articular bulge on the transverse process. Ribs are already present on the cervical vertebrae, with the exception of the first three, but on our slab are unfortunately for the most part indiscernible because of the carbonate encrustation. Thus one sees on the right side of the fifth and sixth cervical vertebrae an adjacent piece of a rib, and the same on the seventh [vertebra] of the same side. Especially well preserved, however, is the right rib of the eights cervical vertebra. It is around 7 cm long, and 5 mm wide at the proximal end, and extends down along the ribs of the first and second dorsal vertebrae, where its distal end pushes in between them. The eighth left cervical rib, which reaches over to the left humerus, and also the seventh left cervical rib are recognizable, if somewhat encrusted. These latter ones already liken the dorsal ribs in form and approach these in length. In contrast, the first three pairs, so far as their encrusted condition permits conclusions, are short and flat, but robust, especially at their proximal ends. Posteriorly, however, they gradually increase in size.

The dorsal ribs, all twenty of which are clearly visible on the right side and in their relationship to the corresponding vertebra, also constitute thin but strong, latterally moderately curved struts of bone [“Knochenspangen”] whose proximal end, as in the cervical ribs, articulated in the indicated manner with the transverse process of the corresponding vertebra. Their lateral surface is convex and provided with small embossments for muscle attachment, the inner one rather smooth and depressed along its length like a channel; the upper edge is rounded off, [whereas] the lower one [is] narrower, somewhat angular toward the proximal end, more rounded again distally. Their length reaches a maximum from the 9th to 15th, where they come to about 10 cm; the 6th, 7th and 8th are each 9.5 cm long; posteriorly, namely from the 16th on, the 18th reaches 3.0 cm in length, the 19th 2.5 cm, and the 20th only 2 cm. Their width comes to a maximum of 5 mm at the proximal end and 2 mm at the distal end; it decreases posteriorly to 2 mm proximally and 1.5 mm distally. One can term the last four rib pairs dorsolumbar ribs like the vertebrae to which they are attached. On the left side the ribs are all more or less strongly pressed against the vertebral centra, whereby the evidence of their belongingness to the corresponding vertebra is sometimes made difficult. The last four ribs of the lefft side have been lost entirely. In that place, namely, the fracture of the stone slab runs close to the vertebral centra and coincides posterior to the second sacral vertebra with a transverse break that destroyed the caudal vertebrae.

The relationship of the dorsal ribs to the sternum, and so the construction of the rib cage and the differentiation into true ribs, which are connected to the sternum, and false ribs, whose distal ends, probably cartilaginous in life, no longer reach the sternum, can no longer be ascertained on our fossil. If one may be permitted to draw a conclusion about from the agreement in the number of cervical ribs and dorsal ribs, which occurs in most cases in the Lacertilia, then one could also suppose that there were five true ribs in our animal, an assumption, however, that can not be established with certainty.

On account of the aforementioned destruction of the shoulder girdle, there are also only very tiny and doubtful traces of the sternum present. Thus one sees, in the region of the right humerus, between it and the distal end of the first two dorsal ribs, a rhomboidal little plate of bone, which continues to the side, in particular to the left, as a thin little rod. It is possibly interpretable as remains of the episternum**, which forms the so-called T-shaped, anterior-most part of the sternum with said rod-like appendage. This appendage is closely appressed against a thin, longish, rather straight bone that is expandedat the ends and could be interpreted as the clavicle. Because the photograph of this detail was unable to represent this clearly, only mention of it will be made here in the text and its position on the original slab noted. For this reason as well there is on the outline-plate (II) neither a more definite circumscription nor a label. That a developed sternum was present is demonstrated by the multiple in-between pieces, or costae intermediae (co. i.), Pl. II, preserved between the dorsal ribs, in particular their vertebral pieces, on the slab, and the rather long sternal pieces, costae sternales (co. st.), Pl. II, which run out from them, angling anteriorly and toward the middle of the body, which certainly were for attaching to the lateral margin of a cartilaginous breast-bone. Such sternal pieces are found, among other places, on the left side of the vertebral column at the tenth dorsal vertebra, where lies in the proximity of the transverse process of the latter and extends anteriorly therefrom along the vertebral column until the middle of the body of the sixth dorsal vertebra. Such sternal rib pieces likewise lie on the right side of the body, e.g., near the transverse processes of the twelfth and eleventh dorsal vertebrae, where one sees them directed anteriorly, crossing the vertebral pieces of the ribs of these and many other vertebrae [“mit den Vertebralstücken der Rippen dieser und mehrerer anderer Dorsalwirbel sich kreuzend”]. Presumably, the position of the mesosternum on our slab, after the compression of the thorax and the displacement of the forelimbs, was finally to be found lateral to the right humerus, because one sees there a number of the aforementioned costae sternales, unfortunately irregularly arrayed, ending like an insertion [“nach Art einer Insertion enden”].
Sacral vertebrae.

The sacrum is composed of two vertebrae in our animal. They are similar to the dorsal vertebrae in form, except that their centra are somewhat shorter, specifically 1.6 cm long each, and their diapophyses more broadly apposed. Their preservational state is not as good, for they were affected by the fracture that also involved the anterior part of the caudal vertebral column immediately behind the sacrum. In an undisturbed position of the animal, the pelvic bones would have to have made an appearance and overlain the two sacral vertebrae. But the same are in greater part missing, so that the under-surface of the sacrum, turned upward at the level of the stone slab, are free. The calcitic encrustation, however, partly renders the upper [observed?] surface of the vertebrae unrecognizable.

On the centrum of the first sacral vertebra (sa1), see Pl. II, as on the dorsal vertebrae, the longitudinal groove can clearly be seen which, together with the vertebral centrum, appears slightly laterally shifted to the left. On the second sacral vertebra (sa2) this channel-like depression is scarcely recognizable any more, and the encrustation on it is greater. Yet one still clearly sees the connection of this second sacral vertebra with the first, a connection that must have been a much tighter, firm one, for the condyle is must less well developed than that of the antecedent ultimate dorsal vertebra, so there can only have been reduced intervertebral movement of the sacrum.

On the left side of the sacral vertebrae there runs the strong fracture of the main or larger part, whereby both the character of these lateral surfaces and that of the robust transverse process of the left side appear to be rendered unrecognizable.

At the posterior end of the second sacral vertebra one sees the angular, edgy location of the aforementioned fracture, which the vertebral column has suffered here. In this regard, a small part of the vertebral centrum was displaced, so that nothing more can be ascertained of the convexity of the condyle.
Caudal vertebrae.

At this point begins the long series of caudal vertebrae, which are distinguished by very well-developed transverse processes (p. tr.) as well as by strongly developed neuro- and haemapophyses (n. and h.) with thier processes, the neural and haemal spines (nsp. and hsp.). Unforunately it is precisely the beginning of this segment of the vertebral column, among all parts of the animal, that was most affected by destructive influences. The vertebral column is truncated just behind the second sacral vertebra, and only incomplete impressions and bony fragments, which moreover are covered in many cases by a calcite crust, permit one to assume, by means of comparison of the part and its counterparts and taking into consideration the dimensions determined for the succeeding remaining vertebral centra—each vertebra on average probably 14 mm in length—the presence of seven vertebrae (Pl. II, ca1 to ca7), counting up to the downwardly directed corner of the triangle that indicates the wholly missing slab.

