(Lat. Reptilia, creeping things, from reptilis; refere, to creep; Gr. g pirav, whence the term " herpetology," for the science dealing with them). In the days before Linnaeus, writers comprised the animals which popularly are known as tortoises and turtles, crocodiles, lizards and snakes, frogs and toads, newts and salamanders, under the name of oviparous quadrupeds or four-limbed animals which lay eggs. Linnaeus, desirous of giving expression to the extraordinary fact that many of these animals pass part of their life in the water and part on land,' substituted the name of Amphibia for the ancient term. Subsequent French naturalists (Lyonnet 2 and Brisson 3) considered that the creeping mode of locomotion was a more general characteristic of the class than their amphibious habits, and consequently proposed the scarcely more appropriate name of Reptiles. As naturalists gradually comprehended the wide gap existing between frogs, toads, &c., on the one hand, and the other oviparous quadrupeds on the other, they either adopted the name of Batrachia for the former and that of Amphibia for the latter, or they restricted the term Amphibia to Batrachians, calling the remainder of these creatures reptiles. Thus the term Amphibia, as used by various authors, may apply (1) to all the various animals mentioned, or (2) to Batrachians only (see Batrachia). The term Reptiles (Reptilia) is used (i) by some for all the animals mentioned above, and (2) by others, as in the present article, for the same assemblage of animals after the exclusion of Batrachians.

Equally varying are the limits of the term Saurians, which occurs so frequently in every scientific treatise on this subject. At first it comprised living crocodiles and lizards only, with which a number of fossil forms were gradually associated. As the characters and affinities of the latter became better known, some of them were withdrawn from the Saurians, and at present it is best to abandon the term altogether.

I. History Of Herpetology Certain kinds of reptiles are mentioned in the earliest written records or have found a place among the fragments of the oldest relics of human art. Such evidences, however, form no part of a succinct review of the literature of the subject such as it is proposed to give here. We distinguish in it six periods: (I) the Aristotelian; (2) the Linnaean (formation of a class Amphibia, in which reptiles and Batrachians are mixed); (3) the period of the elimination of Batrachians as one of the reptilian orders (Brongniart); (4) that of the separation of reptiles and Batrachians as distinct subclasses; (5) that of the recognition of a class Reptilia as part of the Sauropsida (Huxley); (6) that of the discovery of fossil skeletons sufficiently well preserved to reveal, in its general outlines, the past history of the class.

1. The Aristotelian Period. - Aristotle was the first to deal with the reptiles known to him as members of a distinct portion Aristotle. of the animal kingdom, and to point out the characteristics by which they resemble each other and differ from other vertebrate and invertebrate animals. The plan of his ' " Polymorpha in his amphibiis natura duplicem vitam plerisque concessit." 2 Theologie des insectes de Lesser (Paris, 1 745), i. 91, note 5. Regne animal divise en neuf classes (Paris, 1756).

work, however, was rather that of a comparative treatise of the anatomical and physiological characters of animals than their systematic arrangement and definition, and his ideas about the various groups of reptiles are not distinctly expressed, but must be gleaned from the terms which he employs. Moreover, he paid less attention to the study of reptiles than to that of other classes. This is probably due to the limited number of kinds he could be acquainted with, to which only very few extraEuropean forms, like the crocodile, were added from other sources. But while we find in some respects a most remarkable accuracy of knowledge, there is sufficient evidence that he neglected everyday opportunities of information. Thus, he has not a single word about the metamorphoses of Batrachians, which he treats of in connexion with reptiles.

