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Dragons of the Air

<h2>CHAPTER XIV ORNITHOSAURS FROM THE MIDDLE SECONDARY ROCKS</h2> <h4>RHAMPHOCEPHALUS</h4> The Stonesfield Slate in England, which corresponds in age with the lower part of the Great or Bath Oolite, yields many evidences of terrestrial life—land plants, insects, and mammals—preserved in a marine deposit. A number of isolated bones have been found of Pterodactyles, some of them indicating animals of considerable size and strength. The nature of the limestone was unfavourable to the preservation of soft wing membranes, or even to the bones remaining in natural association. Very little is known of the head of Rhamphocephalus. One imperfect specimen shows a long temporal region which is wide, and a very narrow interspace between the orbits; with a long face, indicated by the extension of narrow nasal bones. The lower jaw has an edentulous beak or spear in front, which is compressed from side to side in the manner of the Liassic forms, but turned upward slightly, as in Dorygnathus or Campylognathus. Behind this extremity are sharp, tall teeth, few in number, which somewhat diminish in size as they extend backward, and do not suddenly change [Pg 154] to smaller series, as in the Lias genera. A few small vertebræ have been found, indicating the neck and back. The sacrum consists of five vertebræ. One small example has a length of only an inch. It is a little narrower behind than in front, and would be consistent with the animal having had a long tail, which I believe to have been present, although I have not seen any caudal vertebræ. The early ribs are like the early ribs of a Crocodile or Bird in the well-marked double articulation. The later ribs appear to have but one head. V-shaped abdominal ribs are preserved. Much of the animal is unknown. The coracoid seems to have been directed forward, and, as in a bird, it is 2½ inches long. The humerus is 3½ inches long, and the fore-arm measured 6 inches, so that it was relatively longer than in Dimorphodon. The metacarpus is 1¾ inches long. The wing finger was exceptionally long and strong. Professor Huxley gave its length at 29 inches. My own studies lead to the conclusion that the first finger bone of the wing was the shorter, and that although they did not differ greatly in length, the second was probably the longest, as in Campylognathus. Professor Huxley makes the second and third phalanges 7¾ inches long, and the first only about 3/8 inch shorter, while the fourth phalange is 6½ inches. These measurements are based upon some specimens in the Oxford University Museum. There is only one first phalange which has a length of 7¾ inches. The others are between 5 and 6 inches, or but little exceed 4 inches; so that as all the fourth phalanges which are known have a length of 6½ inches, it is possible that the normal length of the first phalange in the larger species was 5½ inches. The largest [Pg 155] of the phalanges which may be classed as second or third is 8½ inches, and that, I suppose, may have been associated with the 7¾ inches first phalange. But the other bones which could have had this position all measure 5½ and 7¾ inches. The three species indicated by finger bones may have had the measurements:— <div class='center'> <table border="0" cellpadding="2" cellspacing="0" summary=""> <tr><th align='center' colspan='4'>Phalanges of the wing finger</th></tr> <tr><td align='center'>I.</td><td align='center'>II.</td><td align='center'>III.</td><td align='center'>IV.</td></tr> <tr><td align='center'>7¾</td><td align='center'>8½</td><td align='center'>[7?]</td><td align='center'>6½</td><td align='center' rowspan='3'>} length of each bone in inches.</td></tr> <tr><td align='center'>5½</td><td align='center'>7¾</td><td align='center'>5½</td><td align='center'>[4½?]