In the breaking off of the caudal vertebral column, which occurred immediately at its beginning behind the sacrum, the total length of this section of the columna vertebralis was rotated toward the right side of the animal around its own axis, and thus the column [i.e., the set of articulated centra?] itself was somewhat displaced to the left. Thus, the haemal spines should have appeared upwardly directed, as they clearly do on the dorsal vertebrae and on the sacrum because of the inverted position of the animal, to make a rightwardly turning quarter-circle and were embedded in such a position on the entire caudal section at the level of the bedding plane. Accordingly, the neural spines of all caudal vertebrae are contrariwise sunk into this very plane on the side correposonding to the left half of the fossil. In addition to the parts, which are apparent on Pl. I and II by the fractures that developed during the excavation of the stone, three more fragments of the stony slab, Fig. 2, 3 and 4 on Pl. III, comprised the counterpart, just as we already mentioned in the discussion of the head, Fig. 1, Pl. III. They contain sections of the caudal vertebral column, indeed with nearly all the bone pieces that fit as positives to the negatives of the molds or impressions on the part. The state of preservation of two of these, Pl. III, Fig. 3 and 4, makes the determination of the correspondence between their features and those of said fragments of the part doubtlessly possible, which will [“soll”] be dealt with below. The third, Pl. III, Fig. 2, is somewhat more difficult to interpret. It contains three vertebral centra with parts of their neural and haemal spines, as well as their transverse processes, which apparently belong to the anterior-most part of the caudal vertebral column. One could hardly err in seeing them as remains of the third, fourth and fifth caudal vertebrae. The impressions of the remaining vertebral centra and their processes on the fragments of the part belong then to the first, second, and sixth caudal vertebrae. The length of this piece of the vertebral column comes to over 9.5 cm34 on the part, or on average 14 mm for each vertebra, which stands in agreement with the length of the remaining anterior vertebrae figured on Pl. III, Fig. 2, as well as with the length of 16 mm taken for the sacral vertebrae, in comparison with which they [the sacrals] gradually diminish in size.

Now, as mentioned earlier, a triangular piece of the part is missing on whose lower, vor- oder einspringendem corner, i.e., the one toward the junction of three of the remaining pieces of plate, a vertebra was preserved and has been lost together with neighboring parts of the preceeding and succeeding vertebrae.

On the two following pieces—the negatives on the part, the positives on two like overyling counterparts, Pl. III, Fig. 3 and 4—, which belong together and are separated by a break in the stone slab, there lie twelve caudal vertebrae, and furthermore a fragment of another, ca8, anterior (in reference to the animal) one, of which only something of the centrum and of the neural spine was preserved, and then also another, ca21, at the end of this section, of which only half is preserved. On the two counterparts, Fig. 3 and 4, the character of these vertebrae is clearly discernible. Robust neural spines (nsp.), in which the strong, basically [“am Grunde”] 7-mm-wide neurapophyses (n.) terminate and combined reach a height of over 1.5 cm; well-developed, flat, horizontal, gradually tapering transverse processes (p. tr.), here and there broken-off, up to 1 cm in length and around 5 mm in width, and furthermore two side-pieces each of a haemapophysis (h.), which, converging on one another distally, finally are united as one haemal spine (hsp.) each.

On the vertebral centra one sees very clearly, especially on some of them, that, as in other Lacertidae [“Lacertiden”], each haemapophysis articulates with the posterior end of the vertebra itself to which it pertains and not at the place at which any two vertebrae are joined, which latter view is represented by R. Owen35 and C. Gegenbaur.36

The second of the two said positive counterparts of this section of the vertebral column, Fig. 4, is the immediate extension of the one just mentioned, Fig. 3, of identical character and only separated from it by the fracture in the stone. By a small displacement, which clearly occurred after the decomposition of the soft parts (joint capsules and ligaments) that surrounded their articular ends, there arose in said sections of the vertebral column weak kinks that interrupted the continuity of its curvature, whereby the latter does not continue in a completely smooth curve toward the posterior bit of the tail but rather forms a blunt if significant angle. Now we arrive at the place where a piece of the stony slab was entirely lost, the triangular Zwickel terminating ventrally at its point, which we have already encountered in the descrioption of the eighth caudal vertebra. The distance between the two vertebral pieces that bound the lost portion of the caudal vertebral column anteriorly and posteriorly comes to 12 cm, a length measured along the curve of the bending of the tail. In this stretch there were presumably 11 vertebral places, at which count one arrives when one envisages the dimensional length of the vertebrae preceeding the missing piece (11.2 mm) and of those following it (9 mm) and thereby also considers somewhat the intervertebral joints.

The following and final piece of the caudal vertebral column is very well preserved, with the exception of the very last and smallest end-member, both on the part and, in part, on the corresponding counterpart, which, as we have seen above, contains the positive of the head, Pl. III, Fig. 1. The anterior part of the former shows the impression of 16 vertebrae, ca32 to ca47, with a length of 17.5 cm, preceeded and succeeded also by a half[-vertebra]. The counterpart contains the bony substance of its 15. The individual vertebrae, in respect of the length and height of their centra as well as the length of their neural and haemal spines, diminish only very gradually posteriorly. The following end-piece of the caudal vertebral column, in contrast, is like the section preceeding it. It is namely provided to us as a positive on the part, that is, as actual bony material, whereas the corresponding counterpart, which would bear the impression, is missing entirely. The length of this section is 21 cm, including an estimation of the non-preserved smallest elements of the tip of the tail. One can estimate the count of its vertebrae, the latter included, at about 50. One sees, already on the vertebrae preceeding the gap, that the transverse processes become progressively shorter and narrower. Thereafter they disappear entirely, and on the final piece they are no more than weak lateral bosses or are no longer discernible at all. An identical condition of the zygapophyses obtains, which shorten dramatically, so that at last the articular joint is limited to the vertebral centra. At the same time the anterior concave and posterior convex articular surfaces of the vertebral centra become progressively flatter; indeed, near the end of the tail they acquire at last a character reminiscent of the skeleton of fishes. Neural and haemal spines retain their form, but diminish constantly in respect of size unto the smallest element near the tail tip, which finally consists only of very delicate [“zarten”] little rods of bone.


Shoulders and forelimbs.

