B 3 OF LIFE OLOGICAL TIME. W. DAWSON, LL.D. ERS TH] OF THE UNIVERSITY OF CALIFORNIA PRESENTED BY PROF. CHARLES A. KOFOID AND MRS. PRUDENCE W. KOFOID EARTH SCIENCES LIBRARY THE CHAIN OF LIFE GEOLOGICAL TIME A SKETCH OF THE ORIGIN AND SUCCESSION OF ANIMALS AND PLANTS BY SIR J. WILLIAM DAWS ON C.M.G., LL.D., F.R.S., F.G.S., ETC. AUTHOR OF "ACADIAN GEOLOGY," "THE STORY OF THE EARTH," "EGYPT AND SYRIA; THEIR PHYSICAL FEATURES IN RELATION TO BIBLE HISTORY," ETC. THIRD AND REVISED EDITION WITH NUMEROUS ILLUSTRATIONS THE RELIGIOUS TRACT SOCIETY, 56 PATERNOSTER Row; 65 ST. PAUL'S CHURCHYARD, AND 164 PICCADILLY 1888 :. RICHARD CLAY AND SONS, LONDON AND BUNGAV. EARTH SCIENCES LIBRARY PREFACE. QUESTIONS as to the origin and history of life are not at the present time answered by mere philosophical speculation and poetical imagining. Such solutions of these questions as science can profess to have obtained are based on vast accu- mulations of facts respecting the remains of animals and plants preserved in the rocky beds of the earth's crust, which have been successively accumulated in the course of its long geological history. These facts undoubtedly afford the means of attaining to very certain conclusions on many points relat- ing to the history of life on the earth. But, on the other hand, they have furnished the material for hypotheses which, though confidently affirmed to be indisputable, have no real founda- tion in nature, and are indirectly subversive of some of the most sacred beliefs of mankind. In these circumstances it is most desirable that those who are not specialists in such matters should be in a position to judge for themselves; and it does not appear impossible in the actual state of knowledge, to present, in terms intel- ligible to the general reader, such a view of the ascertained sequence of the forms of life as may serve at once to give vi PREFACE, exalted and elevating views of the great plan of creation, and to prevent the deceptions of pseudo-scientists from doing their evil work. Difficulties, no doubt, attend the attempt. They arise from the number and variety of the facts, from the uncertainties attending many important points, from the new views constantly opening up in the progress of discovery, and from the difficulty of presenting in an intelligible form the preliminary data in biology and geology necessary for the understanding of the questions in hand. In order, as far as possible, to obviate these difficulties, the plan adopted in this work has been to note the first known appearance of each leading type of life, and to follow its progress down to the present time or until it became extinct. This method is at least natural and historical, and has commended itself to the writer as giving a very clear comprehension of the actual state of our knowledge, and as presenting some aspects of the subject which may be novel and suggestive even to those who have studied it most deeply. In selecting examples and illustrations, the writer has en- deavoured to avoid, as far as possible, those already familiar to the general reader. He has carefully sought for the latest facts, while rejecting as unproved many things that are con- fidently asserted ; and has endeavoured to avoid all that is irrelevant to the subject in hand, and to abstain from all technical terms not absolutely essential. In a work at once so wide in its scope, so popular in its character, and so limited in its dimensions, a certain amount of hostile criticism on the part of specialists is to be expected, some portion of it per- haps just, other portions arising from narrow prejudices due to limited lines of study. The writer is willing to receive PREFACE. vii such comments with attention and gratitude, but he would deprecate the misuse of them in the interest of those coteries which are at present engaged in the effort to torture nature into a confession of belief in the doctrines of a materialistic or agnostic philosophy. The title of the work was suggested by that of Gaudry's recent attractive book, Les Enchainements du Monde animal. It seemed well fitted to express the connection and succession of forms of life, without implying their derivation from one another, while it reminds us that nature is not a fortuitously tangled skein, and that the links which connect man himself with the lowest and oldest creatures bind him also to the throne of the Eternal. In the few years that have elapsed since the publication of the first edition of this work, great additions have been made to our knowledge of fossil animals and plants. Many new species have been described, and many new facts have been discovered, respecting species previously known. This rapid progress of discovery has, however, invalidated few of the statements made in the first edition, and has certainly estab- lished nothing against the general laws of the succession of life as stated in this work. Perhaps the most interesting phase of recent discovery is the tracing back of certain forms of life to earlier periods of the earth's geological history. Some of the most recent facts of this kind are the finding, by M. Charles Brongniart, of a fossil insect, allied to the Blattae or cockroaches, in the Silurian of Spain, that of true Scorpions in the Upper Silurian of Sweden by Lindstrom, and in the Upper Silurian of Scotland by Peach, who has also described fossil Millipedes from the Lower viii PREFACE. Devonian. The tendency of such discoveries is to carry farther back the origin of highly specialised forms of life, and thus to 'render less probable their origin by any process of gradual derivation. Other discoveries serve to fill up blanks in our knowledge, and thus to render the geological record less imperfect. Of this kind is the close approximation now worked out in Western America between the end of the reign of the great Mesozoic reptiles and the beginning of that of the mammals of the Tertiary a great and abrupt revolution, effected apparently by a- coup de main. I have myself had opportunity to show that a similarly sharp line separates that quaint old Mesozoic flora of pines, cycads and ferns, which extends upward into the Lower Cretaceous, from the rich and luxuriant assemblage of broad-leaved trees of modern aspect, which takes its place in the middle part of the same formation. It is not too much to say that these and similar discoveries, while they serve to bridge over gaps in the succession of organic beings, do not favour the theory of slow modification of types. They rather point to a law of rapid development of new forms under special conditions as yet unknown to science, and this accompanied with the extinction of older species. Recent discoveries also present many remarkable instances of the early introduction of highly specialised types, of higher forms preceding those that are lower in the same class, and of the persistence of certain types throughout geological time without any important change. J. W. D. McGiLL COLLEGE. CONTENTS. CHAP. PAGE I. PRELIMINARY CONSIDERATIONS AS TO THE EXTENT AND SOURCES OF OUR KNOWLEDGE i II. THE BEGINNING OF LIFE ON THE EARTH 21 III. THE AGE OF INVERTEBRATES OF THE SEA 45 IV. THE ORIGIN OF PLANT LIFE ON THE LAND 89 V. THE APPEARANCE OF VERTEBRATE ANIMALS . . . . '. 117 VI. THE FIRST AIR-BREATHERS 137 VII. THE EMPIRE OF THE GREAT REPTILES 165 VIII. THE FIRST FORESTS OF MODERN TYPE 185 IX. THE REIGN OF MAMMALS 207 X. THE ADVENT OF MAN 233 XL REVIEW OF THE HISTORY OF LIFE 253 LIST OF ILLUSTRATIONS. FRONTISPIECE. Life in the Silurian Age .... To face Title. FIG. PAGE 1. Bank of stream or coast, showing stratification ... 4 2. Section at Niagara Falls 4 3. Section obtained by boring, near Goderich, Ontario . . 5 4. Inclined beds, holding fossil plants 6 5. Ideal section of the Apalachian Mountains .... 7 6. Generalised section across England from Menai Straits to the Valley of the Thames 9 7. Generalised section from the Laurentian of Canada to the coal-field of Michigan 9 8. Unconformable superposition of Devonian Conglomerate on Silurian slates, at St. Abb's Head, Berwickshire . . 10 9. Section of Trenton limestone, Montreal 14 10. Diagram showing different state of fossilisation of a cell of a Tabulate Coral 15 11. Cast of erect tree (Sigillaria) in Sandstone 16 12. Protichnites septem-notattts . . . . . . . .17 I2a. Footprints of modem Limulus, or king-crab . . . .18 13. Current markings on shale, resembling a fossil plant . . 1 8 Frontispiece. Magnified and restored section of a portion of Eozoon canadense .......... 20 14. Ideal section, showing the relations of the Laurentian and Huronian ........... 24 15. Small weathered specimen of Eozoon 23 1 6. Nature-printed specimen of Eozoon slightly etched with acid 29 17. Magnified group of canals in supplemental skeleton of Eo- zoon 31 18. Portion of Eozoon magnified 100 diameters . . . .31 19. Magnified portion of shell of Calcarina 32 20. Amceba, a fresh- water naked Rhizopod ; and Actinophrys, a fresh-water Protozoon ........ 34 21. Nonionina t a modern marine Foraminifer .... 34 LIST OF ILLUSTRATIONS. xi FIG. PAGE 22. Stromatopora concenirica 35 23. Caunopora planulata 36 24. Archoeocyathus minganensis. A Primordial Protozoon . . 37 25. Receptaculites. Restored 38 26. Section of Loftusia Persica. An Eocene Foraminifer . . 39 27. Foraminiferal Rock Builders, in the Cretaceous and Eocene 41 Frontispiece. Paradoxides Regina (Matthew) .... 44 28. Group of Cambrian Animals 46 29. Portion of skeleton of Hexactinellid Sponge (Cceloptychium) 49 30. Protospongia fenestrata (Salter) 50 31. Astylospongia prczmorsa (Koemer) 5 1 32. Spicules of Lithistid Sponge (Trichospongia, Billings) . . 51 33. Gldhamia antiqua (Forbes) . . . . . . . 5 2 34. Dictyonema sociale. Enlarged 52 35. Dictyonema Websteri (Dn.) 53 36. Group of modern Hydroids allied to Graptolites ... 54 37. Silurian Graptolitidse 55 38. Central portion of Graptolite, with membrane, or float (Di- ehograpsus octobrachiatus, Hall) 55 39. Ptilodiclya acuta (Hall). Bryozoan 55 390. Fenestella Lye Hi (Dn.). A Carboniferous Bryozoan . . 56 40. Chaetetes fibrosa. A Tabulate Coral with microscopic cells . 56 41. a, Stenopora exilis (Dn.). b, Chaetetes tumidus (Edwards and Haine) 57 42. Living Anthozoan Coral (Astrcea) 58 43. Tabulate Corals (Halisites and Favosites] 59 44. Rugose Coral (Heliophyllum IJalli) 59 440. Zaphrentis prolifica (Billings) 60 45. Rugose Corals (Zaphrentis Minas, Dn., and Cyathophyllum Billingsi, Dn.) 60 46. Modern Crinoid (Rhizocrinus Lofotensis) 6 1 47. Palaaster Niagarensis (Hall) 62 48. Palachinus ellipticus (McCoy) 62 49. Pleurocyslites squamosus . . -63 50. Heterocrinus simplex (Meek) 63 51. Body of Glyptocrinus 63 52. Extracrinus Briar eus ......... 64 53. Pentacrimis caput-mednsa 64 54. Lingula anatina 65 55. Cambrian and Silurian Lingulse 65 56. Terebratula s ace u IMS (Martin) 66 57. Brachiopods ; genus Orthis 66 58. Rhynchonella increbrescens (Hall) 66 59. Spirifer nnicronatus (Conrad) 67 59. Athyris subtilita (Hall) 67 60. Protfuctus cora (D'Orbigny) ........ 68 61. Group of Older Palaeozoic Lamellibranchs .... 69 62. Conularia planicstata (Dn.). A Carboniferous Pteropod . 70 63. Silurian Sea-snails 70 xii LIST OF ILLUSTRATIONS. FIG. PAGE 64. Squid (Loligo) 72 65. Pearly Nautilus (Nautilus pompilius} . . . . . .72 66. Orthoceras ............ 73 67. Gomphoceras 73 68. Lituites 73 69. Nautilus Avonensis (Dn.) 74 70. Goniatites crenistria (Philips) ....... 74 71. Ceratites nodosus (Schloth) 75 72. Ammonites Jason (Reinecke) 76 J2a. Suture of Ammonites componens (Meek) 76 73. Cretaceous Ammonitidae 77 74. Belemnite 78 74#. Belemnotetithis antiquus ......... 78 75. Cambrian Trilobites 79 76. Transverse section of Calymene. A Silurian Trilobite . . 