</td></tr> <tr><td align='center'>4½</td><td align='center'>——</td><td align='center'>——</td><td align='center'>——</td></tr> </table></div> The femur is represented by many examples—one 3¾ inches long, and others less than 3 inches long (29/10). In Campylognathus, which has so much in common with the jaw and the wing bones in size, the upper leg bone is 28/10 inches. Therefore if we assign the larger femur to the larger wing, the femur will be relatively longer in all species of Rhamphocephalus than in Campylognathus. Only one example of a tibia is preserved. It is 3½ inches long, or only 1/10 inch shorter than the bone in Campylognathus, which has the femur 28/10 inches, so that I refer the tibia of Rhamphocephalus to the species which has the intermediate length of wing. These coincidences with Campylognathus establish a close affinity, and may raise the question whether the Upper Lias species may not be included in the Stonesfield Slate genus Rhamphocephalus. The late Professor Phillips, in his Geology of Oxford, attempted a restoration of the Stonesfield Ornithosaur, and produced a picturesque effect (); but no restoration is possible without such attention to the proportions of the bones as we have indicated. [Pg 156] <h4>OXFORD CLAY</h4> A few bones of flying reptiles have been found in the Lower Oxford Clay near Peterborough, and others in the Upper Oxford Clay at St. Ives, in Huntingdonshire. A single tail vertebra from the Middle Oxford Clay, near Oxford, long since came under my own notice, and shows that these animals belong to a long-tailed type like Campylognathus. The cervical vertebræ are remarkable for being scarcely longer than the dorsal vertebræ; and the dorsal are at least half as long again as is usual, having rather the proportion of bones in the back of a crocodile. <h4>LITHOGRAPHIC SLATE</h4> Long-tailed Pterodactyles are beautifully preserved in the Lithographic Limestone of the south of Bavaria, at Solenhofen, and the quarries in its neighbourhood, often with the skeleton or a large part of it flattened out in the plane of bedding of the rock. Fine skeletons are preserved in the superb museum at Munich, at Heidelberg, Bonn, Haarlem, and London, and are all referred to the genus Rhamphorhynchus or to Scaphognathus. It is a type with powerfully developed wings and a long, stiff tail, very similar to that of Dimorphodon, so that some naturalists refer both to the same family. There is some resemblance. The type which is most like Dimorphodon is the celebrated fossil at Bonn, sometimes called Pterodactylus crassirostris, which in a restored form, with a short tail, has been reproduced in many text-books. No tail is preserved in the slab, and I ventured to give the animal a tail for the first time in a restoration () published by the Illustrated London News in 1875, which accompanied a report of a Royal [Pg 157] Institution lecture. Afterwards, in 1882, Professor Zittel, of Munich, published the same conclusion. The reason for restoring the tail was that the animal had the head constructed in the same way as Pterodactyles with a long tail, and showed differences from types in which the tail is short; and there is no known short-tailed Pterodactyle, with wrist and hand bones, such as characterise this animal. The side of the face has a general resemblance to the Pterodactyles from the Lias, for although the framework is firmer, the four apertures in the head are similarly placed. The nostril is rather small and elongated, and ascends over the larger antorbital vacuity. The orbit for the eye is the largest opening in the head, so that these three apertures successively increase in size, and are followed by the vertically elongated post-orbital vacuity. The teeth are widely spaced apart, and those in the skull extend some distance backward to the end of the maxillary bone. There are few teeth in the lower jaw, and they correspond to the large anterior teeth of Dimorphodon, there being no teeth behind the nasal opening. The lower jaw is straight, and the extremities of the jaws met when the mouth was closed. The breast bone does not show the keel which is so remarkable in Rhamphorhynchus, which may be attributed to its under side being exposed, so as to exhibit the pneumatic foramina. The ribs have double heads, more like those of a Crocodile in the region of the back than is the case with the bird-like ribs from Stonesfield. The second joint in the wing finger may be longer than the first—a character which would tend to the association of this Pterodactyle with species from the Lias; a relation [Pg 158] to which attention was first drawn by Mr. E. T. Newton, who described the Whitby skull. The Pterodactyles from the Solenhofen Slate which possess long tails have a series of characters which show affinity with the other long-tailed types. The jaws are much more slender. The orbit of the eye in Rhamphorhynchus is enormously large, and placed vertically above the articulation for the lower jaw. Immediately in front of the eye are two small and elongated openings, the hinder of which, known as the antorbital