Even an only tolerably recognizeable or articulated shoulder girdle is no longer present on our slab. As the rib cage has suffered terrible destruction and the separation of its elements, so that only a few speculations about its construction can be expressed, so too are both shoulder girdles in so poor a preservational state that extremely little can be inferred from the slab about their form. This applies both to the right shoulder and, to an even greater degree, to the left one. Far to the side, over 6 cm from the vertebral column, approximately at the level of the fourth dorsal vertebra, one sees several smooth, uneven, undulating bony plates from the first [the right] girdle, which are partly encrusted and overlain by the capitulum of the humerus. This [the capitulum] is separated from its articular fossa and shoved up onto it next to the ends of ribs and fragments. The edges of the aforementioned bony plates are no longer recognizable; only the little bony plates lying behind the capitulum show a marginal embayment reminiscent of those of the raven-bone, or coracoid. The adjacent little bony plates would then be assigned to the scapula and its partly ossified margin, which continues further. Weak conjecture with regard to the clavicle and the episternum attached to it were previously expressed. On the left side, in the region between the fourth and sixth dorsal vertebrae, there exists only an agglomeration of flat, little bony plates, permeated by cracks or fragmented, under which there appears a somewhat clearer one [agglomeration?]. It [the agglomeration?] is bounded on the right by a half-moon–shaped curve and appears to derive from the scapula, on whose left side a portion appears to indicate the articular fossa for the shoulder joint [“an deren linker Seite ein Antheil davon die Gelenkpfanne für das Schultergelenk zu bezeichnen scheint”], while the posteriorly adjacent part would belong to the coracoid, which, as is known, helps construct the glenoid fossa in combination with the scapula. The upper or proximal articular head of the humerus would then be interpreted as still in situ, that is, lying in the articular fossa, also entirely consistent with its form and surroundings on our object. Posteriorly, triangular pieces of the stone are missing, together with any possible inclusions [i.e., bone], which were apparently lost during the recovery of the slab in the quarry.

Both forelimbs, especially on the left side, are fairly well preserved, each in its three segments, the upper arm, the forearm and the hand. The right one is bent at the elbow, the left stretched out straight. Both turn their anterior or ventral side [“ihre vordere oder ventrale Seite”] to the observer.

The upper arm bone, or humerus (hu.), is a robust, fairly straight long-bone about 4.5 cm in length, expanded at the ends, measuring 1.4 cm proximally, 1.3 cm distally, and nearly half as narrow in the middle, at 7 mm. Its head is oval, little distinct from the diaphysis, and shows lateral tuberosities that served for muscle attachment. The distal end, as clearly discerned on the left side, has a trochanter laterally for articulation with a correspondingly depressed articular fossa on the upper, proximal end of the radius, and a more protrusive one medially, the division of the elbow-joint, probably trochlear, for the proximal end of the ulna. A depressed place on the anterior or ventral side of the humerus, which is visible on the slab, divides these two articular parts, while the dorsal or posterior surface is sunk into the stone and is not visible.

The forearm, or antebrachium, is shorter than the upper arm, namely 3.5 cm long, and consists, as usual, of the radius (ra.) and the ulna (ul.), which are separated by a significant interossial space that distally reaches 8 mm. It is, in particular, only their proximal ends at the elbow that are united, whereas distally carpals are interposed between the ends of the radius and ulna. The radius is somewhat laterally curved, and the ulna extends fairly straight. The mechanism for the rotation of the radius to the ulna at the upper, proximal end of both, which doubtlessly obtained in the living animal, cannot be discerned on the stone. A patella ulnaris or the olecranon of the ulna is likewise not visible, structures that, on the extensor surface of the arm, now lie sunken into the stone slab.

The third section of the forelimb, the hand or manus, is composed of three further pieces: the carpus, the metacarpus, and the fingers or digits.

On the left hand the components of the carpus (cp.) are rather completely preserved, and more clearly in the original than they are reproduced in the photograph. One can see, best on the left side, the two rows of short, thick little bones, three in the proximal row: the radiale, for articualation with the distal end of the radius, and the ulnare, for articulation with the ulna. Between both the intermedium or centrale is interposed like a wedge, while the pisiform attaches to the ulna and simultaneously to the carpals laterally as a so-called sesamoid.

The second, distal row of carpals consists of five pieces; they are terms carpals 1 through 5. Carpal 1 borders the radius proximally, the centrale laterally, and the first metacarpal distally. Carpals 2 through 5 are arrayed one after the other in such a way that they articulate distally with the proximal ends of the neighboring metacarpals and proximally with carpal 1, the centrale and the ulnare.

The metacarpals (mcp.), likewise five in number, are short “long-bones” [“Röhrenknochen”] whose broader proximal ends have deepened articular surfaces for the reception of the carpals just named, while their thickened distal ends show heads [“Gelenkköpfe”], which serve for the articulation of proximal phalanges.

The fingers, or digiti manus, also constitute short “long-bones,” which are somewhat thinner in the middle than at the ends. They are of unequal length. The longest (fourth) finger measures 5.2 cm, the shortest (first) 2.7 cm. Because their bones lie in rows, one after the other, they have received the name phalanges. The first finger has 2, the second 3, the third 4, the fourth 5 and the fifth again 3 phalanges.37 Their proximal ends, broader relative to the diaphyses, have concave articular surfaces for the reception of the convex heads of the appertaining metacarpals; the distal end of the phalanx following the metacarpal, as with all further phalanges, is rounded off and provided with a groove for the articulation of the proximal end of the next member, which shows an eminence corresponding to the groove and neighboring depressions, which constitutes a so-called saddle-joint. A similar joint one also finds, for example, in man between the proximal end of the metatarsus of the thumb and the os multangulum majus that articulates with it. The last phalanx of each finger ends in a small, ventrally curved point, which was provided with a horny claw.


Pelvis and hind-limbs.

Previously, in the description of the sacrum, it was emphasized that the underside of the two sacral vertebrae, like that of the adjacent dorsal vertebrae, are entirely exposed, whereas in the normal condition of the skeleton, they would have been covered by the paired bones of the pelvis—united at a symphysis on the midline—and in particular by the os pubis anteriorly, by the os ischii posteriorly, and further laterally, more or less, by the ilium. The pelvic girdle, however, is not given to us but rather divided into its consituents; these themselves, however, were for the most part dispersed and lost, except for tiny remains. For only the ilia, or ossa ilii, of both sides are still present, if in an entirely different position. They appear as thin, irregularly elongate, triangular plates of bone whose thickened, normally anteroventrally directed base, as is generally known, participated in the building of the acetabulum with the two other pelvic bones, whereas the upper, thinned, somewhat sharp, posterodorsally directed end was connected laterally to the sacrum on its two strong, connected transverse processes, probably by synchrondroses in life. Both ilia lie to the side of the vertebral column, separated from their places of articulation: the right one—on the part—against and beneath the upper leg-bone and, further, on the proximal end of the fibula, with the tip directed posterolaterally; the left one, with its base abutting on the vertebral column, extends laterally and somewhat posteriorly over the proximal piece of the left femur, with the tip directed toward the proximal end of the fibula. The left ilium is located on the adjacent, accessory piece of the part that was separated by the longitudinal break at the vertebral column.

Nothing remains of the os pubis on our slab.

A trace of the os ischii may be indicated, though very dubiously, on the counterpart of the anterior-most piece of the caudal vertebral column, in particular an impression in the calcareous shale. This presents a short, irregular rectangle, its sides somewhat embayed in the middle, the corners on the other hand a little drawn forwards [“die Ecken aber ein wenig vorgezogen”]. It has in form a similarity to the outline of an ischium; only for the uncertainty of the identification did I abstain from indicating it in any way on the figures of the plates.

The hind-limbs themselves, like the forelimbs, consist of three sections each, namely, the upper leg-bone, or femur, the shank, or crus (tibia and fibula), and the foot, or pes. It is their flexor surfaces that are exposed at the level of the stony slab. Thus, the hollows of the knee face upward, the tibiae lateral, the fibulae medial.