80 760. Burrows of Trilobite and of modern King-crab . . .81 77. Silurian Trilobites 82 78. Devonian and Carboniferous Trilobites 83 79. Palaeozoic Ostracod Crustaceans 83 80. Pterygotus anglicus 84 81. Amphipeltis paradoxus (Salter) .85 82. Anthropalamon Hilliana (Dn.) . .... 85 Frontispiece. Cordaites, of the group of Dory-Cordaites . 88 83. Protannularia Harknesni (Nicholson) 91 84. American Lower Silurian Plants 92 86. Fragment of outer surface of Glyptodendron of Claypole . 93 87. Psilophyton princeps (Dn.) 95 88. Trunk of a Devonian Tree-fern ( Caulopteris Lockwoodi, Dn.) 97 89. Frond of ArckceopfeHs Jacksoni (Dn.) 98 90. Portion of a branch of Leptophleum rhombicum (Dn.) . . 98 91. Calamites radiatus (Brongniart) 99 92. A Devonian Taxine Conifer (Dadoxylon ottangondianum, Dn.) IOD 93. Group of Devonian fruity &c 101 94. Structures of the oldest-known Angiospermous Exogen (Sy- ringe Jiy Ion mirabile, Dn.) . . . . . . . .102 95. Asterophyllites parvula (Dn.) and Sphenophyllum antiquum (Dn.) 103 96. Calamites 104 97. Carboniferous Ferns 105 98. Carboniferous Tree-ferns 107 99. Lepidodendron cori-ugatum (Dn.) ....... 108 100. Sigillaricz of the Carboniferous 109 101. Trigonocarpum Hookeri (Dn. ) . . . . . . in Frontispiece. Pteraspis. Restored 116 102. Siluro-Cambrian Conodonts .118 103. Lower Carboniferous Conodont 119 104. a, Head-shield of an Upper Silurian Fish (Cy at has pis}; b, Spine of a Silurian Shark (Onchus tenui-striatus, Agass.) ; c, d, Scales of Thecodus , 121 LIST OF ILLUSTRATIONS. xiii FIG. PAGE 105. Cephalaspis Da-wsoni (Lankester) 122 106. Devonian Placoganoid Fishes (Pterichthys cornutus, Cephal- aspis Lyelli) 123 107. Devonian Lepidoganoid Fishes (Diplacanthus and Osteolepis) 124 108. Modern Dipnoi (Ceratodus Fosteri -. and Lepidosiren annectus) , ....... 124 109. Anterior part of the palate of Dipterus ..... 125 no. Dental plate of Conchodus plicatus (Dn.) . . . . 126 111. Dental plate of Ceratodus Barrandii . . . . . . 126 112. Dental plate of Ceratodus serratus 127 113. Jaws of Dinichthys Hertzeri (Newbeiry) 127 114. Lower Jaw of Dinichthys Hertzeri 128 115. Jaws of ' Lepidosiren 128 1 16. A small Carboniferous Ganoid (Palaoniscus (Rhadinichthys) Modulus, Dn.) 129 117. Teeth and Spines of Carboniferous Sharks .... 130 118. Teeth of Cretaceous Sharks (Otodus and Ptychodus} . . 131 119. Tooth of a Tertiary Shark (Carcharodon) 132 1 20. A Liassic Ganoid (Dapedius} 132 121. Cretaceous Fishes of the modern or Teleostian type (Beryx Lerwesiensis and Portheus molossus, Cope) . . . . 133 122. Modern Ganoids (Polypterus and Lepidosteus} . . . .134 Frontispiece. A Microsaurian of the Carboniferous Period (Hylonomus Lyellt) . . . f 136 123. Wings of Devonian Insects 140 124. Land-snail (Pupa vetusta, Dn.) 143 125. Land-snail (Zonites (Conulus] priscus, Carpenter) . . . 143 126. Millipedes (Xylobius sigillaria, Dn. ; Archiulus Xylobioides, Scudder ; X. farctus, Scudder) 145 127. Wings of Cockroaches 146 128. Wing of May-fly (Haplophlebium Barnesii, Scudder) . . 147 129. A Jurassic Sphinx-moth (Sphinx Snelleri, Weyenburgh) . 148 130. An Eocene Butterfly (Prodryas persephone, Scudder) . . 149 131. Abdominal part of a Carboniferous Scorpion . . . .150 132. Carboniferous Scorpion (Eoscorpius carbonarius, Meek and Worthen) 151 133. Footprints of one of the oldest known Batrachians, probably a species of Dendrerpeton . 152 134. Archegosaurus Decheni 154 135. Ptyonius 154 136. A large Carboniferous Labyrinthodont (Baphetes planiceps, Owen) , 155 137. Baphetes planiceps (Owen) 156 138. A lizard-like Amphibian (Hylonomus aciedentatus) . .157 139. Stelliosaums longicostatus (Fritsch) .158 140. Section showing the position of an erect Sigillaria, contain- ing remains of land animals 1 60 1400. Section of base of erect Sigillaria, containing remains of land animals . 161 xiv LIST OF ILLUSTRATIONS. Frontispiece. Inhabitants of the English Seas in the Age of Reptiles 164 141. Arm of Proterosaurus Speneri 166 142. Skeleton of Ichthyosaurus 167 142^. Head of Pliosaurtis . . 168 142^. Paddle of Plesiosaurus Oxoniensis 168 143. Skeleton of Clidastes 170 144. An Anomodont Reptile of the Trias (Dicynodon lacerticeps, Owen) 170 145. A Theriodont Reptile of the Trias (Lycosaurus} . . .170 146. Skeleton of Pterodochylus crassirostris 170 147. Restoration of Rhamphorhyncus Bucklandi . . . .171 148. A Jurassic bird (Archaopteryx macroura) 172 149. Jaw of a Cretaceous Toothed Bird (Ichthyornis dispar) . 173 150. Jaw of Bathygnathus borealis (Leidy) 174 151. Hadrosaurus Foulkii (Cope) 175 152. Jaws of Megalosaurus 176 153. Tooth of Megalosaurus 177 154. Compsognathus 179 Frontispiece. Lower Cretaceous Leaves 184 155. Sassafras cretaceum (Newberry) 190 156. Liriodendron pnmcevum (Newberry) . . . . . .191 157. Onoclea sensibilis \ .191 158. Davallia tenuifolia . .192 159. Eocene Leaves 194 160. An Ancient Clover (Trifolium palceogceum, Saporta) . . 195 161. An Eocene Maple (Acer sextiamts, Saporta) . . . .195 162. A European Magnolia of the Eocene (M. diance, Saporta) . 195 163. Flower and Leaf of Bombax sepultijlorum .... 196 164. Branch and Fruit of Sequoia Couttsice (Heer) .... 197 165. Cinnamomum Schetichzeri (Heer) 198 Frontispiece. Sivatherium giganteum ...... 206 1 66. Jaw of Dromatherium sylvestre (Emmons) .... 209 167. Myrmecobius Jasciatus ......... 209 1 68. Jaw and Molar of Phascolotherium Bucklandi . . .210 169. Jaw and Pre-molar of Plagiaulax Becklesii . . . .210 170. Restoration of Palceotherium magnum 21 1 171. Skull of a Lower Eocene Perissodactyl (Coryphodon Hamatus) 214 172. Fore-foot of Coryphodon 215 173. Skull of Upper Eocene Perissodactyl (Dinoceras mirabilis} . 216 174. Fore-foot of Dinoceras 217 175. Skull of Miocene Perissodactyl (Brontotherium ingens, Marsh) 217 176. Series of Equine feet 218 177. Skull of generalised Miocene Ruminant (Oreodon major) . 221 178. Lower Jaw of Megatherium 222 179. Ungual Phalanx and Claw-core of Megatherium . . . 222 1 80. Tooth of Eocene Whale (Zettglodon cetioides) .... 223 181. Mastodon ohioticus 225 182. Head of Dinotherium giganteum 226 LIST OF ILLUSTRATIONS. xv FIG. j-AGE 183. Wing of Eocene Bat ( Vesfertilio aquensis) . . . 226 184. Skull of a Cymetar-toothed Tiger (Machairodus cultridens) . 228 185. Lower Jaw of Diyopithecus Fontani 229 Frontispiece. Contemporaries of Post-Glacial Man . . 232 1 86. Klephas primigenius 241 187. Tooth of Elasmotheritim 242 188. Engis Skull 243 189. Outlines of Three Prehistoric European Skulls compared with an American Skull 244 190. Flint Implement found in Kent's Cavern, Torquay . 245 191. Bone Harpoon (Palseocosmic) 246 192. Sketch of a Mammoth carved on a portion of a Tusk of the same Animal 249 TABULAR VIEW OF GEOLOGICAL PERIODS AND OF LIFE-EPOCHS. GEOLOGICAL PERIODS. ANIMAL LIFE. VEGETABLE LIFE. [ ) Modern. Age of Man Quaternary. (Post-Glacial. CAINOZOIC and modern Mammals. Age of or { /Pleistocene or NEOZOIC. ] 1 Glacial. Tertiary . . I Pliocene. 1 Miocene. Age of Extinct Mammals. (Earliest Placental Angiosperms and Palms. k \Eocene. Mammals.) f i Upper, Cretaceous . < Lower, or (Neocomian. Age of Reptiles (Earliest TW/zcwV f Oolite. MES070IC. \s uf ' \Lias. and Birds. Modern Trees.) Age of \ T - ( Middle, or ' Muschelkalk. \ VLower. (Earliest Marsupial Mammals.) Cycads and Pines. Permian . . l M ' agn ^ L ; mes tone. (Earliest \ Lower. true Reptiles.) (Upper Coal-Formation. Carboniferous 1 Coal-Formation _ I Carboniferous Limestone. VLower Coal-Formation. (Upper. PALEOZOIC. *>****> (j^f Age of A mphibians and Fishes. Age of Acrogens and Gymnosperms. (Earliest Land Plants.) Silurian . . {L^' Age of A Igte. Siluro-Cam- | Upper Age of Ordovician. ' ower< Mollusks, Corals, and TUpper. Cambrian . < Middle. VLower. Crustaceans. ^Z. Age of Protozoa. (First animal remains.) Indicatiins of Plants notdeterminable. vBojian. THE CHAIN OF LIFE. CHAPTER I. PRELIMINARY CONSIDERATIONS AS TO THE EXTENT AND SOURCES OF OUR KNOWLEDGE. IT is of the nature of true science to take nothing on trust or on authority. Every fact must be established by accurate observation, experiment, or calculation. Every law and prin- ciple must rest on inductive argument. The apostolic motto, " Prove all things, hold fast that which is good," is thoroughly scientific. It is true that the mere reader of popular science must often be content to take that on testimony which he cannot personally verify ; but it is desirable that even the most cursory reader should fully comprehend the modes in which facts are ascertained and the reasons on which conclusions are based. Failing this, he loses all the benefit of his reading in so far as training is concerned, and cannot have full assurance of that which he believes. When, therefore, we speak of life- epochs, or of links in a chain of living beings, the question is at once raised What evidence have we of the succession of such epochs? This question, with some accessory points, must engage our attention in the present chapter. B 2 THE CHAIN OF LIFE. Geology as a practical science consists of three leading parts. The first and most elementary of these is the study of the dif- ferent kinds of rocks which enter into the composition of those parts of the earth which are accessible to us, and which we are in the habit of calling the crust of the earth. This is the sub- ject of Lithology> which is based on the knowledge of minerals, and has recently become a much more precise department of science than heretofore, owing to the successful employment of the microscope in the investigation of the minute structure and composition of rocks. The second is the study of the arrange- ment of the materials of the earth on the large scale, as beds, veins, and irregular masses ; and inasmuch as the greater part of the rocks known to us in the earth's crust are arranged in beds or strata, this department may be named Stratigraphy. A more general name sometimes employed is that of Petro- graphy. The third division of geology relates to the remains of animals and plants buried in the rocks of the earth, and which have lived at the time when those rocks were in process of formation. These fossil remains introduce us to the history of life on the earth, and constitute the subject of Paleontology. It is plain that in considering what may be learned as to epochs in the history of life we are chiefly concerned with the last of these divisions. The second may also be important as a means of determining the relative ages of the fossils. With the first we have comparatively little to do. Previous to observation and inquiry, we might suppose that the kinds of animals and plants which now inhabit the earth are those which have always peopled it ; but a very little study of fossils suffices to convince us that vast numbers of creatures once inhabitants of this world have become extinct, and can be known to us only by their remains buried in the earth. When we place this in connection with stratigraphical facts, we further find that these extinct species have succeeded each other at different times, so as to constitute successive dynasties PRELIMINARY CONSIDERATIONS. 