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The History of Fossil Crocodiles

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The History of Fossil Crocodiles 288 A. SMITH WOODWARD ON THE THE HISTORY OF FOSSIL CROCODILEB. By ARTHUR SMITH WOODWARD, F.G.S., Assistant in the Geological Department, British Museum (Natural History.) 1.-INTRODUCTION. The rapid progress of every department of scientific research, and the unfortunate scattering of the various contributions to the same subject in the different journals and periodicals throughout the world. renders the task of the compiler and historian of no small importance, " Stock-taking" in science is as important in its own way as is the same procedure in commercial undertakings, and occasional reviews of the present state of any particular branch of inquiry are of the greatest value in indicating the most promising paths for further traverses, and the most profitable domains for future conquest. The abstruse nature of much of the literature, too, and the out-of­ the-way character of most of the older, and even some of the newer, works and memoirs, make it often advisable to possess a less de­ tailed and more accessible account of what they contain; and it is the remembrance of such considerations as these that has induced the writer to offer the present communication as a slight attempt towards providing, in the form of a connected narrative, an outline of what is already known concerning one small group of fossil ver­ tebrate animals-that of the Crocodilia. It is proposed to enumerate, in order, the more important extinct types of this ancient reptilian class whose remains are met with in the various successive formations, and, while referring more par­ ticularly to those discovered in British deposits, to make usc of all available materials to elucidate the developmental history of the race. And if, at times, more attention is given to fragmentary fossils than their small importance may be deemed to justify, we plead, as an excuse, that such relics frequently fall into the hands of collectors, and the desultory notices of them to be found in ordi­ nary text-books are usually much too brief and indefinite to satisfy the curiosity of those who pursue their collecting in a thoroughly philosophical manner. HISTORY OF FOSSIL CROCODILES. 289 n.-RECENT CROCODILES. To render the fossils intelligible, however, and to bring out their features of greatest interest, it is necessary, in the first place, to take a general glance at the tribes of Crocodiles as they exist in the liv­ ing fauna of the globe;* or, in the words of modern biology, to consider the culmination of the Crocodilian" phylum" as it is to be observed at the present day. With regard to the number of species still surviving, it is almost impossible to make any definite statement, on account of the great differenees of opinion that prevail as to what really constitute specific characters; and the same remark applies to the generic divisions; t but some or other members of the class are still to be met with in nearly all tropical and subtropical regions of the earth's surface t-notwithstanding their rapidly approaching extinction in many parts through the interference of man-and there are at least two, if not three, well-marked types or families still to be distin­ guished. There are the Gavials or Gharials specially adapted for an aquatic existence; and the Crocodiles and Alligators, more accus­ tomed to an amphibious mode of life. According to Dr. Gray, the true Gavials (Gharials) are at present exclusively confined to the rivers of the interior of India, and are there found only in the lower regions where the climate is hot. Their most prominent and striking superficial peculiarity con­ sists in the great elongation and slenderness of the snout (Fig. 1), which is armed with a numerous series of sharp, equal-sized teeth, admirably adapted for the capture of slippery prey; this peculiarity, * For good illustrations of the osteology of living Crocodiles, the follow­ ing works may be consulted :-G. Ouvier, 'Ossemens Fossiles;' De Blain­ ville, 'Osteographie j' d'Alton and Burmeister, 'Del' Fossile Gavial von Boll,' Halle, 1854 j' and" Synopsis," by J. E. Gray, in 'Traus. Zool. Soc., Vol. vi, quoted below. t T. H. Huxley, "Notes on the Specific and Generic Characters of Recent Crocodilia."-' Proc. Linn. Soc.,' Vol. iv (Zoology), 1860, p. 1; J. E. Gray, "Synopsis of the Species of Recent Crocodilians or Emydosaurians."­ 'Trans. Zool. Soc.,' Vol. vi (1862), pp. 125.169, pls. xxxi- xxxiv ; J. E. Gray, " On the Change of Form of the Heads of Orocodiles."-' Brit. Assoc. Re­ ports,' 1862, p. 109; J. E. Gray, "Catalogue of Shield Reptiles in the British Museum," Part 2, 1872. t For the geographical distribution of recent Crocodilia, see A. Strauch, "Synopsis del' gegenwart.ig lebenden Crocodiliden."