The upper leg-bone, or femur (fe.), at 5.5 cm long, is the most robust long bone of the entire skeleton. It exceeds the humerus in length by 10 mm. On our (main and accessory) parts the osseous tissue is preserved in part, namely, somewhat less than half in each, and in particular, the proximal end of the right one and the distal end of the left. The remainder is only represented by more or less distinct impressions. The proximal, upper end of the right femur clearly shows a part of the spheroidal capitulum, then the neck and, under it, two of the trochanters as muscle processes [“als Muskelfortsätze”]; these are, namely, a large one laterally, the T. medius, and a smaller one medially, the T. anterior. Further ventrally, the body or middle piece [diaphysis], provided with longitudinal grooves and streak-like eminences [i.e., flutings], narrows somewhat and is then broken transversely, from which the impression of its posterior surface continues further backward over the fibula, on top of which the femur had been shoved. As a result of this condition, as well as the aforementioned apposition of the ilium, the interpretation of its relations is made rather difficult. On the left side, the proximal end of the femur is partly covered by the left ilium and partly encrusted; thereafter follows a part of the impression of its middle piece [diaphysis] and further still its distal end with the articular surface for the tibia and laterally for the fibula.

The shank has the same length as the forearm, 35 mm, but is 20 mm shorter than the femur and consists, as usual, of the lateral tibia and the medial fibula. Both are separated by an interosseal space, just like the bones of the forearm, so that their lower, distal ends do not connect and only the upper, proximal ends articulate with one another.

The shin-bone, or tibia (ti.), is conspicuously more robust than the fibula—twice as wide as it—and proximally it shows the two articular tuberosities of the knee and, between them, a depression; on the middle piece [diaphysis] are flutings, and distally there is a longish, transverse articular surface for the tibiotarsal or the astragalus.

The fibula (fi.) has almost the same length as the tibia. Its rounded, upper or proximal end, on both the right and left sides, is more or less displaced and removed from its connection with the femur and the tibia. Distally, it connects with the fibulotarsal (calcaneum). One of the small bones that, in lizards, are present in and around the knee joint is clearly visible on the left side between the two upper capitula of the crural bones; the knee-cap, or patella tibialis, appears to project beneath the ball-like articular surface at the femur.

The ankle bones, or ossa tarsi (ta.), are somewhat removed from their normal position by the pulling apart of the distal ends of the two leg bones. One recognizes, however, especially on the right side, the bone, shifted inward (medially) and therefore separated from the tibia and clinging to the fibula, that represents the first row of tarsals and according to the older view of Cuvier and Owen38, in the sense of human anatomy, was regarded as astragalus (tibiotarsal) and as calcaneus (fibulotarsal), tightly linked or united as one bone [fused]. According to the studies of Gegenbaur39, however, it arose from a fusion of four primary elements—the tibiale, intermedium and centrale on the one side, the fibulare (calcaneus) on the other—and was termed by him the calcaneo-astragalo-scaphoideum. One sees its connection with the fibula quite distinctly on the right foot. The connection was in any case, together with the tibia, a rather tight one, for the movement of the foot at the lower shank, like in all related animals, took place at the intertarsal joint.

The second row of tarsal bones on the left is clearly distinguished from the metatarsals, but on the right side they are shoved up somewhat at their proximal ends, such that here, individual little bones are removed from their position and connections. One sees, however, beginning medially, i.e. on the fibular side, a massive bone, the cuboid (digito-tarsal 4–5 tum Brühl), which follows on the third tarsal (ectocuneiform). Nothing can be discerned of the second tarsal (mesocuneiform) or the first. These may well already have fused with the proximal ends of metatarsals II and I, especially the first tarsal. A short, conspicuously wide and arched bone borders laterally on the cuboid and is viewed by Hoffmann40 as the fifth tarsal. Then the fifth toe would only have three phalanges, because the immediately following element would be considered as metatarsus. Otherwise this bone is given in comparative anatomy as metatarsal V, whereby four phalanges are attributed to the fifth toe.

The first four middle foot-bones, or ossa metatarsalia (mta.), are rather uniform in their shape and form. There are longish, thin “long-bones” [“Röhrenknochen”] whose proximal ends articulate with the tarsals. On the right side, as noted, on account of the resultant displacement [“wegen erfolgter Verschiebung”], these joints are not visible on our object. On the left side, in contrast, the same are more distinct, and one sees their proximal ends in connection with the articular surfaces of the tarsus; namely, metatarsals I and II articulate with the tibiotarsal (astragalus), and metatarsals II and III with tarsal 3, the latter of which also borders medially on the cuboid, and this cuboid is distally in connection with the fourth metatarsal and medially with the fifth. The last [metatarsal] is, as just mentioned, short and wide, with a medial bend that produces a protrusion [“Vorsprung”] and is consequently totally different from the other metatarsals I–IV. The length of the latter [four] is very unequal; they come to 13 mm for the first and shortest, 15 mm for the second, 18 mm for the third, and 19 mm for the fourth and longest.

The toes, or digiti pedis, are all of a concordant nature with one another and with the fingers; only their phalanges are somewhat longer. The fourth toe, including the metatarsals [sic], measures 63 mm; it is thus 11 mm longer than the fourth and longest finger, which measures only 52 mm. Because the fifth toe, if one does not share the view of Hoffmann, mentioned above, of a fibular appendage, has four phalanges—that is, one more than the fifth finger—the phalangeal formula for the foot, from the first through fifth toes, is expressed by the number series 2, 3, 4, 5, 4, counted from the lateral or outer side to the medial [as preserved], whereas for the fingers of the hand in succession the numbers 2, 3, 4, 5, 3 apply. The terminal phalanges are, like in the forelimbs, slightly ventrally curved, pointed, and somewhat laterally compressed and were in life provided with horny claws.

If one compares the two pairs of anterior limbs [sic] with one another, the forelimb appears somewhat shorter than the hind-limb. The length ratio of fore- to hind-limbs is 9:11. Even the hand is somewhat smaller than the foot; the ratio of these is 11:15. The upper arm is shorter than the upper leg, and the ratio of the lengths of the two is 9:11. The forearm and shank are equally long, 3.5 cm. Their appropriately structured [“zweckmässig gegliederten”] fingers and toes, five each in number, with their strong claws made them adequate [“befähigten sie”] for walking on land; likewise, their broad and long palms and soles were for movement in water and were also strongly supported by the long, tall, powerful, and probably laterally compressed tail, that is, an excellent rudder organ.

* *

*

With regard to the systematic position of our animal, there can be no doubt that it must be counted among the order of the scaled lizards, Lepidosauria [=Squamata of today], although nothing remains to us of its integument.



The procoelous nature of its vertebrae, the moveable quadrate attached only proximally by means of a suspensorium, the simple joint at which the unicapitate ribs articulate with the transverse processes of the vertebrae, and finally the absence of so-called gastralia, i.e., ossified strips of connective tissue, such as occur in other Sauria, for example in the Rhynchocephalia, from which they [the “Lepidosauria,” i.e., Squamata] probably evolved and which they liken in the external appearance of the body and in the structure of the interior.