3 of life. On the one hand, when we know the successive ages of fossil forms, these become to us, like medals or coins to the historian, evidences of periods in the earth's history. On the other hand, we are obliged in the first instance to ascertain the ages of the medals themselves by their position in the succes- sive strata which have been accumulated on the surface. The series of layers which explorers like Schliemann find on the site of an ancient city, and which hold the works of successive peoples who have inhabited the place, thus present on a small scale a faithful picture of the succession of beds and of forms of life on the great earth itself. Our leading criterion for estimating the relative ages of rocks is the superposition of their beds on each other. The beds of sandstone, shale, limestone, and other rocks which constitute the earth's crust have nearly all been deposited thereon by water, and originally in attitudes approaching to horizontality. Hence the bed that is the lower is the older of any two beds. Hence also, when any cutting or section reveals to us the succession of several beds, we know that fossil remains contained in the lower beds must be of older date. We can scarcely walk by the side of a stream which has been cutting into its banks, or at the foot of a sea-cliff, or through a road-cutting, without observing illustrations of this. For instance, in the section represented in Fig. i, we see at the surface the vegetable soil, below this layers of gravel and sand, below this a bed of clay, and below this hard limestone. Of these beds a is the newest, d the oldest ; and if, for example, we should find some marine shells in d, some fresh- water shells in c, bones of land animals and flint arrow- heads in b, and fragments of modern pottery in a, we should be able at once to assign their relative ages to these fossils, and to form some idea of the succession of conditions and cf life which had occurred in the locality. On a somewhat larger scale, we have in Fig. 2 a section of the beds cut through by the great Fall of Niagara. All of B 2 4 THE CHAIN OF LIFE. these except that marked a are very ancient marine rocks, holding fossil shells and corals, but now forming part of the interior of a continent, and cut through by a fresh-water river. FIG. i. Bank of stream or coast, showing stratification. a, Vegetable soil. t>, Gravel and sand, c, Clays, d, Limestone rock, slightly inclined. In deep mines and borings still more profound sections may be laid open, as in Fig. 3, which represents the sequence of beds ascertained by boring with the diamond drill in search FIG. 2. Section at Niagara Falls, showing the Fall. Thickness of beds ta^cut through by the action of the u725o feet. a, Boulder clay and gravel Post-pliocene. d, Clinton limestone j e, Medina sandstone / marine shells and corals. of rock salt near Goderich in Canada. Here we have a suc- cession of 1,500 feet of beds, some of which must have been formed under very peculiar and exceptional conditions. The beds of rock salt and gypsum must have been formed by the drying up of sea-water in limited basins. Those of Dolomite PRELIMINARY CONSIDERATIONS. 5 imply precipitation of carbonate of lime and magnesia in the sea-bottom. The marls must have been formed largely by the driftage of sand and clay, while some of the limestone was 78' 9" 278' 3" 276" of I'o" I 3 2'o" Total 1,517 ft. FIG. 3. Section obtained by boring with the diamond drill, near Goderich, Ontario, Canada, in the Salina series of the Upper Silurian. From a memoir by Dr. Hunt in the Report of the Geological Survey of Canada for 1876-7. No. i, Clay, gravel, and boulders Post-pliocene. Nos. 2, 4, 7, 9, 13, Dolomite or magnesian limestone, with layers of marl, limestone, and gypsum. No. 3, Lime- stone with corals Favosites, etc. Nos 5, n, 15, 17, Ma'Hs with layers of Dolomite and anhydrous gypsum. Nos. 6, 8, 10, 12, 14, 16, Rock salt. 6 THE CHAIN OF LIFE. produced by accumulation of corals and shells. Such deposits must not only have been successive, but must have required a long time for their formation. In Fig. 4 we have a bed of coal and its accompaniments. The coal itself was produced by the slow accumulation ot vegetable matter on a water-soaked soil, and this was buried FIG. 4. Inclined beds, holding fossil plants. Carboniferous. South Joggins, Nova Scotia. i. Shale and sandstone. Plants with Spirorbis attached ; rain marks (?). le, 8 feet. Erect Calamites. ) An erect coniferous (?) tree, rooted on the shale, passes up through 15 feet of the sandstones and shale. 2. Sandstone and shal 3. Gray sandstone, 7 feet. 4. Gray shale, 4 feet. 5. Gray sandstone, 4 feet. 6. Gray shale, 6 inches. Prostrate and erect trees, with rootlets, leaves, Naiadites, and Spirorbis on the plants. 7. Main coal-seam, 5 feet coal in two beds. 8. Underclay, with rootlets. under successive beds of sand and clay, now hardened into sandstone and shale, some of the beds holding trees and reed- like plants, which still stand on the soils on which they grew, and which must have been buried in sediment deposited in inundations or after subsidence of the land. In this section we may also observe that the beds are somewhat inclined ; PRELIMINARY CONSIDERATIONS. 7 and that this is not their original position is shown by the posture of the stems of trees, once erect, but now inclined with the beds. This leads to a consideration very important with reference to our present subject ; namely, that as our con- tinents are mostly made up of beds deposited under water and afterwards elevated, these beds have in this process experienced such disturbances that they rarely retain their horizontal posi- tion, but are tilted at various angles. When we follow such inclined strata over large areas, we find that they undulate in great waves or folds, forming what are called anticlinal and synclinal lines, and that the irregularities of the surface of the land depend to a great extent on these undulations, along with the projection of hard beds whose edges protrude at the sur- face. In point of fact, as shown in Fig. 5, mountain ranges depend on these crumplings of the earth's crust; and the FIG. 5. Ideal section of the Apalachian Mountains showing folding of the earth's crust. a, Anticlinal axes, b, Overturned strata, c, Synclinals, d, Unconformable beds. primary cause of these is probably the shrinkage of the mass of the earth owing to contraction in cooling. When the dis- turbances of beds are extreme, they often cause intricacies of structure difficult to unravel ; but when of moderate extent they very much aid us in penetrating below the surface, for we can often see a great thickness of beds rising one from beneath another, and can thus know by mere superficial ex- amination the structure of the earth to a great depth. It thus happens that geologists reckon the thickness of the stratified 8 THE CHAIN OF LIFE. deposits of the crust of the earth at more than 70,000 feet, though they cannot penetrate it perpendicularly to more than a fraction of that depth. The two sections, Figs. 6 and 7, showing the sequence of beds in England and in the northern part of North America, will serve, if studied by the reader, to show how, by merely travelling over the surface and measuring the upturned edges of beds, many thousands of feet of deposits may be observed, and their relative ages distinctly ascertained. In studying any extensive section of rock we find that its members may more or less readily be separated into distinct groups. Sometimes these are distinguished by what is termed unconformability, that is, the lower series has been disturbed or inclined before the upper has been deposited upon it. This is seen on a grand scale in the section Fig. 7, in the case of the Laurentian and Cambrian formations, and on a smaller scale in Fig. 8 in the unconformable superposition of Devonian conglomerate on Silurian slates at St. Abb's Head. In the last section it is quite evident that the beds of the lower series have been bent into abrupt folds and worn away to a con- siderable extent before the deposition of the overlying series. In such a case we know not merely that the upper series is newer than the lower, but that some considerable time must have elapsed after the deposition of the one before the other was laid down ; and we are not surprised to find that the fossils in the groups thus unconformable to each other are very different. But even when the beds are conformable, they can usually be separated into groups, depending upon differences of mineral character, or changes which have occurred in the mode of deposition. One group of beds, for example, may be largely composed of limestone, another of sandstone or shale. One group may be distinguished by containing some special mineral, zs, for example, rock salt or coal, while others may be destitute of such special minerals. One group may show by its fossils that it was deposited in the sea, others may be estuarine or PRELIMINARY CONSIDERATIONS. 5: ;3- io THE CHAIN OF LIFE. lacustrine. Thus we obtain the means of dividing the rocks of the earth into groups of different ages, known as " Forma- tions," and marking particular periods of geological time. By tracing these formations from one district or region to another, we learn the further truth that the succession is not merely local, but that, though liable to variation in detail, its larger subdivisions hold so extensively that they may be regarded as world-wide in their distribution. Putting together the facts thus obtained, we can frame a tabular arrangement of the earth's strata, as in the table pre- fixed to this chapter ; and when we add the further discovery, very early made by geologists, that the successive formations FiG. 8. Unconfjrmable superposition of Devonian conglomerate on Silurian slates, a St. Abb's Head, Berwickshire. After Lyell. differ from each other in their fossil remains, we have the means of recognising any particular formation by its fossils, even when the stratigraphical evidence may be obscure or wanting. Thus our knowledge of Epochs of Life, and indeed of the whole geological history of the earth, is based on the superposition of beds in the earth's crust, and on the diversity of fossil re- mains in the successive beds so superimposed on each other ; and it is on these grounds that we are enabled to construct a Table of Geological Formations representing the whole series of beds as far as known, with the characteristic groups of fossils of each period. Here I might close these preliminary con- siderations, but there are a few accessory questions, important to our clear comprehension of the subject, which may profitably occupy our attention for a short time. PRELIMINARY CONSIDERATIONS. n One of these relates to the absolute duration of the time represented by the geological history of the earth. Such estimates as our present knowledge enables us to form are very indefinite. Whether we seek for astronomical or geo- logical data, we find great uncertainty. To such an extent is this the case, that current estimates of the time necessary to bring the earth from a state of primitive incandescence to its present condition have varied from fifteen millions of years to five hundred millions. Of the various modes proposed, perhaps the most satisfactory as well as instructive is that based on the rate of denudation of our present continents, as indicated by the amount of sediment carried down by great rivers. The Mississippi, draining a vast and varied area in temperate latitudes, is washing away the American land at the rate of one foot in 6,000 years. The Ganges, in a tropical climate and draining many mountain valleys, works at the rate of one foot in 2,358 years. The mean of these two great rivers would give one foot in 4,179 years, at which rate our continents would be levelled with the waters in about six millions of years. But the land has been in process of renewal as well as of waste in geological time ; and a better measure will be afforded by the amount of beds actually deposited. The entire thickness of all the stratified rocks of Great Britain has been calculated by Ramsay at 72,000 feet. Now, if we suppose the waste in all geological time to have been on the average the same as at present, and that this material has been deposited to the thickness of 72,000 feet on a belt of sea margin 100 miles in width, we shall have about 86 millions of years as the time required. 