Aristotle makes a clear distinction between the scute or scale of a reptile, which he describes as 40Xis, and that of a fish, which he designates as AE7ris. He mentions reptiles (I) as oviparous quadrupeds with scutes, viz. Saurians and Cheloniai s; (2) as oviparous apodals, viz. Snakes; (3) as oviparous quadrupeds without scutes, viz. Batrachians. He considered the first and second of these three groups as much more nearly related to each other than to the third. Accurate statements and descriptions are sadly mixed with errors and stories of, to our eyes, the most absurd and fabulous kind. The most complete accounts are those of the crocodile (chiefly borrowed from Herodotus) and of the chameleon, which Aristotle evidently knew from personal observation, and had dissected himself. The other lizards mentioned by him are the common lizards (aavpa), the common seps (X aXKis or "ctyvis) and the gecko (aaKaXa(3c. Trts or Kop8bXos). Of snakes (of which he generally speaks as 6 s) he knew the vipers ( g xis or g XLBva), the common snake (155pos), and the blindworm (roc/Aim ac/as), which he regards as a snake; he further mentions the Egyptian cobra and dragons (5peuc v) - North-African serpents of fabulous size. Of Chelonians he describes in a perfectly recognizable manner land tortoises (XEAc ? vrt), freshwater turtles (Eµos) and marine turtles (XEAc.vn OaXarria). Passing over eighteen centuries, we find the knowledge of reptiles to have remained as stationary as other branches of natural history, perhaps even more so. The reptile fauna of Europe was not extensive enough to attract the energy of a Belon or Rondelet; popular prejudice and the difficulty of preserving these animals deterred from their study; nor was man sufficiently educated not to give implicit credence to the fabulous tales of reptiles in the i 5th and 16th centuries. The art of healing, however, was developing into a science based upon rational principles, and consequently not only those reptiles which formed part of the materia medica but also the venomous snakes became objects of study to the physician, though the majority of the writers were ignorant of the structure of the venom-apparatus, and of the distinction between non-venomous and venomous snakes.

Nothing can show more clearly the small advance made by herpetology in this long post-Aristotelian period than a glance at the celebrated work, De Differentiis Animalium Libri decem (Paris, 1552), by Edward Wotton (1492 '555). Wotton treats of the reptiles which he designates as Quadrupedes oviparae et Serpentes in the sixth book of his work. They form the second division of the Quadrupedes quae sanguinem habent, and am subdivided in the following " genera ": Crocodilus et scincus (cap. cv.); Testudinum genera (cvi.); Ranarum genera (cvii.); Lacertae (cviii.); Salamandra et seps quadrupes (cix.); Stellio (cx.); Chamaeleo (cxi.); Serpentes (cxii.), a general account, the following being different kinds of serpents: Hydrus et alii quidam serpentes aquatiles (cxiii.); Serpentes terrestres et primo aspidum genera (cxiv.); Vipera, dipsas, cerastes, et hammodytes (cxv.); Haemorrhus, sepedon, seps, cenchris, et cenchrites (cxvi.); Basiliscus et alii quidam serpentes quorum venenum remedio caret (cxvii.); Draco, amphisbaena, et alii quidam serpentes quorum morsus minus affert periculi (cxviii.).

Wotton's work might with propriety be termed " Aristoteles redivivus." The plan is the same, and the observations of the Greek naturalist are faithfully, sometimes literally, reproduced.

It is surprising that even the reptiles ^f his native country were most imperfectly known to the author.

With the enlargement of geographical knowledge that of reptiles was also advanced, as is sufficiently apparent from the. large encyclopaedic works of Gesner, Aldrovandi and Johnston. The last-named author especially, who published the various portions of his Natural History in the middle of the 17th century, was able to embody in his compilations notices of numerous reptiles observed by Francisco Hernandez in Mexico and by Marcgrave and Piso in Brazil. As the author had no definite idea of the Ray-Linnaean term " species," it is not possible to give the exact number of reptiles mentioned in his work. But it may be estimated at about fifty, not including some marine fishes and fabulous creatures. He figures (or rather reproduces the figures of) about forty - some species being represented by several figures.

2. Linnaean Period: Formation of a Class Amphibia.- Within the century which succeeded these compilatory works (1650-1750) fall the labours which prepared the way sors of for and exerted the greatest influence on Ray and Linnaeus. Although original researches in the field of herpetology were limited in extent and in number, the authors had freed themselves from the purely literary or scholastic tendency. Men were no longer satisfied with reproducing and commenting on the writings of their predecessors; the pen was superseded by the eye, the microscope and the knife, and statements were tested by experiment. This spirit of the age manifested itself, so far as the reptiles are concerned, in Chara's and Redi's admirable observations on the viper, in Major's and Vallisnieri's detailed accounts of the anatomy of the chameleon, in the researches of Jacobaeus into the metamorphoses of the Batrachians and the structure of lizards, in Dufay's history of the development of the salamander (for Batrachians are invariably associated with reptiles proper); in Tyson's description of the anatomy of the rattlesnake, &c. The natural history collections formed by institutions and wealthy individuals now contained not merely skins of crocodiles or serpents stuffed and transformed into a shape to correspond with the fabulous descriptions of the ancient dragons, but, with the discovery of alcohol as a means of preserving animals, reptiles entire or dissected were exhibited for study; and no opportunity was lost of obtaining them from travellers or residents in foreign countries. Fossils also were now acknowledged to be remains of animals which had lived before the Flood, and some of them were recognized as those of reptiles.