With regard to the suborder into which it should be placed, the presence of limbs rules out the limbless [“fusslosen”] Ophidia; it further distances it [the animal] also from the Pythonomorpha, whose limbs—flippers [“Ruderfüsse”] especially adapted for swimming, both anteriorly and posteriorly (less so here) formed like oars, exquisitely shortened, broad and flattened—are composed of flat elements never capable of rotation and lying throughout in one plane (one surface). Our Lesina fossil has five-toed, claw-bearing walking-feet [“Gehfüsse”].

The fossil from Lesina does indeed agree in the important and singular character of the remarkable dentition with the Pythonomorpha, although these also display a toothed pterygoid; only the last-named, extinct, mighty sea-monsters still differ most considerably in their extraordinarily elongate body-form and an enormously long body cavity surrounded by many short, strongly curved ribs, whereas in our animal this cavity is not conspicuously long and is surrounded by numerous but not long ribs. The Pythonomorpha also usually have, like the snakes, special processes for the articular connection of the vertebrae, the zygosphenes and zygantra,41 which the Lacertilia (exc. Iguanidis [except Iguanidae?]) lack.

As examples of the noted great length of the body, just Mosasaurus at 8 m (25 ft), Clidastes at 11 m, Tylosaurus dispelor Cope at well over 30 m (100 ft) length may be named; most impressive dimensions, which other species of this suborder also have, and from which our fossil, in constrast, at only 1.4 m length, strongly departs. Moreover, the sacrum of the Pythonomorpha consists of only one vertebra, that of our animal of two; the Pythonomorpha lack the columella [epipterygoid], which is present here.42

The suborder Dolichosauria, or long-necked lizards, which possess a greater number of cervical vertebrae, always at least over nine [“stets mindestens über neun”], can likewise not be considered for our lizard, for it shows only eight cervical vertebrae.

That leaves, therefore, only the suborder Lacertilia into which we can place the fossil from Lesina. The body shape, not stretched in any conspicuous way and of a moderate length that also occurs in living forms, the extremely long tail, the rod-like columella, the limbs constructed as obvious walking-feet, with five claw-bearing toes, all amply justify this placement.43

But of all Lacertilia, our animal is closest to the monitor lizards, or Varanidae, in its form and body-shape, insofar as the skeleton permits one to determine them, and in particular in the elongate head, the smooth anterior frontal and neighboring posterior parietal regions, the rounded orbits, the azygous parietal, in the supratemporal fossa that extends up onto the dorsal surface of the skull, in the nearly quadrilateral occiput and the long parotic processes that extend from it, in the lack of lumbar vertebrae, and finally in the suspensorium for the quadrate.

Because of the construction of the skeleton, one would be nearly tempted to assign it [our animal] immediately to this speciose [“formenreich”] lineage, if not for the nature of its teeth, which are attached to supporting bases and wholly resemble those of the Pythonomorpha; their nature namely excludes our animal from the genus Varanus and all other Lacertilia. Our animal, on account of this remarkable character, is so very special and peculiar among the latter [Lacertilia], that it must be seen as its own genus, if not as representative of its own family.

The characters presented in the previous description render this fossil superior, preeminent among all those extinct animal forms thus far excavated on the island of Lesina, and in particular among the Reptilia, in respect of its size and beauty as well as its state of preservation and finally the special connection to related groups in the zoological system. It thus deserves, by all rights and above all others, the term Lesina-Saurian, xατ᾿ ξοχήν, in the true sense. But the expression “lesinensis” is now already multiply in use for fishes, ferns, and the like and was assigned by me myself as a species name in 1873 to Hydrosaurus (description loc. cit.), so another appellation might more appropriately be coined, except that something like “Lesinosaurus,” bilingually composed, would not fully adhere to the more stringent linguistic requirements of word-construction. The Italian name for the island, however, derives from a comparison that is supposed to have been made between an awl or Pfriem, which tool in Italian is called “lesina,” and the island’s peculiar, long, stretched-out shape. The Greeks named this tool “τ πήτιον,” which would give Opetiosaurus for the name of the new genus and which I now take the liberty of proposing, so cleaving to the generic name a commemoration of the splendid southern island with its precious, exquisite paleontological discoveries. Concerning the name of the species, the obvious course of action is to preserve in it [the name] our grateful remembrance of the active and highly honored student of varied fields of natural science, Mr. Gregor Bucchich of Lesina, to whose most laudably known eagerness and lively interest in the extraction and preservation of the “mementos of creation” [“Denkmünzen der Schöpfung”] we owe, beside many others, also this marvelous petrfaction, Opetiosaurus Bucchichi, one of the most precious treasures of the splendid collections of the Royal Imperial Geological Institute.

Although the special peculiarity of Opetiosaurus—in its remarkable dentition—distinguishes it in a salient way from all Varanidae and strongly differentiates it from all fossil and Recent Lacertilia known thus far, so that an identification with other Sauria previously found in the Chalk formation, be they in the plattenkalke of Lesina or the somewhat older carbonaceous shales of Komen (Barremian) and so forth, cannot be considered, yet a short differentiation with regard to other characters of these already known fossil saurians may be given.



Mesoleptos Zendrini Cornalia, 1851 from Komen, in the colleciton of the Museo civico di Storia naturale in Milan, belongs among the older discoveries. It is characterized by its posteriorly conspicuously narrowed vertebrae, which attribute also found expression in its name. Opetiosaurus in no way shows the same. A similar petrefaction from Lesina, which Kramberger44 mentions, is found in the collection of the instructor-widow Mrs. Antonia Novak in Eso Grande at [“bei”] Zara.

Acteosaurus Tomasinii Herm[ann] von Meyer, 1860, from the same locality, in the Museo civico in Trieste, is a dolichosaur with 27 dorsal vertebrae, nearly exactly equally long ribs, and furthermore with extremely shortened forelimbs, whereby the ratio of the length of the humerus to that of the femur is 1:2, and that of the middle to the proximal sections of the forelimb is 5:7, and 4:7 in the hind-limb, whereas in Opetiosaurus the ratio of the humerus to the femur is 9:11 and those of the aforementioned sections 7:9 and 7:11.

Hydrosaurus lesinensis Kornhuber, 1873, in the collections of the Royal Imperial Geological Institute, has 30 dorsal vertebrae and much more weakly developed forelimbs and a smaller head than Opetiosaurus.

Adriosaurus Suessi Seley [sic, for “Seeley”], 1881, from Komen, in the geological collection of the University of Vienna, is similar to the latter two but is a much smaller animal, although full-grown, and is distinguished from Opetiosaurus by short, compact [“gedrungene”] vertebrae that are convex length-wise [“der Länge nach convexe”] and by very robust ribs that are rather alike.

Carsosaurus Marchesettii Kornhumber, 1893, from Komen, in the Museo civico of Trieste, has nearly equally developed upper arm- and leg-bones, with a ratio of 16:17, whereas the forarm and shank have the same length, and the ratio of forearm to upper arm is 5:8.