1 This has the merit of approxi- mating to Sir William Thomson's calculation, based on the rate of cooling of the earth, that a minimum of 100 millions of years may represent the time since a solid crust first began to form. As it is more likely that the rate of denudation has on 1 Croll has elaborated this calculation in his work, Climate and Time, 12 THE CHAIN OF LIFE. the average been greater in former geological periods than at present, we may perhaps estimate fifty or sixty millions of years as the time required for the accumulation of all our formations. Some geologists object to this as too little, but in this some of them are influenced by the exigencies of theories of evolution, and others appear to have no adequate conception of the vast lapse of time represented by such numbers, in its relation to the actual rates of denudation and deposition. It should be mentioned here, however, that, on certain theories now somewhat generally accepted, respecting the nature and source of solar heat, the absolute duration of geological time would be much reduced below the estimate of Sir Wm. Thomson, Prof. Tait has based on such data an estimate of fifteen millions of years. Prof. Simon Newcomb says that " on the only hypothesis science will now allow us to make respecting the source of the solar heat " (the gravitation hypothesis of Helmholtz) " the earth was, twenty millions of years ago, enveloped in the fiery atmosphere of the sun." Dr. Kirkwood has called attention to these results in con- nection with the planetary hypothesis of La Place, in the Proceedings of the American Philosophical Society^ Should such views prove to be well-founded, geological calculations as to the time required for the successive formations may have to be revised. If now we attempt to divide this time among the formations known to us, according to their relative thicknesses, we have, according to an elaborate estimate of Professor Dana, the time ratios of 12, 3, and i for the Palaeozoic, Mesozoic, and Cainozoic periods respectively. Taking the whole time since the begin- ning of the Cambrian as forty-eight millions of years, we should thus have for the Palaeozoic thirty-six millions, for the Mesozoic nine, and for the Tertiary three. Another calculation, recently 1 Sept. 1879. PRELIMINARY CONSIDERATIONS. 13 made by Professors Hull and Haughton, gives the following ratios : Azoic ... . . . 34 '3 per cent. Palaeozoic ....... 42*5 ,, Mesozoic and Cainozoic . 23*2 ,, This calculation is, however, based on the absolute thickness of the several series as ascertained in Great Britain, without reference to the nature of the beds, as indicating different rates of accumulation. Under either estimate it will be seen that the Palaeozoic time greatly exceeds the Mesozoic and Caino- zoic together, and consequently that changes of life seem to have proceeded at an accelerated rate as time wore on. Another inquiry of some importance relates to the manner of preservation of fossils, and the extent to which they consti- tute the material of rocks. This inquiry is doubly important, as it bears on the genuineness of fossil remains, and on the means we have of understanding their nature. Some rocks are entirely made up of matter that once was alive, or formed part of living organisms. This is the case with some limestones, which consist of microscopic shells, or of larger shells, corals, and similar calcareous organisms, either entire or broken into fragments and cemented together with pasty or crystalline limestone filling their interstices. This may be seen in Fig. 9, which represents a magnified slice of a Silurian limestone. Coal in like manner consists of carbonised vegetable matter, retaining more or less perfectly its organic structure, and sometimes even the external forms of its consti- tuent parts. More frequently, fossils are dispersed more or less sparsely through the substance of beds composed of earthy matter ; and they have usually been more or less affected by chemical changes, or by mechanical pressure, or are mineral- ised by different substances which have either filled their pores by infiltration or have more or less completely replaced their substance. Of course, as a rule, the softer and more putrescible organic matters have perished by decay, and it is only the 14 THE CHAIN OF LIFE. harder and more resisting parts that remain. Even these have often yielded to the enormous pressure to which they have been subjected, and if at all porous, have been changed by the slow action of percolating water charged with various kinds of mineral matter in solution. It thus happens that many fossils are infiltrated with mineral matter. Wood, for example, may have the cavities of its cells FIG. 9. Section of Trenton limestone, magnified, showing that it is composed ot fragments of corals, crinoids, and shells. Montreal. and vessels filled with silica or silicates, with sulphide or car- bonate of iron, or with limestone, while the woody walls of the cells may remain either as coaly matter or charcoal. I have often seen the microscopic cells of fossil wood not only filled in this way, but presenting under a high power successive coats of deposit, like the banded structure of an agate. In some cases not only are the pores filled with mineral PRELIMINARY CONSIDERATIONS. 15 matter, but the solid parts themselves have been replaced, and the whole mass has actually become stone, while still retaining its original structure. Thus silicified wood is often as hard and solid as agate, and under the microscope we see that the wood has entirely perished, and is represented by silica or flint, dif- fering merely in colour from that which fills the cavities. In this case we may imagine the wood to have been acted on by n FIG. 10. Diagram showing different state of fossilisation of a cell of a tabulate coral (Dawson's Dawn of Life). a Natural condition, wall calcite cell empty, b Wall calcite, cells filled with the same. c Walls calcite, cells filled with silica or a silicate, d Wall silicified, cells filled with calcite. e Wall silicified, cell filled with silica. water holding in solution silica, combined with soda or potash, in the manner of what is termed soluble glass. The wood, in decay, would be converted into carbon dioxide, and this as formed would seize on the potash or soda, leaving the silica in an insoluble state, to be deposited instead of the carbon. Thus each particle of the carbon of the wood, as removed by decay, would be replaced by a particle of silica, till the whole be- came stone. By similar chemical changes corals and shells are often represented by silica, or by pyrite, which has taken the place of the original calcareous matter ; and still more remarkable changes sometimes occur, as when the siliceous spicules of sponges have been replaced by carbonate of lime. The organic matter present in the fossils greatly promotes these changes, by the substances produced in its decay, and thus it often happens that the shells, corals, etc., contained in lime- stone have been replaced by flint, while the inclosing lime- stone is unchanged. Fig. 10 shows the various conditions i6 THE CHAIN OF LIFE. which a coral may assume under these different modes of treatment. The substance of a fossil may be entirely removed by decay or solution, leaving a mere mould representing its external form, and this may subsequently be filled with mineral matter, so as to produce a natural cast of the object. This is very common in the case of fossil plants ; and large trunks of trees . may sometimes be found represented, as seen in Fig. n, by stony pillars retaining nothing of the original wood except per- haps a portion of the bark in the state of coal. It sometimes happens that the substance of fossils has been removed, leaving mere empty cavities, sometimes con- taining stony cores repre- senting the internal chambers of the fossils. Again, cal- careous fossils imbedded in hard rocks are often removed by weathering, leaving very perfect impressions of their forms. For this reason the fossil remains contained in some hard resisting rocks can be best seen as impressed moulds on the weathered surfaces. Lastly, we sometimes have impressions or footprints repre- senting the locomotion of fossil animals, rather than the fossils themselves. In this way some extinct creatures are known to us only by their footsteps on sand or clay, once soft, but now FIG. ii. Cast of erect tree (Sigillaria) in sandstone, standing on a small bed of coal, South Joggins, Nova Scotia (Daw son's Acadian Ge< PRELIMINARY CONSIDERATIONS. 17 hardened into stone ; and in the case of some of the lower animals the trails thus made are often not easily interpreted (Figs. 12, 120). It has been found that even sea-weeds drifted FIG. 12. Protichnites septem-notatus. A supposed series of crustacean foot-prints n.acL in sand, now hardened into sandstone. Cambrian. After Logan. by the tide make impressions of this kind, which, when they occur in old rocks, are very mysterious. Even rain-drops are capable of being permanently impressed on rocks, and con- stitute a kind of fossils. Besides these we have many kinds of imitative markings which simulate fossils, as those of concre- tions or nodules, which are often very fantastic in shape, those of [dendritic crystallisation giving moss-like forms, and the c iS THE CHAIN OF LIFE, complicated tracery produced on muddy shores by the little rills of water which follow the receding tide (Fig. 13). Such things are often mistaken by the ignorant for fossil remains, but are easily distinguished by a practised eye. The reader who has followed these, perhaps somewhat dry, FIG. i2, Internal cavity. with spicules ; but these I have ascertained to be merely acci- dental, and will be referred to in the next chapter. The true structure of Archseocyathus consists of radiating calcareous plates enclosing chambers connected by pores. Archaeocyathus came in with the Later Cambrian, and seems to have died out in the Siluro- Cambrian. The only more modern things which at all resemble it are the Foraminifera called Dactylopora, which belong to the Tertiary period. Receptaculites is a still more complex organism. It has a THE BEGINNING OF LIFE ON THE EARTH. 39 sack-like form, often attaining a large size, and the double walls are composed of square or rhombic plates, connected with each other by hollow tubes from which proceed canals perforating the plates (Fig. 25). This curious structure is confined to the Siluro-Cambrian, and is so dissimilar from modern forms that its affinities have been subject to grave doubts. We thus have presented to us the remarkable fact that in FIG. 26. Section of Loftitsia Persica. An Eocene Foraminifer. Magnified five diameters. After Carpenter and Brady. the Palaeozoic age we have no precise representative of Eozoon, but instead three divergent types, differing from it and from each other, all apparently specialised to particular uses, all temporary in their duration ; while in later times nature seems to have returned nearer to the type of Eozoon, though on a smaller scale, and separating some characters conjoined in it. Some portion of this curious result may be due to our ignorance ; and it would be interesting to know, what we may know some 40 THE CHAIN OF LIFE. day, how this type of life was represented in the long interval between the Huronian and the Upper Cambrian, when perhaps there may have been forms that would at least enable us to connect Eozoon and Stromatopora. Another link in the chain of being remains to be noticed here. In the Laurentian limestones we meet with numerous minute spherical bodies and groups of spheres with calcareous tubulated tests. 