vacuity, is often slightly smaller than the nostril, which is placed in the middle length of the head, or a little further back, giving a long dagger-shaped jaw, which terminates in a toothless spear. The lower jaw has a corresponding sharp extremity. The teeth are directed forward in a way that is quite exceptional. Notwithstanding the massiveness and elongation of the neck vertebræ, which are nearly twice as long as those of the back, the neck is sometimes only about half the length of the skull. All these long-tailed species from the Lithographic Stone agree in having the sternum broad, with a long strong keel, extending far forward. The coracoid bones extend outward like those of a Crocodile, so as to widen the chest cavity instead of being carried forward as the bones are in Birds. These bones in this animal were attached to the anterior extremity of the sternum, so that the keel extended in advance of the articulation as in other Pterodactyles. The breadth of the sternum shows that, as in Mammals, the fore part of the body must have been fully twice the width of the region of the hip-girdle, where the slenderer hind limbs were attached. The length of the fore limb was enormous, for although the head [Pg 159] suggests an immense length relatively to the body, nearly equal to neck and back together, the head is not more than a third of the length of the wing bones. The wing bones are remarkable for the short powerful humerus with an expanded radial crest, which is fully equal in width to half the length of the bone. Another character is the extreme shortness of the metacarpus, usually associated with immense strength of the wing metacarpal bone. The hind limbs are relatively small and relatively short. The femur is usually shorter than the humerus, and the tibia is much shorter than the ulna. The bones of the instep, instead of being held together firmly as in the Lias genera, diverge from each other, widening out, though it often happens that four of the five metatarsals differ but little in length. The fifth digit is always shorter. The hip-girdle of bones differs chiefly from other types in the way in which those bones, which have sometimes been likened to the marsupial bones, are conditioned. They may be a pair of triangular bones which meet in the middle line, so that there is an outer angle like the arm of a capital Y. Sometimes these triangular bones are blended into a curved, bow-shaped arch, which in several specimens appears to extend forward from near the place of articulation of the femur. This is seen in fossil skeletons at Heidelberg and Munich. It is possible that this position is an accident of preservation, and that the prepubic bones are really attached to the lower border of the pubic bones. Immense as the length of the tail appears to be, exceeding the skull and remainder of the vertebral column, it falls far short of the combined length of the [Pg 160] phalanges of the wing finger. The power of flight was manifestly greater in Rhamphorhynchus than in other members of the group, and all the modifications of the skeleton tend towards adaptation of the animals for flying. The most remarkable modification of structure at the extremity of the tail was made known by Professor Marsh in a vertical, leaf-like expansion in this genus, which had not previously been observed (). The vertebræ go on steadily diminishing in length in the usual way, and then the ossified structures which bordered the tail bones and run parallel with the vertebræ in all the Rhamphorhynchus family, suddenly diverge downward and upward at right angles to the vertebræ, forming a vertical crest above and a corresponding keel below; and between these structures, which are identified with the neural spines and chevron bones of ordinary vertebræ, the membrane extends, giving the extremity of the tail a rudder-like feature, which, from knowledge of the construction of the tail of a child's kite, may well be thought to have had influence in directing and steadying the animal's movements. There are many minor features in the shoulder-girdle, which show that the coracoid, for example, was becoming unlike that bone in the Lias, though it still continues [Pg 161] to have a bony union with the elongated shoulder-blade of the back. [Pg 162] <div class="figcenter"> FIG. 56.   RESTORATION OF THE SKELETON OF RHAMPHORHYNCHUS PHYLLURUS From the Solenhofen Slate, partly based upon the skeleton with the wing membranes preserved <ANTIMG src="images/i_188.jpg" width-obs="1024" height-obs="630" alt="FIG. 56." title="FIG. 56." /></div> [Pg 163] <div class="figcenter"> FIG. 57.   