There is now only Aigialosaurus dalmaticus of Prof. Dr. Carl Kramberger-Gorjanović [sic] in Agram to consider. This animal likewise derives from the fish- and reptile-producing plattenkalke of the island of Lesina, was discovered by a peasant [“Bauer”] of Vrbanj, and came into the possession of the now-deceased Mr. J. Novak, teacher in Zara. Prof. Kramberger-Gorjanović [sic] gave a careful description of it with illustrations.45 Aigialosaurus has some similarities to our new fossil; however, its head is much more tapered anteriorly (is sharply wedge-shaped), the parietal is broader in the middle, the mandible much lower [“niedriger”], the anteriorly displaced quadrate thinner and provided dorsally with a posteriorly directed, triangular process, only seven cervical vertebrae (whereas the dorsal vertebral count agrees with that of our animal, namely, twenty), the ribs are significantly shorter and the caudal vertebrae lower [“niedriger”]. Nothing appears to have been preserved of the teeth. At least, there is no mention of them in Kramberger. In case one should like so to refer to a steep, cone-shaped structure with a flat (broken-off?) tip on the dentary of the right mandible, then one could attribute to Aigialosaurus a very different dentition than in our new fossil, just like in other Lacertilia. It is not at all easy, without examination of the original slab, to form a sound judgment of one’s own from observation of the lithographic figure alone. Thus, one could be disposed to view the structures joined to the cervical vertebrae of Prof. Kramberger’s Aigialosaurus and which he identified as hypapophyses instead as cervical ribs, because the same arise from the anterior end of the vertebral centra at the transverse processes, which lie next to the prezygapophyses, exactly as is apparent in the cervical ribs of the fifth through seventh cervical vertebrae from the labels on the same plate, whereas the hypapophyses of the Lacertidae, Scincoidea and so forth, except for the Agamidae,46 and furthermore those of the Pythonomorpha, are always applied beneath the articular condyle, and thus posteriorly on the vertebra.47 Whatever the case may be, the hypapophyses of our new petrefaction have a form and position completely different from that of Aigialosaurus, as described earlier, and occur until the eighth cervical vertebra.

According to the reconstruction of Kramberger, his Aigialosaurus, in my opinion, could belong with no little probability to the family Varanidae D[úmeril] et B[ibron], where, of course, occur many fossil annd Recent forms conforming to “Sauri longo, acuto capite et aequis fere extremitatibus” and in which the emphasized diagnostic characters [“unterscheidend hervorgehobenen Charaktere”: sic?] of Kramberger’s group Ophiosauria48 are fully met. Kramberger49 namely finds these differences, which at the same time are supposed to reflect the transition of Aigialosaurus to the Pythonomorpha, to be: “1. in the greatly stretched body, 2. in the clearly distinct tendency toward reduction of the limbs, 3. in the presence of hypapophyses and 4. in the nature of the quadrate.” To begin with, as far as the stretching of the body is concerned, the length of Aigialosaurus, at 1.34 m—that is, a body-length that, in Varanidae, is both common among and exceeded (Varanus arenarius 1 m, V. niloticus 1.5 to 1.9 m, V. bivittatus 1.5 m, V. albogularis [sic] 1.7 m)—has no similarity to the enormous body lengths of 10 to 30 m in the Pythonomorpha50 and so cannot support any phylogenetic relationship of Aigialosaurus to these latter. In the limbs of both, however, there exists such a great contrast between the clearly distinct, claw-bearing walking-feet of Aigialosaurus and the unclawed flippers (“fins” of Cope51) of the Pythonomorpha, in which, moreover, the forelimbs generally exceed the hind-limbs, the reverse of the condition in Varanidae, so that this character also breaks down, particularly as the limbs show no apparent tendency toward reduction in Aigialosaurus either but rather have the same ratios as in the majority of monitor lizards.

Regarding further the presence of the hypapophyses on the cervical vertebrae, this constitutes no similarity with the order Pythonomorpha, for these [hypapophyses] do not represent a diagnostic character of the latter but rather also occur in a similar way in the Lacertilia. If these appear more conspicuous in the genus Clidastes, where Cope52 has figured them, this is probably explained by the significant size of this prehistorical American sea-monster. These hypapophyses appear so constantly in the Lacertilia that Calori53 even bases on them the distinction between cervical vertebrae, which possess them, and dorsal vertebrae, which lack them. C. K. Hoffmann54 also gives these as a general character of the cervical vertebrae in Sauria. I myself55 already described them in the year 1873, and not merely, as Kramberger loc. cit. p. 18 (91) and 30 (103) says, on the second, but also on the other cervical vertebrae of Hydrosaurus lesinensis. They are really true hypapophyses and not, like Kramberger states, something like the parapophyses of Owen.56 For the latter appear laterally on the centrum, and paired; they correspond to the little rib capitula or the lower transverse process (Joh. Müller), which is the anterior root of the processus transversus (Soemering).57 The hypapophyses, in constrast, arise, like in our saurian, on the midline beneath the vertebral centrum and are unpaired. Fr. Siebenrock58 as well has expressed his view on the hypapophyses in the same sense.

Lastly, I am not in a position to add to the ascribed diagnostic value of the “entirely different form,” as Kramberger stated, of the quadrate in Aigialosaurus, for the quadrate takes on the most various, often recurring [“wiederkehrend”] forms in the different suborders, families, genera and species of the Lepidosauria. Thus, the form of quadrate in our new Lesina fossil, as already described in the observations on the head, as well as that of Aigialosaurus probably has a similarity with the bone of the same name in the Pythonomorpha, which Cope59 figures from no fewer than 17 species in various positions, of which v. Zittel, after Owen and Cope, provides outstanding illustrations in Fig. 546 and 547 on p. 616 of his Palaeozoologie III and Grundzüge der Palaeontologie, Munich and Lepzig 1895, p. 645. But v. Zittel also noted, in the same place, that this quadrate “is most reminiscent in its form to that of the lizards and varies enormously in the various genera.” Indeed, we are reminded of it in Monitor niloticus, where it possesses an even more prismatic form, by the shallow excavation on its lateral surface (creusé en demi canal Cuvier60). In the genus Lacerta it was fittingly compared with an outer ear.61 Lacerta agilis has a curved, dorsally rounded, D-shaped quadrate,62 and Lacerta viridis has a completely similar one.63 One cannot therefore recognize, in this character, any special phylogenetic relationship of the Lesina saurians to the Pythonomorpha, particularly as Cope himself, in the diagnostic characters on which he based this new order in the subclass Streptostylica,64 made no mention of it.

Thus, the erection of a monotypic family “Aigialosauridae” cannot be deemed advisable for the reasons stated. A similarity of the Krambergerian saurian with the Pythonomorpha appears to me to lie in the formation of the peculiarly tapering, wedge-shaped head, which is reminiscent of Cope’s65 representation. This condition, in addition to the characters listed above, may well, in complete agreement in features with the family Varanidae, justify the adoption of a new genus “Aigialosaurus.”

The foregoing, elaborated argument will have provided the proof that the saurians discovered thus far in the chalk formation of Istria and Dalmatia generally bear all the varanid characters, apart from special features, and that even Kramberger’s Aigialosaurus as well as the new Opetiosaurus must be counted among them. If a transitional group between the Varanidae and the Pythonomorpha should be adopted, then such would best be represented by the new genus Opetiosaurus, for this animal, in its special, distinctive dentition among all other varanid-characters, embodies a completely natural, closer phylogenetic connection to the Pythonomorpha.