The contributions to a positive knowledge of the animal kingdom became so numerous as to render the need of a methodical arrangement of the abundance of new facts more and more pressing. Of the two principal systematic attempts made in this period the first ranks as one of the most remarkable steps of the progress of natural history, whilst the second can only be designated as a signal failure, which ought to have been a warning to all those who in after years classified animals in what is called an, " artificial system." As the latter attempt, originating with Klein (1685-1759), did not exercise any further influence on herpetology, it will be sufficient to have merely Ray. mentioned it. John Ray (1628-1705) had recognized R the necessity of introducing exact definitions for the several categories into which the animals had to be divided, and he maintained that these categories ought to be characterized by the structure of animals, and that all zoological knowledge had to start from the " species " as its basis. His definition of reptiles as " animalia sanguinea pulmone respirantia cor unico tantum ventriculo instructum habentia ovipara " fixed the class in a manner which was adopted by the naturalists of the succeeding hundred and fifty years. Nevertheless, Ray was not a herpetologist; his knowledge of reptiles is chiefly derived from the researches of others, from whose accounts, however, everything not based upon reliable demonstration is critically excluded. He begins with a chapter treating of frogs (Rana, with two species), toads (Bufo, with one species) and tortoises' (Testudo, with fourteen species). The second group comprises the Lacertae, twenty-five in number, and includes the salamander and newts; and the third the Serpentes, nine species, among which the limbless lizards are enumerated.

Except in so far as he made known and briefly characterized a number of reptiles, our knowledge of this class was not advanced by Linnaeus. That he associated in the 12th edition cartilaginous and other fishes with the reptiles under the name of Amphibia Nantes was the result of some misunderstanding of an observation by Garden, and is not to be taken as a premonitory token of the recent discoveries of the relation between Batrachians and fishes. Linnaeus places reptiles, which he calls Amphibia, as the third class of the animal kingdom; he divides the genera thus: Order I. Reptiles. - Testudo (15 species); Rana (17 sp.); Draco (2 sp.); Lacerta (48 sp., including 6 Batrachians).

Order 2. Serpentes. - Crota1US (5 species); Boa (10 sp.); Coluber (96 sp.); Anguis (15 sp.); Amphisbaena (2 sp.); Caecilia (2 sp.).

None of the naturalists who under the direction or influence of Linnaeus visited foreign countries possessed any special knowledge of or predilection for the study of reptiles; all, however, contributed to our acquaintance with tropical forms, or transmitted well-preserved specimens to the collections at home, so that Gmelin, in the 13th edition of the Systema Naturae, was able to enumerate three hundred and seventy-one species. The man who, with the advantage of the Linnaean method, - first treated of reptiles monographically, was Laurenti. In a small book 2 he proposed a new division of these animals, of which some ideas and terms have survived into our times, characterizing the orders, genera and species in a much more precise manner than Linnaeus, giving, for his time, excellent descriptions and figures of the species of his native country. Laurenti might have become for herpetology what Artedi was for ichthyology, but his resources were extremely limited.

The circumstance that Chelonians are entirely omitted from his Synopsis seems due rather to the main object with which he engaged in the study of herpetology, viz. that of examining and distinguishing reptiles reputed to be poisonous, and to want of material, than to his conviction that tortoises should be relegated to another class. He divides the class into three orders: I. Salientia, with the genera Pipa, Bufo, Rana, Hyla, and one species of " Proteus," viz. the larva of Pseudis paradoxa. 2. Gradientia, the three first genera of which are Tailed Batrachians, viz. two species of Proteus (one being the P. anguinus), Triton and Salamandra; followed by true Saurians- Caudiverbera, Gecko, Chamaeleo, Iguana, Basiliscus, Draco, Cordylus, Crocodilus, Scincus, Stellio, Seps. 3. Serpentia, among which he continues to keep Amphisbaena, Caecilia and Anguis, but the large Linnaean genus Coluber is divided into twelve, chiefly from the scutellation of the head and form of the body.

The work concludes with an account of the experiments made by Laurenti to prove the poisonous or innocuous nature of those reptiles of which he could obtain living specimens.