1 These may either be small Foraminiferae, distinct from Eozoon, or may be germs or detached cells from its surface. Similar bodies are found in the lower part of the Siluro-Cambrian, in the Quebec group at Point Levis ; and there they are filled with a species of glauconite constituting a sort of greensand rock. Still higher, in the Carboniferous, there are very numerous species of Foraminifera, presenting forms very similar to those in the modern seas, so that in the smaller shells of this group we seem to have evidence of a continuous series all the way from the Laurentian to the present time. The greater laminated forms co-exist with these up to the Eocene Tertiary. Throughout the whole of geological time from the formation of the Laurentian limestones to that of the chalky ooze accumulating in the modern ocean these humble creatures have been among the chief instruments in seizing on the calcareous matter of the waters and depositing it in the form of limestone. I have said nothing of the development of higher forms of animal life from Eozoon, simply because I know nothing of it. We shall see in the next chapter that these are introduced seemingly in an independent manner. We may be content to trace foraminiferal life along its own line of development, waxing and waning, but ever confined within the same general boundaries, from the Laurentian to the present time. It is likely that if, in any of the ages constituting this vast lapse of time, a dredge had been dropped into the depths of ocean, 1 Archaesfheriiut of the author. THE BEGINNING OF LIFE ON THE EARTH. 41 it would have brought up Foraminifera not essentially different in form and structure. If any one asks to what extent the suc- cessive species constituting this almost endless chain may be descendants one of the other, we have no absolutely certain information to give. On the one hand, it is not inconceivable thas such forms as Stromatopora or Nummulina may have de- FIG. 27. Foraminiferal Rock Builders, in the Cretaceous and Eocene. a, Nummulites Icevigata Eocene, b, The same, showing chambered interior, c, Milioline limestone, magnified Eocene, Paris, a, Hard Chalk, section magnified Cretaceous. scended from Eozoon. On the other hand, it is equally con- ceivable that the same power which produced Eozoon at first, whether from dead matter or from some unknown lower form of life, may have repeated the process in later times with modifications. In any case it is probable that the Foraminifera have experienced alternations of expansion and shrinkage, 42 THE CHAIN OF LIFE. of elevation and decadence, in the lapse of geological time. There were times in which many new forms swarmed into ex- istence, and times in which old forms were becoming extinct without being replaced by others. In so far as the areas of the continents and the adjacent waters are concerned, those periods when the land was subsiding under the ocean must have been their times of prosperity, those in which the crust of the earth shrunk and raised up large areas of land must have been their times of decay. Still this lowest form of animal life has never perished, but has always found abundant place for itself, however pressed by physical change and by the intro- duction of higher beings. Paradoxides Regina (Matthews). Lower Cambrian of New Brunswick. Jth Nat. Size. CHAPTER III. THE AGE OF INVERTEBRATES OF THH SEA. IF the middle portion of the Laurentian age was really a time of exuberant and abounding life, either this met with strange reverses in succeeding periods, or the conditions of preservation have been such as to prevent us from tracing its onward history. Certain it is, that according to present ap- pearances we have a new beginning in the Cambrian, which introduces the great Palaeozoic age, and few links of connection are known between this and the previous Eozoic. At the beginning of the Palaeozoic we have reason to believe that our continents were slowly subsiding under the sea, after a period of general continental elevation which was con- sequent on the crumbling of the earth's crust at the close of the Eozoic ; and on the new sea-bottoms formed by this subsi- dence came in, slowly at first, but in ever-increasing swarms, the abundant and varied life of the early Palaeozoic. In the oldest portion of the Cambrian series in Wales, Hicks has catalogued species of no less than seventeen genera, em- bracing Crustaceans, the representatives of our crabs and lobsters, bivalve and univalve shell-fishes of different types, worms, sea-stars, zoophytes, and sponges. If we could have walked on the shores of the old Cambrian sea, or cast our dredge or trawl into its depths, we should have found repre- sentatives of most of the humbler forms of sea life still extant, 4 6 THE CHAIN OF LIFE. though of specific forms strange to us. Perhaps the nearest approach to such experience which we can make is to examine the group of Cambrian animals delineated in Fig. 28, and to notice, under the guidance of the geologist above named, the sections seen at St. David's, in South Wales. Here we find a nucleus of ancient rocks supposed to be Laurentian, though in mineral character more nearly akin to FIG. 28. Group of Cambrian Animals (from Nicholson). a, Arenicolites didymus, worm tubes, b, Lingulella ferruginea. c, Theca Davidii. d, Modiolopsis solvensis. e, Orthis Hicksii. f, Obolella sagittalis. g, Hymen- ocaris vermicauda. h, Trilobite, Olenus micrurus. the Huronian, but which have hitherto afforded no trace of fossils. Resting unconformably on these is a series of par- tially altered rocks, regarded as Lower Cambrian, and also destitute of organic remains. These have a thickness of almost 1,000 feet, and they are succeeded by 3,000 feet more of similar rocks, still classed as Lower Cambrian, but which have afforded fossils. The lowest bed which contains indica- tions of life is a red shale, perhaps a deep-sea bed, and possibly THE AGE OF INVERTEBRATES OF THE SEA. 47 itself partly of organic origin, by that strange process of de- composition or dissolution of foraminiferal ooze and volcanic fragments, going on in the depths of the modern ocean, and described by Dr. Wyville Thomson as occurring over large areas in the South Pacific. The species are two Lingulellce, a Discina and a Leperditia. Supposing these to be all, it is remarkable that we have no Protozoa or Corals or Echinoderms, and that the types of Brachiopods and Crustaceans are of com- paratively modern affinities. Passing upward through another 1,000 feet of barren sandstone, we reach a zone in which no less than five genera of Trilobites are found, along with Ptero- pods and a sponge. Thus it is that life comes in at the base of the Cambrian in Wales, and it may be regarded as a fair specimen of the facts as they appear in the earlier fossiliferous beds succeeding the Laurentian. Taking the first of these groups of fossils, we may recognise in the Leperditia a two- valved Crustacean closely allied to forms still living in the seas and fresh waters. The Lingulellse, whether we regard them as molluscoids, or, with Professor Morse, as singularly specialised worms, represent a peculiar and distinct type, handed down, through all the vicissitudes of the geological ages, to the present day. The Pteropods and the sponge are very similar to forms now living. The Trilobites are an extinct group, but closely allied to some modern Crustaceans. Had the primordial life begun with species altogether inscrutable and unexampled in succeeding ages, this would no doubt have been mysterious ; but next to this is the mystery of the oldest forms of life being also among the newest. Whatever the origin of these creatures, they represent families which have endured till now in the struggle for existence without either elevation or degradation. Yet, though thus vast in their duration, they seem to have swarmed in together and in great numbers, in the Cambrian, without any previous preparation. From the Cambrian onward, throughout the whole Palaeozoic, there is no decided break in the continuity of marine life ; and 48 THE CHAIN OF LIFE. already in the Silurian period the sea was tenanted with all the forms of invertebrate life it yet presents, and these in a teeming abundance not surpassed in any succeeding age. Let us now, in accordance with our plan, select some of these ancient inhabitants of the waters and trace their subsequent history. Remains of sea-weeds are undoubtedly present in the Cam- brian rocks. One of the lowest beds in Sweden has been named from their abundance the Fucoidal Sandstone; and wherever fossiliferous Cambrian rocks occur, some traces, more or less obscure, of these plants may be found. Nearly all that we can say of them, however, is, that, in so far as their remains give any information, they are very like the plants of the same group that now abound in our' seas. In the fucoidal sandstone of Sweden certain striated or ribbed bodies have been found, which have even been regarded as land plants ; 1 but they seem rather to be trails or marks left by sea-weeds dragged by currents over a muddy bottom. The plants of the sea thus precede those of the land, and they begin on the same level as to structure that they have since maintained. I agree with Nathorst, however, in holding that the Bilobites" and many other forms believed by some to be sea-weeds, are really trails and tracks of animals. 2 The Foraminifera of the Palaeozoic we have noticed in the last chapter ; but we now find a new type of Protozoan that of the Sponge. Sponges as they exist at present may be defined to be composite animals, made up of a great number of one-celled or gelatinous zoids, provided with vibrating threads or cilia, and so arranged that currents of water are driven through passages or canals in the mass, by the action of the cilia, bringing food and aerated water for respiration. To support these soft sarcodic sponge-masses, they secrete fibres of horny matter and needles (spiculae) of flint or of 1 Eophyton Linn&Unum (Torrell). 2 See Paper on " Footprints and Impressions of Animals," Am. Journal of Science, 1873. THE AGE OF INVERTEBRATES OF THE SEA. 49 limestone, forming complicated fibrous and spicular skeletons, often of great beauty. They abound in all seas, and some species are found in fresh waters. With the exception of a very few species destitute of skeleton, and which we cannot expect to find in a fossil state, the sponges may be roughly divided into three groups : i, those with corneous or horny skeleton, like our common washing sponges ; 2, those with skeletons composed of silicious needles of various forms and arrangement ; 3, those with calcareous spicules. Of these, the second or silicious group has pre- FIG. 29. Portion of skeleton of Hexactinellid Sponge {Coeloptychii.m}. After Zittel. Magnified. cedence in point of time, beginning in the Early Cambrian, and continuing to the present. Two of its subdivisions are especially interesting in their range. The first is that of the Lattice-sponges (Hexcutinellida), in which the spicules have six rays placed at right angles, and are attached to each other by their points, so as to form a very regular network (Fig. 29). The second is that of the Stone-sponges (Lithistidcz), in which 50 THE CHAIN OF LIFE. the spicules are four-rayed or irregular, and are united by the branching root-like ends of the rays. The most beautiful of all sponges, the Venus Flower-basket (Euplectella), is a modern Hexactinellid, and the wonderful weaving of its spicules is as marvellous a triumph of constructive skill as its general form is graceful. The Lithistids are less beautiful, but are the densest and most compact of sponges, and are represented by several species in the modern seas. Both of these types go back to the Early Cambrian, and have continued side by side to the present day, as representatives of two distinct geometrical methods for the construction of a spicular skeleton. Many years ago the keen eye of the late lamented Salter FIG. 30. Protosfiongia fenestrata (Salter). Menevlan group, a. Fragment showing the spicules partially displaced. &, Portion enlarged. detected in a stain on the surface of a slab of Cambrian slate the remains of a sponge ; and minute examination showed that its spicules crossed each other, and formed lattice-work on the hexactinellid plan. Salter boldly named it Protospongia (the first sponge), and it is still the earliest that we know (Fig. 30). Thus the type whose skeleton is the most perfect in a mechanical point of view takes the lead. It is continued in the Silurian in many curious forms, of which the stalkless sponges (Astylospongia) are the most common (Fig. 31). It perhaps attains its maximum in the Cretaceous, from which the beautiful example in Fig. 29 is taken, and it still flourishes, giving us the most beautiful of all recent forms. Before the THE AGE OF INVERTEBRATES OF THE $EA. 51 close of the Cambrian there were other sponges of the Lithistid type. Fig. 32 represents a group of spicules from FIG. 31. Astylospongia. pr&morsa (Roemer). Niagara group. After Hall, a, Spicules magnified. the Calciferous (Lowest Silurian or Upper Cambrian) of Mingan, 1 and which probably belong to a large Lithistid sponge FIG. 32. Spicules of Lithistid sponge ( Trichospongia of Billings). From the Cambrian of Labrador. of that early time. The Lithistids have been recognised in the Upper Silurian and Carboniferous, and continuing upward to the Cretaceous, there become vastly numerous, while their 1 They probably belong to a large sponge named by Billings Tricho- spongia sericea. E 2 52 THE CHAIN OF LIFE. modern representatives are by no means few. The silicious sponges with simple spicules appear to have existed as far back as the Siluro-Cambrian, and there is believed to be almost as early evidence of horny or corneous sponges. The calcareous sponges have been recognised as far back as the Silurian. 