Finally, I cannot refrain from addressing [“begrüssen”] again most graciously those monsieurs who underwrote my work with their kind support in so many ways. Above all I am indebted to the Director of the Royal Imperial Geological Institute, my highly honored friend of many years, Mr. Hofrath Dr. G. Stache, for the allocation of the object; to Mr. Bergrath Friedr. Teller, for much good advice and for his friendly agency in the execution of the lithography in Plates II and III; and to Mr. Librarian Dr. Anton Matosch for multiple, kind Mühewaltung; further to the royal and imperial Director of the k. k. naturhistorischen Hofmuseum Mr. Hofrath Dr. Franz Steindachner and the royal and imperial Curator of the same, Mr. Friedr. Siebenrock for the allowance of literature and of comparative material of skeletal parts; finally to my excellent, erstwhile pupil, dear friend and colleague, Mr. Hofrath Prof. Dr. Franz Toula, for the work-room most kindly offered to me in the offices [“Örtlichkeiten”] of the Lehrkanzel for Mineralogy and Geology at Wiener k. k. Polytechnic, surrounded by the rich book treasures of the same, which made it possible and much easier for me to execute, in my dermalen and intermittent stays in Vienna, the undertaken task. I fulfill a dear and pleasant duty in allowing myself, in conclusion, to give expression to my feelings of most heart-felt and sincere gratitude to the honored sirs named above.


—————————

Comprehensive summary

of

Admeasurements of the more important parts of the skeleton of Opetiosaurus Bucchichi and ratios of some of the same



———

Meters


Length of the head 0.150

Width “ “ “ at maximal separation of the two jugals 0.060

“ “ “ “ at the posterior end (separation of the ends of the mandibles) 0.075

Length of the parietal 0.025

Width “ “ “ between the proximal ends of the two supratemporal processes 0.012

Smallest width of the parietal between the two concave lateral margins 0.007

Greatest “ “ “ “ (and frontals) at the frontoparietal suture 0.030

Length of the frontal on the counterpart, circa 0.040

“ “ “ “ on the negative of the part from nasal to frontoparietal suture 0.050

Width of the pair of frontals between the two prefrontals in the middle 0.009

“ “ “ “ “ “ “ the inner orbital margins 0.012

Length of the prefrontal 0.027

Greatest width of the prefrontal 0.008

Length of the postorbitofrontal (greatest, with the processes) 0.025

Width “ “ “ (greatest, with the processes) 0.015

Length of the orbit 0.035

Width “ “ “ 0.024

Length of the supraoccipital 0.005

“ “ “ quadrate, so far as it appears over the articular 0.022

Greatest width of the quadrate 0.017

Width of the quadrate at the position of the articular 0.012

Length of the suspensorium of the quadrate (su. te., sq., pr. parot.) 0.035

Distance of the upper (proximal) margin of the quadrate from the median groove

of the parietal 0.020

Thickness of the maxilla at the broken place (on the counterpart, Pl. III) 0.006

Width of the skull at this fracture 0.035

Length of the impression of the palate corresponding to the lost piece of the snout 0.045

Distance of the tip of the snout from the anterior end of the supraorbital [palpebral?] 0.055

“ “ “ “ “ “ “ from the frontoparietal suture 0.093

“ “ “ “ “ “ “ from the posterior end of the parietal 0.120

Length of the maxilla (on the negative) from the tip of the snout to the prefrontal 0.050

“ “ “ lower jaw [mandible] 0.150

Height of the lower jaw from the tip of the coronoid process straight down to the inferior

margin of the angular 0.028

Length of the dentary from the tip to the middle suture in the mandibular ramus 0.082

“ “ “ opercular from the tip to the middle suture in the mandibular ramus 0.083

Height of the coronoid from tip to base 0.013

Greatest length of the coronoid near its base 0.024

Height of the mandibular ramus at the fracture in front of the quadrate 0.007

Length of the mandibular ramus, so far as bone is present 0.031

“ “ “ “ “ (impression) from the fracture to the middle suture 0.036

Height of the angular (maximal) on the negative of the main slab 0.008

“ “ “ surangular (maximal) on the negative of the main slab 0.005

“ “ “ anterior half of the Md (d + op) from the middle, angled suture 0.018

“ “ “ “ “ “ “ “ “ in the middle of this half 0.008

“ “ “ “ “ “ “ “ “ anteriorly near the tip 0.003

Distance between the two posterior ends of the mandibular rami 0.075

Measurements of different vertebrae, their processes and ribs may be found in the text.

———

Number of cervical vertebrae on the counterpart, attached to the occiput 3



“ “ “ “ “ “ main slab, very distorted 5

“ “ dorsal vertebrae 20

“ “ lumbar vertebrae 0

“ “ sacral vertebrae 2

“ “ all vertebrae of the torso 30

“ “ “ caudal vertebrae, the anterior-most on the main slab 7

“ “ “ caudal vertebrae, the following ones, that is, the 8th, 9th and

10th, on the smaller slab comprehensively 3

Number of caudal vertebrae on the following two broken pieces of the slab, to

which two counterparts, Pl. III, Fig. 3 and 4, also correspond 10

“ “ all caudal vertebrae, that probably were present in the cleft but now

are missing entirely 11

“ “ all posterior caudal vertebrae in the caudal piece of the vertebral

column stretched out straight on the main slab, the anterior negative

part (impression of the counterpart), 1/2 then 16 then 1/2 again 17

“ “ all posterior-most caudal vertebrae, present as positives (the osseous

tissue) on the main slab, with the small, last, distal-most element of the

tail tip, which can only be counted approximately, circa 50

“ “ all caudal vertebrae, therefore, 98, or rounded to 100

“ “ all vertebrae combined 130


Length of the cervical section of the vertebral column (1st through 3rd vertebrae

0.020, 4th through 8th vertebrae 0.085) 0.105

“ “ “ dorsal section of the vertebral column 0.380

“ “ “ sacrum 0.032

“ “ “ torso, therefore 0.517

“ “ “ tail, anterior piece through the lost part of the stony slab (the cleft) 0.260

“ presumed of this cleft itself, along the direction of the curvature of the tail 0.120

“ “ “ tail section on the main part, anterior part as an impression (negative)

on this slab 0.175 [sum:]

“ “ “ tail section, posterior (end-)part as a positive (osseous tissue) 0.21 0.385

“ “ “ whole tail 0.765

“ “ “ whole animal skeleton (the head, at 0.150, included) 1.432

Length of the upper arm bone (humerus) 0.045

Width “ “ upper arm bone at the proximal end 0.014

“ “ “ “ “ “ in the middle 0.007

“ “ “ “ “ “ at the distal end 0.013

Length of the forearm (antebrachium) 0.035

Width of the ulna at the proximal end 0.0075, of the radius there 0.005

“ “ “ “ in the middle ...........0.004, “ “ “ “ 0.003

“ “ “ “ at the distal end ......0.006, “ “ “ “ 0.004

“ of the interosseal space as great as 0.008

Ratio of the length of the forearm to the upper arm 7:9

Length of the carpus, width unclear 0.005

Longest finger (the fourth) 0.052

Shortest finger (the first) 0.027

Length of the unguals, from 0.005–0.007

“ “ “ hand 0.055

“ “ “ forelimb 0.135

———
Length of the ilium 0.025

Width “ “ “ , proximally 0.008, distally 0.003

Length of the upper leg-bone (femur) 0.055

Width “ “ “ “ “ at the proximal end 0.013

“ “ “ “ “ “ in the middle 0.008

“ “ “ “ “ “ at the distal end 0.013

Length of the shank (crus) 0.035

Width of the tibia at the proximal end 0.09, of the fibula there 0.005

“ “ “ “ in the middle...........0.006, “ “ “ “ 0.003

“ “ “ “ at the distal end......0.008, “ “ “ “ 0.007

Interosseal space between the two, as wide as 0.008

Ratio of the length of the shank to the upper leg 7:11

“ “ “ “ “ “ upper arm to the upper leg 9:11

“ “ “ “ “ “ forearm to the shank 1:1

“ “ “ “ “ “ proximal and middle sections of the forearm to the length

of the proximal together with the middle sections of the hind-limb 8:9

Length of the tarsus 0.011, its width 0.015

Longest toe (the fourth), without the metatarsal 0.044, with the metatarsal 0.063