RESTORATION OF THE SKELETON OF SCAPHOGNATHUS CRASSIROSTRIS Published in the Illustrated London News in 1875. In which a tail is shown on the evidence of the structure of the head and hand <ANTIMG src="images/i_190.jpg" width-obs="640" height-obs="408" alt="FIG. 57." title="FIG. 57." /></div> The great German delineator of these animals, Von Meyer, admitted six different species. Mr. Newton and Mr. Lydekker diminish the number to four. It is not easy to determine these differences, or to say how far the differences observed in the bones characterise species or genera. It is certain that there is one remarkable difference from other and older Pterodactyles, in that the last or fourth bone in the wing finger is usually slightly longer than the third bone, which precedes it. There is a certain variability in the specimens which makes discussion of their characters difficult, and has led to some forms being regarded as varieties, while others, of which less material is available, are classed as species. I am disposed to say that some of the confusion may have resulted from specimens being wrongly named. Thus, there is a Rhamphorhynchus called curtimanus, or the form with the short hand. It is represented by two types. One of these appears to have the humerus short, the ulna and radius long, and the finger bones long; the other has the humerus longer, the ulna much shorter, and the finger bones shorter. They are clearly different species, but the second variety agrees in almost every detail with a species named hirundinaceus, the swallow-like Rhamphorhynchus. This identification shows, not [Pg 164] that the latter is a bad species, but that curtimanus is a distinct species which had sometimes been confounded with the other. While most of these specimens show a small but steady decrease in the length of the several wing finger bones, the species called Gemmingi has the first three bones absolutely equal and shorter than in the species curtimanus, longimanus, or hirundinaceus. In the same way, on the evidence of facts, I find myself unable to join in discarding Professor Marsh's species phyllurus, on account of the different proportions of its limb bones. The humerus, metacarpus, and third phalange [Pg 165] of the wing finger in Rhamphorhynchus phyllurus are exceptionally short as compared with other species. Everyone agrees that the species called longicaudus is a distinct one, so that it is chiefly in slight differences in the proportions of constituent parts of the skeleton that the types of the Rhamphorhynchus are distinguished from each other. I cannot quite concur with either Professor Zittel (Fig. 58, 3) or Professor Marsh (Fig. 58, 2) in the expansion which they give to the wing membrane in their restorations; for although Professor Zittel represents the tail as free from the hind legs, while Professor Marsh connects them together, they both concur in carrying the wing membrane from the tip of the wing finger down to the extremity of the ankle joint. I should have preferred to carry it no further down the body than the lower part of the back, there being no fossil evidence in favour of this extension so far as specimens have been described. Neither the membranous wings figured by Zittel nor by Marsh would warrant so much body membrane as the Rhamphorhynchus has been credited with. I have based my restoration () of the skeleton chiefly on Rhamphorhynchus phyllurus. <div class="figcenter"> FIG. 58.   SIX RESTORATIONS <ANTIMG src="images/i_191.jpg" width-obs="600" height-obs="620" alt="FIG. 58." title="FIG. 58." /> 1. Ramphocephalus. Stonesfield Slate. John Phillips, 1871 2. Rhamphorhynchus. O. C. Marsh, 1882 3. Rhamphorhynchus. V. Zittel, 1882 4. Ornithostoma. Williston, 1897 5. Dimorphodon. Buckland, 1836. Tail then unknown 6. Ornithocheirus. H. G. Seeley, 1865 </div> <h4>THE SHORT-TAILED TYPES</h4> The Pterodactylia are less variable; and the variation among the species is chiefly confined to relative length of the head, length of the neck, and the height of the body above the ground. The tail is always so short as to be inappreciable. Many of the specimens are fragmentary, and the characters of the group are not easily determined without careful comparisons and measurements. The bones of the [Pg 166] fore limb and wing finger are less stout than in the