1 Thus from the close of the Palaeozoic all the types of sponges seem to have existed side by side ; and in the Cretaceous period. FIG* 33. OldJtamia antiqnct (Forbes). Frc. 34. Dictyonema soctale. Enlarged. Lingula flags. After Salter. ] when such large areas of our continents were deeply submerged, they attained a wonderful development, perhaps not equalled in any other era of the earth's history. Sponges may be regarded as the highest or most complex of the Protozoa or the lowest of the Coelenterates. We have no links wherewith to connect them with the lower Protozoa of the Eozoic period ; .and through their long history, though very numerous in genera and species, they show no closer relation- ship with the Foraminifera below, and the Corals above, than do their successors in the modern seas. They thus stand very 1 Amphispongia. THE AGE OF INVERTEBRATES OF THE SEA. 53 much apart ; and modern studies of their development and minute structures do not seem to remove them from this isola- tion. Though we are treating here of inhabitants of the sea, it may be proper to mention that Geinitz has described two species from the Permian which he believed to be early precur- sors of the Spongillse, or fresh-water sponges ; but more recently he seems to regard them as probably Algae. Young has, how- ever, recently found true spicules of Spongilla in the Purbeck beds. 1 A stage higher than the sponges are those little polyp-like animals with sac-like bodies and radiating arms or tentacles, which form minute horny or calcareous cells, and bud out into c*. FIG. 35. Dictyontma Websteri (Dn ). Niagara formation, a, Enlarged portion (Acadian Geology). branching communities, looking to untrained eyes like delicate sea-weeds the sea-firs and sea-mosses of our coasts (Fig. 36). These belong to a very old group, for in the oldest Cambrian we have a form referred to this type (Fig. 33), and in the Upper Cambrian another still more decided example (Fig. 34). 2 This style of life, once. introduced, must have increased in variety and extended itself with amazing rapidity, for in the Siluro-Cambrian age we find it already as characteristic as in our modern seas, and so abundant that vast thicknesses of shale are filled and blackened with the debris of forms allied to the sea-firs, and masses of limestone largely made up of the 1 Geological Magazine, May, 1878. 2 It is regarded as somewhat doubtful whether these are Hydroids or Bryozoa. 54 THE CHAIN OF LIFE. more calcareous forms of the sea-mosses. As examples of the former we may take the Graptolites, so named from their re- semblance to lines of writing, and of which several forms are represented in Fig. 37. The little teeth on the sides of these were cells, inhabited probably by polyps, like those repre- sented in the modern Sertularia in Fig. 36. Some of them were probably attached to the bottom. In others the branches radiated from a central film which may have been a hollow FIG. 36. Group of modern Hydroids allied to Graptolites. Magnified, and natural sire. a, Sertularia. b, Tubiilaria. c, Campanularia vesicle or float, enabling them to live at the surface of the water (Fig. 38). These Graptolites are specially character- istic of the Upper Cambrian and Lower Silurian. The netted ones (Dictyonemd), as may be seen from Figs. 34 and 35, came in before the close of the Cambrian, and continue unchanged to the Silurian, where they disappear. The branching forms, seen in Fig. 37, have scarcely so great a range. They thus form most certain marks of the period to which they belong, THE AGE OF INVERTEBRATES OF THE SEA. 55 and being oceanic and probably floaters, they diffused them- selves so rapidly that they appear to indicate the same geological time in countries so widely separated as Europe, North America, FIG. 37. Silurian Graptolitidae. a, Graptolithus. b, Diplograpsus. c, Phyllograpsus. d, Tetragrapsus. e, Didyjnograpsus. and Australia. It is curious, too, that while the Graptolites thus mark a definite geological time, and seem to disappear abruptly and without apparent cause, they are the first link in the long chain of the Hydroids, which, though under different FIG. 30. Ptibdictya ac-uta (Hall). Bryo- zoan. Siluro-Cambrian. FIG. 38. Central portion of Graptolite, with membrane, or float (Dicho- grapsus octobrachiatus, Hall). family forms, continue to this day, apparently neither better nor worse than their perished Palaeozoic relatives. There is a group of little Stony Corals (Monticuliporida), which were possibly also THE CHAIN OF LIFE. the cells of Hydroids, that have a similar history. They are the only known Corals that date so far back as the Upper Cambrian ; and they continue under very similar forms all FIG. ypc.Fenestetta: Lye Hi (Dawson). A Carboniferous Bryozoart through the Palaeozic, and are represented by the millepore corals of the present day. Fig. 40 represents a form found at the base of the Siluro-Cambrian, and Fig. 41 shows forms characteristic of the Carboniferous Lime- stone. If we turn now to the sea-mosses (Bryozoa). we have a group of minute polyp-like animals inhabiting cells not unlike those of the Hy- droids, and which form plant-like aggregates. But the animals themselves are so different in structure that they are considered to be nearer allies of the bivalve shell-fishes than of the Corals. They are, in short, so different, that the most ardent evolutionist would scarcely hold a community of origin between them and such creatures as the Graptolites and Millepores, though an ordinary observer might readily confound the one with the other. These animals appear at the beginning of the Siluro-Cambrian, and such forms as that represented in FIG. 40 Chaetetes fibrosa. A tubulate coral with micro- SQopic cells. Siluro- Cambrian. THE AGE OF INVERTEBRATES OF THE SEA. 57 Fig. 39, very closely allied to some now living, are large consti- tuents of some of the limestones of that period. Other forms, like that represented in Fig. 390, are very characteristic of the Carboniferous. These animals, individually small, though com- plicated in structure and branching into communities, scarcely ever of any great magnitude, humble creatures which have never played any great part in the world, have, nevertheless, been so persistent that, though specific and generic forms have been changed, the group may be said to be in the modern seas exactly what it was in those of the early Palaeozoic, nor can it be affirmed to have originated in anything different, or to have produced anything. FIG. 41. a, Stenopora e xilis (Dawson). 6, Chaetetes tumidus( Edwards and Haine). Carboniferous. The true Stony Corals (Anthozoa] are as yet unknown in the Cambrian. They entered on the stage in immense abundance in the Siluro-Cambrian, where considerable limestones are largely composed of their remains, mixed, however, and some- times overpowered with those of Bryozoa and Hydroids. An ordinary coral, such as those of which coral reefs are built the red coral, used for ornament is not quite similar is the skeleton of an animal constructed on the plan of a sea anemone ; with a central stomach surrounded by radiating chambers, and having above a crown of tentacles. The stony coral sur- rounds and protects the soft body of the animal, and may either be a single cell, for one animal, or an aggregation of 58 THE CHAIN OF LIFE. such cells, .constituting a rounded or branching mass. The modern star coral, represented in Fig. 42, is an instance of the latter condition. It shows nineteen or twenty animals, each with a central mouth and fringe of short tentacles, aggregated together, and two of them showing the spontaneous division by which the number of animals in the mass is progressively increased. The living coral shows only the soft animals and the animal matter connecting them ; but if dead there would be a white stony mass with a star-like cell or depression corresponding to each animal. In their general plan, the oldest Corals were precisely of this character, but they presented some differences in detail, which have caused them to be divided into two groups, which are FIG. 42. Living Antho2oan Coral (Astr&a}. eminently characteristic of the Palaeozoic age the tabulate or floored corals, and the rugose or wrinkled corals. In the former (Fig. 43) the cells are usually small and thin- walled, often hexagonal, like a honeycomb, and are floored across at intervals with tabulae or horizontal plates. A few modern corals present a similar arrangement, 1 but this kind of structure was far more prevalent in the Palaeozoic. In the second type the animals are usually larger and often solitary, the cell has strongly marked radiating plates, while the horizontal floors are absent or subordinate, and there is usually a thick exter- nal rind or outer coat (Figs. 44, 45). In general plan, these 1 Ileliopora, an Alcyonarian ; Pccillopora, an Anthozoan. THE AGE OF INVERTEBRATES OF THE SEA. 59 rugose corals closely resemble those of our modern reefs ; but they differ in their details of structure, and only a very few FIG. 43. Tabulate Corals. a, Halisites* and b, Favosites, Upper Silurian. modern forms from the deep sea are regarded as actual modern representatives. 1 One curious point of difference is that their FIG. 44. Rugose Coral (Heliophyllum Halli). Devonian. radiating laminae begin with four, and increase by multiples of that number, while in modern corals the numbers are six and multiples of six ; a change of mathematical relation not easily accounted for, and which assimilates them to Hydroids on the 1 Haflophyllia, Guynia, Duncania^ of Pourtales. 