The next general work on reptiles is by Lacepede. It appeared in the years 1788 and 1790 under the title Histoire naturelle des quadrupedes ovipares et des serpens (Paris, Lacepede. 2 vols. 4to). Although as regards treatment of details and amount of information this work far surpasses the modest attempt of Laurenti, it shows no advance towards a more natural division and arrangement of the genera. The author depends entirely on conspicuous external characters, and classifies the reptiles into (1) oviparous quadrupeds with a tail, (2) oviparous quadrupeds without a tail, (3) oviparous 1 In associating tortoises with toads, Ray could not disengage himself from the general popular view as to the nature of these animals, which found expression in the German Schildkrote (" Shieldtoad ").

Specimen medicum exhibens Synopsin Reptilium emendatam cum experimentis circa venena et antidota Reptilium Austriacorum (Vienna. 1768, 8vo, pp. 214, with 5 plates).

XXIII. 5 'a ' 0v 4 - Der , I 3, 1 31 "4Â°, lac,i.ti,ng,1 3 cj, +004d, 14 0,14 t. ??&????41????`"s'? V/6-ct 4'?,1a1, 1???1?? ^l; n,i?' 1s t ?ll y ?S i f,4,1 merely regarded as Amphibia because they closely resemble the genera which are proved to have been gill-breathers when immature. All these genera, however, so far as known, agree with the existing Amphibia in the production of their large parasphenoid bone as far forwards as the vomers to form a rigid and complete basicranial axis (fig. i, A). Those genera which less resemble the typical Labyrinthodonts are characterized by the reduction of the parasphenoid bone so that it no longer reaches the vomers; in these animals the weakened skull exhibits a secondary basicranial axis formed by the approximation of the pterygoids to the median line (fig. i, B). The latter condition is universal in existing reptiles, and may therefore perhaps be regarded as a diagnostic feature. If so, the oldest known undoubted reptile is Palaeohatteria, from the Lower Permian of Saxony.

In the structure of the skull Palaeohatteria is much like the existing Sphenodon, the cheek-plates which cover the temporal and masseter muscles on each side being pierced by two great vacuities, one superior-temporal, the other lateral-temporal. The majority of the earliest reptiles, however, either resemble the Labyrinthodonts in having the biting muscles completely covered with a roof of bony plates, or exhibit a slight shrinkage of this investment so that a superior-temporal vacuity appears. As the various groups or orders become differentiated, this shrinkage or reduction continues, while the shape of the ossifying ear-capsule changes, and the squamosal bone, which covers the organ of hearing in the fishes, and presumably also in the Palaeozoic Batrachia, is gradually thrust outwards from all connexion with this capsule except at its hinder angle. The resultant modifications are diagrammatically represented in fig 2. In one series of orders, comprising the Anomodontia, Chelonia, Sauropterygia and Ichthyopterygia (fig. 2, B, C), the superior-temporal vacuity (s) first appears, and the cheekplates in the broad temporal arch thus formed may be variously fused together, sometimes even irregularly perforated - showing at first, indeed, the usual inconstancy of a new and not completely established feature. From the earliest members of this series of reptiles, palaeontology seems to demonstrate that the Mammalia (with one robust temporal arcade or zygomatic arch) arose. In a second series, comprising the orders Rhynchocephalia, Dinosauria, Crocodilia and Ornithosauria (fig. 2, D), the broad arch of cheek-plates is regularly pierced by a lateraltemporal vacuity, which leaves a narrow bar above, another narrow bar below, and uncovers the middle part of the quadrate bone. By the constant loss of the lower, and the frequent loss of the upper, bar, some members of this series eventually pass into the order Squamata (Lacertilia+Ophidia), in which the quadrate bone is completely exposed and loosely attached to the skull (fig. 2, E); other reptiles exhibiting a similar modification may readily have acquired the typical Avian skull (fig. 2, F) by the loss of the upper and the retention of the lower temporal bar in question.

In view of these and other palaeontological considerations, the Reptilia may be classified into orders as follows: - Orders Of Class Reptilia I. Anomodontia. - Bones of postero-lateral region of skull forming a complete roof over the temporal and masseter muscles, or contracted into a single broad zygomatic arch, leaving a superior-temporal vacuity. Pineal foramen present. Ribs completely or imperfectly doubleheaded. No abdominal ribs. A large separately ossified epicoracoid. Limbs for support as well as progression; third and fourth digits with not more than three phalanges. Dermal armour feeble or absent. Range. - Permian and Triassic.

2. Chelonia. - Postero-lateral region of skull as in Anomodontia, except bones of ear-capsule more modified. No pineal foramen. Ribs single-headed. No sternum. Pectoral and pelvic arches unique in being situated completely inside the ribs. No epicoracoid. Abdominal ribs replaced by three or four pairs of large plates, which, with the clavicles and interclavicle, form a plastron. Limbs only for progression; third and fourth digits with not more than three phalanges. A regular dorsal carapace of bony plates intimately connected with the neural spines, and ribs of seven to nine dorsal vertebrae. Range. - Upper Triassic to Recent.