-' Mem. Acad, Imp. Sci. St. Petersbourg ' [7J, Vol. x, No. 13 (1866). 290 A. SMITH WOODWARD ON THE in fact, being one distinguishing the Gavial family from those of the Crocodiles and Alligators, and the other most essential distinctions being also found in tbe same parts. In the lower jaw of the Gavial (Fig. 1), for example, the two halves (or "rami") are united together by a very long suture, and scarcely any of the teeth arc fixed to the jaw behind this" symphysial area;" again, the two bones that contribute largely towards roofing in the front part of the nasal cavity in most vertebrates, and hence called" nasals," are fixed quite at the commencement of the snout, and do not reach the external nostril at all; and there are also some other minor features that may be equally looked upon as distinctive. 2. FIG. 1. Mandible of Gavialis gangeticus. FIG. 2. Mandible of Crocodilus palustris. Turning next to the true Crocodiles, we find their geographical range much more extensive; they occur both in India, Africa, and the tropical parts of America, and at least two species, also, have their haunts in the rivers and lakes of Australia. They are easily distinguished from the Gavials by the form and character of the skull (Figs. 3, 9) ; the jaws are relatively shorter, and gently rounded in front, while the edges, instead of being straight, have a " fes- HISTORY OF FOSSIL CROCODILES. 291 tooned" arrangement, and the powerful teeth are much more unequal in size. The nasal bones, too, reach-or closely approach -the hinder border of the external nostril, and the two rami of the lower jaw (Fig. 2) are only united in front for a very short dis­ tance. The form and proportions of the head, in fact, are more adapted for the seizure of great mammalian quadrupeds than for the slippery fishes upon which the Gavial feeds. The Alligators-with the exception of one lately discovered in. China-are exclusively confined to America, and the single species inhabiting the Mississippi is particularly noteworthy on account of its being occasionally met with in the oft-frozen regions of the North. Two other genera, occurring chiefly in the tropics of the Southern Continent, are also of great interest to the palreontologist, because, of still living forms, they alone are completely encased all round in the bony armour that formed such a conspicuous feature in the majority of extinct Crocodilian genera. On a superficial glance, it is by no means easy to distinguish the members of this family from the true Crocodiles, and, as has already been remarked, some zoologists are inclined to unite them together much more closely than has long' been customary. The Alligators differ, how­ ever, in the circumstance that the fourth tooth of the lower jaw on each side-often called the" canine," from its great relative size­ is received in a socket in the upper jaw, while the corresponding tooth of the Crocodile simply fits in a groove and is visible when the mouth is closed. The dentition of the Alligators is further characteristic from the peculiarity of the lower teeth passing on the inner side of the upper teeth in biting, instead of fitting into the interspaces of the upper row. Summarising the more important of these differential char­ acters," it may be convenient to have them arranged in tabular form, and the following brief comparative statement will, perhaps, facilitate future reference. (See next page.) * T. H. Huxley, "Notes on the Specific and Generic Characters of Recent Crocodilia."-' Proo, Linn. Soc.,' Vol. iv (Zoology), 1860, p.T, 292 A. SMITH WOODWARD ON THE CROCODI L ID.E. GAVIAL I D.E . ALLIGATORI D.E . C n o c o Dl LI D.-E PROP ER. Skull short and broad. Skull more or less Snout very long and short and broad. slender. Nas al bones reach ing Nasal bones r eaching Nas al bones short, and external nostr il, external nostril, or not r eaching the pre­ sometimes di viding (at least) the pre­ maxillre (except in it. maxillas. Tomist oma), Teeth stout, and vari­ Teeth stout, and more Teeth slen de r, shar p' able in size. or less var ia ble in edged, a nd subequal. size. Anterior pair of m an­ Anter ior pa ir of man­ An terior pair of man­ di bular teeth, and dibular teeth re ­ di bul ar teeth, and fourth pai r, receivcd ceived in pits of four th pair, pass in. in pit s in the mar­ pramaxillas, but to gr ooves of pre' g ins of the premax­ fourth pair passing maxillm. Illeoand maxillee, only in to grooves a t junction of pre . maxillas wit h max. Illes, Mandibular teeth be. Mandihular teeth be­ Mandibular teeth be­ hind the fourth pass hi nd th e fourth pass hind the fourth pass inside the maxillary between the maxil­ bet ween the maxil­ t eeth.
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