6o THE CHAIN OF LIFE. one hand, and to a higher group, the Alcyonids, on the other, both of which prefer four and eight to six, or have had these numbers chosen for them. In the Mesozoic period the tabulate and rugose corals were replaced by others, the porous and solid FIG- 44#. Zaphrentis prolifica (Billings). Devonian. corals of the modern seas ; but, in so far as we know, the animals producing these, though differing in some details, were neither more nor less elevated than their predecessors, and they took up precisely the same role as reef-builders in the sea, FIG. 45. Rugose Corals. a, Zaphrentis Minos (Dn.), and b, Cyathophyllum Billingsi (Dn.). Carboniferous. though with probably more tendency to the accumulation of great masses of coral limestone in particular spots. Leaving the corals, we may turn to the sea-stars and sea- THE AGE OF INVERTEBRATES OF THE SEA. 61 urchins. These merely put in an appearance in the Early Cambrian, but become vastly multiplied in the Silurian, where the stalked feather stars (Crinoids) (Fig. 46) seem to have covered great areas of sea-bottom, and multiplied so rapidly that thick sheets of limestone are largely made up of FIG. 4 . Modern Crinoid (Rhizocrinus Lofotensis). After Sars. the fragments of their skeletons. The ordinary star-fishes appear first in the Silurian (Fig. 47). The sea-urchins begin in the Upper Silurian, the early species having numerous and loosely attached plates, like some of those now found in the deep sea l (Fig. 48). 1 Pal&chinus. 62 THE CHAIN OF LIFE. The most curious history in this group is that of the feather- stars. In the Early Cambrian they are represented by a few species known to us only in fragments, and these belong to a humble group (Cystideans) resembling the larval or immature condition of the higher Crinoids. Fig, 49 shows one of these animals of somewhat later age. They have few or rudimentary arms and short stalks, and want the beautiful radial symmetry of the typical star-fishes. In the Silurian these creatures are reinforced by a vast number of beautiful and perfect feather- stars (Figs. 50, 51). These continue to increase in number and beauty, and apparently culminate in the Mesozoic, where gigantic forms exist, some of them probably having more FIG. 47. Pal&aster Niagarensis FIG. 48. Palcechinus ellipticus (McCoy). (Hall). One of the oldest star fishes. One of the oldest types of sea-urchins. complicated skeletons, in so far as number of distinct parts is concerned, than any other animals. Buckland has calculated that in a crinoid similar to that in Fig. 52 there are no less than 150,000 little bones, and 300,000 contractile bundles of fibres to move them. In the modern seas the feather-stars have somewhat dwindled both in numbers and complexity, and are mostly confined to the depths of the ocean. On the other hand, the various types of ordinary star-fishes and sea- urchins have increased in number and importance. We thus find in this group a certain advance and improvement from the Cystideans of the Early Palaeozoic to the sea-urchins and their allies. This advance is not, however, along one line for the -THE AGE OF INVERTEBRATES OF THE SEA. 63 Cystideans continue unimproved to the end. The Crinoids culminate in the Mesozoic, and are not known to give origin to anything higher. The star-fishes and sea-urchins commence independently, before the culmination of the Crinoids, and, though greatly increased in number and variety, still adhere very closely to their original types. Fis. 49. Pleurocystites FIG. 50. Heterocrinus FIG. 51. Body of Glypto- squamosus. Siluro-Cam- simplex (Meek). One of crz'nus. Siluro-Cambrian. brian. After Billings. the least complex crinoids of that period. Siluro- Cambrian. The great sub -kingdom of the Mollusca, including the bivalve and univalve shell-fishes, makes its first appearance in the Cambrian, where its earliest representatives belong to a group, the Arm-bearers or Lamp shells (Brachiopods), held by some to be allied to worms as much as to mollusks. The oldest of all these shells are allies of the modern Lingulcz (Fig. 54), some of 6 4 THE CHAIN OF LIFE. the earliest of which are shown in Fig. 55. The modern Lin- gula is protected by a delicate two-valved shell, composed, unlike that of most other mollusks, of phosphate of lime or bone earth. It lives on sand-banks, attached by its long flexible stalk, which it buries like a root in the bottom. Its food consists of microscopic organisms, drifted to its mouth by cilia placed on two arm-like processes, from which the group derives its name. In the modern world about one hundred FIG. 52. Extracrinus Briarens. Reduced. Jurassic. FIG. 53. P entacrinns caput-inedii Reduced. Modern. species of Brachiopods are known, belonging to about twenty genera, some of which differ considerably from the Lingulse. The genus Terebratula, represented at Fig. 56, is one of the most common modern as well as fossil forms, and has the valves unequal, with a round opening in one of them for the stalk, which is attached to some hard object, and there is an internal shelly loop for supporting the arms. -THE AGE OF INVERTEBRATES OF THE SEA. 65 These curious, and in the modern seas, exceptional shells, were dominant in the Palaeozoic period. Upwards of three thousand fossil species are known, of which a large proportion belong to the Cambrian and Silurian, nine genera appearing in the Cambrian, and no less than fifty-two in the Silurian. The history of these creatures is very remarkable. The Lingulae, which are the first to appear, continue unchanged and with the same phosphatic shells to the present day. Morse, who has FIG. 54. Lingula anatina. With flexible muscular stalk. Modern. FIG- 55. Cambrian and Silurian Lingulae. a> Lingiilella Matthewi (Hartt). Acadian group* b, Lingula, quadrata (Hall). Siluro-Cambrian, c, Lingulella prima (Hail). Potsdam. , Its dorsal fold. c, Hood. o, Eye. t, Tentacles, f, Funnel, g, Air chambers. ^, Siphuncle. collections, though its mechanical structure and advantages for the struggle for existence seem of the highest order. But in the old world of past geological time the case was altogether different. The Nautiloid shell-fishes burst suddenly upon us in the beginning of the Siluro-Cambrian, or Lower Silurian, Barrande's second fauna ; and this applies to all the countries where THE AGE OF INVERTEBRATES OF THE SEA. 73 they have been studied. In this formation alone about 450 species are known, and in the Silurian these increase to 1,200 ; and here the group culminates. It returns in the Devonian to about the same number with the Lower Silurian, diminishes in the Carboniferous to 350, and in the Mesozoic, where the Nautiloid forms are replaced by others of the type FIG. 67. Gomphocer^s. FI-.J. 66.Orthoceras. Siluro-Cam- brian. The dotted line shows the position of the siphuncle. FIG. *8>.Lituites. of the Ammonites, becomes largely reduced. In the Tertiary there are but nineteen species, and, as already stated, in the modern world three. These statements do not, however, re- present the whole truth. In the Palaeozoic, in additon to the genus Nautilus, we have a great number of other genera, some with perfectly straight shells, like Orthoceras (Fig. 66), others 74 THE CHAIN OF LIFE. bent (Cyrtoceras), others differing in the style of siphuncle, or aperture, or chambers (EndocertR^ Gomphoceras, Lituites> FIG. 69. Nautilus Avonensis (Dn.). Carboniferous. a, Shell, reduced, b, Section, showing siphuncle. Figs. 67 to 69), or inflated into sac-like forms (Ascoceras). There is, besides, the family of the Goniatida (Fig. 70), with the FIG. 70. Goniatites crenistria (Philips). Carboniferous. chambers thrown into angular folds and the siphuncle at the back. 'Further, some of the early forms, as the Orthoceratidae, THE AGE OF INVERTEBRATES OF THE SEA. 75 attain to gigantic dimensions, being six feet or more in length, and nearly a foot in diameter. Thus the idea that we should naturally form from the study of the Nautilus, that it represents a type suited for much more varied and important adaptations than those that we now see, is more than realised in those Palaeozoic ages when these animals seem to have been the lords of the seas. When we leave the Palaeozoic and enter the Mesozoic, though the Nautiloid shells still abound, we find them superseded, in great part, by a nobler form, that of the Ammonitidce (Figs. 7 1, FIG. 71. Ceratites nodosus (Schloth). Triassic. 72). These are remarkable for the ornate markings on the surfaces of their shells, and for the beautifully waved edges of the partitions (Fig. 720), which, by giving a much more com- plete support to the sides of the shell, must have contributed greatly to the union of lightness and strength so important to the utility of the shell as a float This type admits of all the same variety of straight, bent, and curled forms with the simpler Nautiloid type, and some of the species are of great size, Am- monites being known three feet or more in diameter. These animals, unknown in the Palaeozoic, appear in numerous species 7 6 THE CHAIN OF LIFE. in the Early Mesozoic, culminate in hundreds of beautiful species in the middle of that era, and disappear for ever at its close, leaving no modern successors. Many and beautiful species of Ammonites and their allies have been obtained from the FIG. 72. Ammonites Jason (Reinecke). Jurassic. Mesozoic rocks of British Columbia and other parts of the west coast of North America, perfectly representing this group as it occurs at the same period in Europe, and closely resembling FIG. jza. Suture of Ammonites componens (Meek), of British Columbia. Showing the complicated folding of tbe edges of the chambers to give strength to the shell. Cretaceous. the Mesozoic Ammonites of India. These animals have all perished, yet the Atlantic and the Pacific roll between, apparently with conditions as favourable for their comfortable existence as those of any previous time. They perished long ago, at the - THE AGE OF INVERTEBRATES OF THE SEA. 77 dawn of the Tertiary ; yet the genus Nautilus, one of the oldest and least improved of the whole, survived, and still testifies to the wonderful contrivance embodied in these animals. These are merely general considerations, but Barrande, in his Etudes Generates, goes much farther. He sums up all the known facts in the most elaborate manner, considering first the em- bryonic characters of the shell in the different genera, then their FIG. 73. Cretaceous Ammonitidae. a, Baculites. b, Ancyloceras. c, Crioceras. d, Turrilites. distribution in space and time, then all the different parts and characters of the shells in the different groups the whole with reference to any possible derivation of the species ; and he finds that all leads to the result that in every respect these shells seem to have been so introduced as to make any theory of evolution with respect to them altogether untenable. In his concluding sentence this greatest of Palaeozoic palaeontologists 78 THE CHAIN OF LIFE. affirms that, "The theoretical evolution of the Cephalopods is, like that of the Trilobites, a mere figment of imagination, without any foundation in fact." 1 I have reserved no space to notice the geological history of the other and higher group of Cephalopods, including the true Cuttles and Squids. This is perhaps less to be regretted, as, from the absence of external shells, they are likely to be much FIG. 74. Belemnite. After Philips. less perfectly known as fossils. So far as known, they are vastly younger than the Nautiloids, for no examples whatever have been found in the Palaeozoic. They appear abundantly in FIG. 74. Belemnoteuthis antiquus. Supposed to be a Belemnite, with soft parts preserved. Jurassic. After Mantell. the Mesozoic, but are there represented principally by an extinct group of squids (Belemnites and their allies, Figs. 74, 74nit (Dn..). B, S. elegans (Brongniart). B 1 , &c. Leaf and Leaf-scars. i io THE CHAIN OF LIFE. roots are the so-called Stigmarice, so abundant in every coal-field, and especially filling the " under-clays" of the coal-beds, which are the soils on which the plants forming these beds were sup- ported. The true botanical position of the Sigillaria has been a matter of much controversy. Some of them undoubtedly have structures akin to those of the tree-like Club-mosses, as Williamson has well shown, and may have been