3. Sauropterygia. - Bones of postero-lateral region of skull contracted into a single broad zygomatic arch, leaving a superior-temporal vacuity. Pineal foramen present. No fused sacral vertebrae. All dorsal ribs single-headed, articulating with transverse processes of the neural arches. Abdominal ribs forming dense plastron. Apparently no sternum. Coracoid, pubis and ischium in form of much-expanded plates. Limbs modified as paddles, with not more than five digits, of which the third and fourth always have more than three phalanges; all digits usually consisting of numerous phalanges. No dermal armour. Range. - Upper Triassic to Cretaceous.

4. Ichthyopterygia. - Bones of postero-lateral region of skull contracted into a single broad zygomatic arch, leaving a superiortemporal vacuity. Pineal foramen present. Vertebral centra short and deeply biconcave, with feeble neural arches which are almost or completely destitute of zygapophyses. No fused sacral vertebrae. Cervical and dorsal ribs double-headed, articulating with tubercles on the vertebral centra. Abdominal ribs forming dense plastron. Apparently no sternum. Coracoid an expanded plate, probably with cartilaginous epicoracoid. Pelvis very small, not connected with vertebrae. Limbs modified as paddles, with digits of very numerous short phalanges, which are closely pressed together, sometimes with supplementary rows of similar ossicles. No dermal armour. A vertical triangular caudal fin, not supported by skeletal rays. Range. - Triassic to Cretaceous.

5. Rhynchocephalia. - Bones of postero-lateral region of skull contracted into two slender zygomatic bars, leaving a superiortemporal and a lateral-temporal vacuity, and partly exposing the quadrate bone from the side. Pineal foramen present or absent. Ribs single-headed. Abdominal ribs present. Sternum present. Epicoracoid cartilaginous. Limbs only for progression; third and fourth digits with four or five phalanges. Dermal armour feeble or absent. Range. - Lower Permian to Recent.

6. Dinosauria. - Postero-lateral region of skull as in Rhynchocephalia. No pineal foramen. Cervical and dorsal ribs doubleheaded. Rarely abdominal ribs. Sternum present, but apparently no clavicular arch. Limbs for support as well as progression; third and fourth digits with four and five phalanges respectively, Dermal armour variable. Range. - Triassic to Cretaceous.

7. Crocodilia. - Postero-lateral region of skull as in Rhynchocephalia. No pineal foramen. Cervical and dorsal ribs doubleheaded. Abdominal ribs present. Sternum present; also interclavicle, but no clavicles. Limbs only for progression on land or swimming; third and fourth digits with four or five phalanges. Dermal armour variable. Range. - Lower Jurassic to Recent.

Ornithosauria. - All bones extremely dense, light and hollow, the organism being adapted for flight. Postero-lateral region of skull as in Rhynchocephalia. No pineal foramen. Cervical and dorsal ribs double-headed. Abdominal ribs present. Sternum present, and keeled for attachment of pectoral muscles; no clavicular arch. Fifth digit of hand much elongated to support a wingmembrane, but with only four phalanges. Hind limb feeble. No dermal armour. Range. - Lower Jurassic to Cretaceous.

0000 0 N l A After Credner. After C. W. Andrews.

FIG. I. - A, Palate of Palaeozoic Amphibian (Archegosaurus decheni). B, Palate of Mesozoic Reptile (Plesiosaurus macrocephalus). b.occ, basioccipital; bs, basisphenoid; eept, ectopterygoid; i. pt, interpterygoid vacuity; j, j ugal; mx, maxilla; pas, parasphenoid; pl, palatine; pmx, premaxilla; pt, pterygoid; pt. nar, posterior p ares; qu, quadrate; s.o, suborbital vacuity; v, vomer.

Bible Encyclopedias
Reptiles

1911 Encyclopedia Britannica

5. Period of the Recognition of a Class of Reptilia as Part of the Sauropsida

6. Period of the Consideration of Skeletons of Extinct Reptiles

8

Crocodiles

AuTHORITIES

Chelonians

The Fore Limbs

The Pelvic Girdle

The sternum and further modifications of the ribs of the trunk.

2. Taste

3. Nose

Ear.

Special Modifications of the Lids

Tortoises

Lizards

Snakes

Tortoises.

Tongue

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