Shortest toe (the first), “ “ “ 0.017, “ “ “ 0.034

Length of the unguals, from 0.006–0.008

“ “ “ foot 0.075

“ “ “ hind-limb 0.165

Ratio of the length of the forelimb to the hindlimb 9:11

“ “ “ “ “ “ hand to the foot 11:15


—————————
Plate I.

A. Kornhuber: Opetiosaurus Bucchichi, a new fossil lizard from the lower chalk of Lesina in Dalmatia

———

Total view of the pieces of the part of the rock with the remains of the animal or their impressions, arranged according to their mutual relations and embedded in plaster of Paris, reproduced as a photograph at a ratio of nearly 9:10 of the natural size.


Plate I.

The fossil figured here was discovered, according to the exact statements of Dr. U. Söhle (Jahrb[uch] d[er] k[aiserlich-]k[öniglichen] geol[ogischen] R[eichs]a[nstalt] 1900, Vol. 50, Part 1, p. 43), who in the year 1899 devoted some time to geological studies on Lesina, in August of the indicated year and in particular in the pit belonging to Marino Vidos, 1.5 km NW from Mte. Hum near Verbosca. This stone quarry is, according to Söhle, about 10 m high and long and 6 m wide, and the layers of the plattenkalk dip 20–30º to the north.

The object, upon which the the photograph on Pl. I is based, consists of an 8- to 10-mm-thick calcareous shale plate, which, during excavation in the pit, was broken up into a larger main plate of 42 cm width and 33 cm height (in relation to the figure), that is, the most significant part of the picture, and into three smaller accessory plates. The main plate contains the head in the upper left, mostly as an impression (negative) of its upper surface, namely the skull roof, then the anterior part and the posterior end of the left mandibular ramus and the right mandiblewith the extremely remarkable, characteristic dentition. Beneath it also lies the posterior part of the tail (34 cm long in the figure), of which the anterior section (16 cm in length in the picture) is preserved as a negative and the remainder, up to the end of the tail, as a positive—that is, retaining the osseous tissue. Further down there follows the larger part of the torso. The animal being inverted, this appears with the under or abdominal surface of the 5 last cervical, 20 dorsal and 2 sacral vertebrae turned toward the observer. The cervical section of the vertebral column is strongly bent the left and posteriorly, the anterior dorsal vertebrae show a weak bending to the right, the following ones a moderate curve to the left. Of the caudal vertebrae, the 1st through 7th are only partly preserved, suggested on the lower left corner of this piece of the slab. Well preserved on the latter are the right forelimb, which, bent, lies near the tip of the tail, then the left forelimb, partly displaced onto the vertebral column and stretched out straight, and finally the right hind-limb with the especially beautifully formed bones of the foot. On a small, nearly equally schenkelig, triangular stone platelet, which adjoins the main plate diagonally and on the front-most part, is the left, hind-limb up to the tarsus and remnants of the caudal piece named above. Finally, in the left corner of the picture, two fragments of the plate are present, separated on the right by a fracture, which simultaneously forms the base of the aforementioned triangle and continues further posteriorly. They [the fragments] contain the left foot and the caudal vertebrae from the 8th to the 21st. The fracture, along which these two fragments abut, runs through the body of the 15th caudal vertebra. There follows a not insignificant cleft that arose from the loss of a piece of the slab, on which 11 caudal vertebrae were present and followed by the long, relatively straight piece of the tail found on the main slab.

The annotated Plate II provides more information on different particularities of the parts mentioned here.

The photograph of the stone slabs with the petrifaction—reassembled, set in plaster of Paris, and surrounded by a frame—was prepared with a ratio to natural size of 54:61. The size of the figure, therefore, is somewhat less than nine tenths of the original; that is, it is somewhat over one tenth reduced in the picture. Because the photograph and the schematic outline-drawing on Plate II, with the labels and so forth, are executed at precisely the same scale, an allowance should generally, and particularly in comparison with the figures on the lithographic Plate III, be made. The figures on this latter plate are namely produced at natural size and so in a relation of 1/1 with the stone.

My highly honored friend and colleague, Mr. Hofrath Prof. Dr. Jos[eph] Maria Eder, always happily prepared when it comes to advancing scientific goals, had the special kindness to have the photography of the stone slab executed in the Royal Imperial Graphical Teaching and Experimental Institut [“k. k. graphischen Lehr- und Versuchsanstalt”] in Vienna, which is under his direction. The taking of the photograph took place on the 5th of October 1900 in the most careful manner by monsieurs August Albert, k.k. Professor, and Anton Massak, Werkmeister, both active [“thätig”] in reproductive photography. The gelatin-plates were also prepared in this institute based on the negative of the very successful photography, and from them, test-photographs. The reproduction of the photographs then took place in the press of Max Jaffé in Vienna. It is an honor for me to express here the warmest, sincere gratitude of the k. k. Geological Institute and of myself to the monsieurs of that institute.


—————————
Plate II.

A. Kornhuber: Opetiosaurus Bucchichi, a new fossil lizard from the lower chalk of Lesina in Dalmatia

———

Outline of the skeleton with labeling of its individual parts, or outline-drawing.


Plate II.

In the representation of the anatomical units of the skeleton of the fossil, the basic outlines were first reproduced from the photograph. However, it was from the smaller plates appended as a lithographic figure on Pl. III, which constitute the counterpart to portions of the larger part, that all that appeared useful for the more exact presentation of the bony framework was incorporated and drawn in the appropriate place in the outline-plate. Thus, many skull bones show only weak impressions in the photograph, or are partly encrusted by calcite and therefore difficult to recognize, whereas the counterpart, Pl. III, contains the positive of the skull, that is, the substance of the bone, and also reproduces each negative impression that appears more or less clearly on the part. The same also applies to the portions of the tail piece of the vertebral column, which extend from the 1st to 7th, then from the 8th to 14th and from the 15th to 21st caudal vertebrae. Here as well, the picture is made much clearer by a combination of drawings from the part and counterparts. The elements of the skeleton are designated with the first letters of their Latin names, as used in the text; the numbers indicate the succession of elements of the same name. They are provided in order of anatomical succession. In a footnote on page 15 of the contribution [original pagination; here, endnote 37], attention was previously called to a mistake in the labeling of Plate II, where on the fifth finger of the right hand an extra phalanx—four instead of three—was represented.


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