Rhamphorhynchus type, while the femur is generally a little longer than the humerus, and the wing finger is short in comparison with its condition in Rhamphorhynchus. These short-tailed Pterodactyles give the impression of being active little animals, having very much the aspect of birds, upon four legs or two. The neck is about as long as the lower jaw, the antorbital vacuity in the head is imperfectly separated from the much larger nasal opening, the orbit of the eye is large and far back, the teeth are entirely in front of the nasal aperture, and the post-orbital vacuity is minute and inconspicuous. The sternum is much wider than long, and no specimens give evidence of a manubrium. The finger bones progressively decrease in length. The prepubic bones have a partially expanded fan-like form, and never show the triradiate shape, and are never anchylosed. About fifteen different kinds of Pterodactyles have been described from the Solenhofen Slate, mostly referred to the genus Pterodactylus, which comprises forms with a large head and long snout. Some have been placed in a genus (Ornithocephalus, or Ptenodracon) in which the head is relatively short. The majority of the species are relatively small. The skull in Ornithocephalus brevirostris is only 1 inch long, and the animal could not have stood more than 1½ inches to its back standing on all fours, and but little over 2½ inches standing as a biped, on the hind limbs. A restoration of the species called Pterodactylus scolopaciceps, published in 1875 in the Illustrated London News in the position of a quadruped, shows an animal a little larger, with a body 2½ inches high [Pg 167] and 6 to 7 inches long, with the wing finger 4½ inches long. Larger animals occur in the same deposit, and in one named Pterodactylus grandis the leg bones are a foot long; and such an animal may have been nearly a foot in height to its back, standing as a quadruped, though most of these animals had the neck flexible and capable of being raised like the neck of a Goose or a Deer (), and bent down like a Duck's when feeding. <div class="figcenter"> FIG. 59.   RESTORATION OF THE SKELETON OF PTENODRACON BREVIROSTRIS From the Solenhofen Slate. The fourth joint of the wing finger appears to be lost and has not been restored in the figure. (Natural size) <ANTIMG src="images/i_194.jpg" width-obs="877" height-obs="768" alt="FIG. 59." title="FIG. 59." /></div> The type of the genus Pterodactylus is the form originally described by Cuvier as Pterodactylus longirostris (). It is also known as P. antiquus, that name having been given by a German naturalist after Cuvier had invented the genus, and before he had [Pg 168] named the species. There are some remarkable features in which Cuvier's animal is distinct from others which have been referred to the same genus. Thus the head is 4½ inches long, while the entire length of the backbone to the extremity of the tail is only 6½ inches, and one vertebra in the neck is at least as long as six in the back, so that the animal has the greater part of its length in the head and neck, although the neck includes so few vertebræ. Nearly all the teeth—which are few in number, short and broad, not exceeding a dozen in either jaw—are limited to the front part of the beak, and do not extend anywhere near the nasal vacuity. This is not the case with all. In the species named P. Kochi, which I have regarded as the type of a distinct genus, there are large teeth in the front of the jaw corresponding to those of Pterodactylus, and behind these a smaller series of teeth extending back under the nostril, which approaches close to the orbit of the eye, without any indication of a separate antorbital vacuity. On those characters the genus Diopecephalus was defined. It is closely allied to Pterodactylus; both agree in having the ilium prolonged forward more than twice as far as it is carried backward, the anterior process covering about half a dozen vertebræ, as in Pterodactylus longirostris. A great many different types have been referred to Pterodactylus Kochi, and it is probable that they may eventually be distinguished from each other. The species in which the upper borders of the orbits approximate could be separated from those in which the frontal interspace is wider. <div class="figcenter"> FIG. 60.   CYCNORHAMPHUS SUEVICUS FROM THE SOLENHOFEN SLATE SHOWING THE SCATTERED POSITION OF THE BONES Original in the Museum at Tübingen <ANTIMG src="images/i_196.jpg" width-obs="854" height-obs="768" alt="FIG. 60." title="FIG. 60." /></div> [Pg 169] <div class="figcenter"> FIG. 61.   CYCNORHAMPHUS SUEVICUS RESTORATION SHOWING THE FORM OF THE BODY AND THE WING MEMBRANES <ANTIMG src="images/i_198.jpg" width-obs="957" height-obs="768" alt="FIG. 61." title="FIG. 61." /></div> It is a remarkable feature in these animals that the middle bones of the foot, termed instep bones or metatarsals, are usually close together, so that the toes diverge from a narrow breadth, as in P. longirostris, P. Kochi, and other forms; but there also appear to be splay-footed groups of Pterodactyles like the species which have been named P. elegans and P. micronyx, in which the metatarsus widens out so that the bones of the toes do not diverge, and that condition characterises the Ptenodracon (Pterodactylus brevirostris), to which genus these species may possibly be referred. Nearly all who have studied these animals regard the singularly short-nosed species P. brevirostris as forming a separate genus. For that genus Sömmerring's descriptive name Ornithocephalus, which he used for Pterodactyles generally, might perhaps have been retained. But the name Ptenodracon, suggested by Mr. Lydekker, has been used for these types. Some of the largest specimens preserved at Stuttgart and Tübingen have been named Pterodactylus suevicus and P. Fraasii. They do not approach the species P. grandis in size, so far as can be judged from the fragmentary remains figured by Von Meyer; for what appears to be the third phalange of the wing finger is 7½ inches long, while in these species it is less than half that length, indicating an enormous development of wing, relatively to the length of the hind limb, which would probably refer the species to another genus. Pterodactylus suevicus differs from the typical Pterodactyles in having a rounded, flattened under surface to the lower jaw, instead of the common condition of a sharp keel in the region of the symphysis. The beak also seems flattened and swan-like, and the teeth are limited to [Pg 170] the front of the jaw. There appear to be some indications of small nostrils, which look upward like the nostrils of Rhamphorhynchus, but this may be a deceptive appearance, and the nostrils are large lateral vacuities, which are in the position of antorbital vacuities, so that there would appear to be only two vacuities in the side of the head in these animals. The distinctive character of the skeleton in this genus is found in the extraordinary length of the metacarpus and in the complete ossification of the smaller metacarpal bones throughout their length. The metacarpal bones are much longer than the bones of the fore-arm, and about twice the length of the humerus. The first wing phalange is much longer than the others, which successively and rapidly diminish in length, so that the third is half the length of the first. There are differences in the pelvis; for the anterior process of the ilium is very short, in comparison with its length in the genus Pterodactylus. And the long stalk of the prepubic bone with its great hammer-headed expansion transversely in front gives those bones a character unlike other genera, so that Cycnorhamphus ranks as a good genus, easily distinguished from Cuvier's type, in which the four bones of the wing are more equal in length, and the last is more than half the length of the first; while the metacarpus in that genus is only a little longer than the humerus, and much shorter than the ulna. The Pterodactylus suevicus has the neck vertebræ flat on the under side, and relatively short as compared with the more slender and narrower vertebræ of P. Fraasii. <div class="figcenter"> FIG. 62.   CYCNORHAMPHUS SUEVICUS Skeleton restored from the bones in Fig. 60 <ANTIMG src="images/i_201.jpg" width-obs="599" height-obs="480" alt="FIG. 62." title="FIG. 62." /></div> <div class="figcenter"> FIG. 63.   RESTORATION OF SKELETON CYCNORHAMPHUS FRAASI SHOWING THE LIMBS ON THE RIGHT SIDE From a specimen in the Museum at Stuttgart <ANTIMG src="images/i_202.jpg" width-obs="768" height-obs="911" alt="FIG. 63." title="FIG. 63." /></div> [Pg 171] <div class="figcenter"> FIG. 64.   CYCNORHAMPHUS FRAASI RESTORATION OF THE FORM OF THE BODY <ANTIMG src="images/i_204.jpg" width-obs="768" height-obs="894" alt="FIG. 64." title="FIG. 64." /></div> <hr style="width: 65%;" /> [Pg 172] <div style="break-after:column;"></div>

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