cryptogamous. Others have structures of higher character, akin to those of the modern Cycads, and seem to have borne nutlets allied to those of these plants. Yet the external forms of these diverse sorts are so similar that no definite separation of them has yet been made. Either these anomalous trees constitute a link connecting the two great series of the vegetable kingdom, or we have been confounding two distinct groups, owing to imperfect information. Another curious, and till recently little understood, group of Carboniferous trees is that known as Cordaites, which existed already in some of its species in the Devonian. Their leaves are long, and often broad as well, and with numerous delicate parallel veins, resembling in this the leaves of grasses. Corda long ago showed that one species at least has a stem allied to the Club-mosses. More recently Grand' Eury has found in the South of France admirably preserved specimens, which show that others more resembled in their structure the Pines and Yews, and were probably Gymnosperms, approaching to the Pines, but with very peculiar and exceptional foliage, of which the only modern examples are the broad-leaved Pines of the genus Dammara (Frontispiece to Chapter). Here again we have either two very distinct groups, combined through our ignorance, or a connecting link between the Lycopods and the Pines. The Yews and their allies among modern trees, while mem- bers of the great Cone-bearing order, bear nut-like seeds in fleshy envelopes, sometimes, as in the Ginkgo of Japan, consti- tuting edible fruits. Seeds of this type seem to have been THE ORIGIN OF PLANT LIFE ON THE LAND, ill extremely abundant in the Carboniferous age in all parts of the world, and were probably produced by trees of several genera (Dadoxylon, Sigillaria^ Cordaites, etc.) (Fig. 101). Charles Brongniart has recently described no less than seven- teen genera of these seeds from the coal-field of St. Etienne alone, and it would be a low estimate to say that we probably FIG. 101. Trigonocarpum Hookeri (Dn.). A Gymnospermous seed. a, Testa, b, Tegmen. c, Nucleus, d, Embryo. know as many as sixty or seventy species in all, while the trunks of great coniferous trees allied to Taxineae, and showing well-preserved structure, are by no means uncommon in the Devonian and Carboniferous. Had these great Yews appeared for the first time in the Coal-formation, we might have supposed that they had been developed from such Lycopods as Lepidodendra, and that the Cordaites are the intermediate 112 THE CHAIN OF LIFE. forms ; but unfortunately the Pines go almost as far back in geological time as the Lycopods, and it does not help us, when in search of evidence of evolution, to find the link which is missing or imperfect in the Early Devonian supplied in the Coal-formation, where, for this purpose at least, it is no longer needed. We have said something of what was in the Palaeozoic flora ; but what of that which was not ? We may answer : Nearly all that is characteristic of our modern forests, whether in the ordinary Exogens, which predominate so greatly in the trees and shrubs of temperate climates, or in the Palms and their allies, which figure so conspicuously within the tropics. The few rare, and to some extent doubtful, representatives of these types scarcely deserve to be noted as exceptions. Had a botanist searched the Palaeozoic forests for precursors of the future, he would probably have found only a few rare species, while he would have seen all around him the giant forms and peculiar and monotonous foliage of tribes now degraded in magnitude and structure, and of small account in the system of nature. It must not be supposed that the Palaeozoic flora remained in undisturbed possession of the continents during the whole of that long period. In the successive subsidences of the conti- nental plateaux, in which the marine limestones were de- posited, it was to a great extent swept away, or was restricted to limited insular areas, and these more especially in the far north, so that on re-elevation of the land it was always peopled with northern plants. Thus there were alternate restrictions and expansions of vegetation, and the latter were always signalised by the introduction of new species, for here, as elsewhere, it was not struggle, but opportunity, that favoured improvement. In the Lower Silurian such plants as existed must have ex- perienced great restriction at the age of the Niagara or Wen- lock limestone. Those of the Upper Silurian suffered a similar reverse at the time of the Lower Helderberg or Ludlow lime- THE ORIGIN OF PLANT LIFE ON THE LAND. 113 stones. This recurred at the close of the Devonian and in the time of the Lower Carboniferous limestone ; and finally the Palaeozoic flora disappeared altogether in the Permian, to be replaced by new types in the Mesozoic. While, therefore, there is a great general similarity in the successive Palaeozoic floras, there are minor differences, so that the Devonian plants are for the most part distinct specifically from those of the Lower Car- boniferous, those of the Lower Carboniferous from those of the Coal-formation, and those of the latter from those of the Permian. With all these vicissitudes it is to be observed that there is no apparent elevation of type in all the long ages from the Devonian to the Permian, that the Acrogens and Gymnosperms of these periods are in some respects superior, in all respects equal, to their modern successors, and that their history shows a decadence toward the modern period ; that intermediate forms arrive too late to form connecting links in time, that several distinct types appear together at the beginning, and that all utterly and apparently simultaneously perish at the end of the Palaeozoic, to make way for the entirely new vegetation of the succeeding age. Theories of evolution receive no support from facts like these, though their practical signifi- cance, as parts of the one great uniform scheme of nature, is sufficiently manifest. Of what use then were these old floras? To the naturalist, vegetable life, with regard to its modern uses, is the great accumulator of pabulum for the sustenance of the higher forms of vital energy manifested in the animal. In the Palae- ozoic this consideration sinks in importance. In the Coal period we know few land animals, and these not vegetable feeders, with the exception of some insects, millipedes, and snails. But the Carboniferous forests did not live in vain, if their only use was to store up the light and heat of those old summers in the form of coal, and to remove the excess of carbonic acid from the atmosphere. In the Devonian period 114 THE CHAIN OF LIFE. even these utilities fail, for coal does not seem to have been accumulated to any great extent, and the petroleum of the Devonian appears to have been produced from aquatic vege- tation. Even with reference to theories of evolution, there seems no necessity for the long continuance and frequent changes of species of acrogenous plants without any perceptible elevation. We may have much yet to learn of the life of the Devonian ; but for the present the great plan of vegetable nature goes beyond our measures of utility ; and there remains only what is perhaps the most wonderful and suggestive correlation of all, namely, that our minds, made in the image of the Creator, are able to trace in these perished organisms structures similar to those of modern plants, and thus to reproduce in imagination the forms and habits of growth of living things which so long preceded us on the earth. We may indeed proceed a step further, and hold that, independently of human appreciation, these primitive plants commended themselves to the approval of their Maker, and perhaps of higher intelli- gences unknown to us ; and that in the last resort it was for His pleasure that they were created. 1 CHAPTER V. THE APPEARANCE OF VERTEBRATE ANIMALS. /^"CONFESSEDLY the highest style of animal is that which \^s possesses a skull and backbone, with brain and nerve system to match, and which embodies the general plan of structure employed in man himself. Yet among the fishes, which constitute the lowest manifestation of this type, are some so rudimentary that the brain is scarcely developed, and the skeleton is merely a cord of gristle. These are represented in the modern world only by the Lancelot, 1 a creature which has sometimes been mistaken for a worm, and by a slightly more advanced type, that of the Lampreys. 2 In these animals the Vertebrates make the nearest approach to the lower domains of the animal kingdom, collectively known as Inverte- brates. We should naturally expect that since the vertebrates succeed the inferior animals in time, their lower types should appear first, and that these should be aquatic rather than terrestrial. On the other hand, as the oldest fishes that are certainly known ' are strongly protected with bony armour, and had to contend against formidable Crustaceans and Cuttles, we might suppose that the Lancelot and the Lampreys are rather degraded types belonging to the modern period, than the true precursors of the other fishes. 1 Amph^o.\^ts. 2 Fetromyzon, &c. Ii8 THE CHAIN OF LIFE. But if fishes like the Lancelot preceded all others, we may never find in a fossil state any traces of their soft and perish- able bodies ; and even the Lampreys have no hard parts except small horny teeth, which might easily escape observation. But palaeontologists have sharp eyes, and it has not escaped them that certain microscopic tooth-like bodies are somewhat widely distributed in the older rocks. In Russia, Pander has found in the Upper Cambrian and Lower Silurian, and also in the Devonian and Carboniferous, minute conical and comb like FIG. 102. Siluro-Cambrian Conodonts. Magnified. After Pander. teeth, to which he has given the name of Conodonts (Fig. 102), and which he supposes to be the teeth of ancient Lampreys. Similar teeth have been found by Moore and others in the Carboniferous of England, and by Newberry in Carboniferous shales in Ohio. In point of form, these bodies certainly re- semble the teeth of the humble fishes to which they have been referred. In the case of the Carboniferous specimens from Ohio the only ones I have had an opportunity to examine APPEARANCE OF VERTEBRATE ANIMALS. 119 the material is calcium phosphate, and the structures are more like those of teeth of Sharks than of Lampreys, so that there can be no doubt that they are really teeth of fishes, and pro- bably of fishes of somewhat higher grade than the Lampreys. 1 The Cambrian and Silurian specimens are said to be composed of calcium carbonate, which would render it more probable that, as has been suggested by Prof. Owen, they may have been teeth of some species of Sea-snail destitute of shell. It is, however, possible that they may have originally been horny, and that the animal matter has been replaced by carbonate of lime. Rohon and Zittel have recently shown that many of these are more allied to the teeth of worms than of any other animals. 2 FIG. 103. Lower Carboniferous Conodont. Magnified. After Newberry. If these older Conodonts were really teeth of fishes, they carry the introduction of these nearly as far back as that of the Mollusks and Crustaceans. If they were not, then the earliest known representatives of this class belong to a much later age, that of the Silurian. Here we have undoubted remains of fishes belonging to two of the higher orders of the class ; and in the succeeding Devonian these became multiplied and extended exceedingly. 1 Dr. Newberry has kindly furnished me with specimens, and Dr. Har- rington has submitted to analysis portions of shale filled with these little teeth, the result giving 2'$& of calcium phosphate for the whole, which indicates that the Conodonts are really bone. Their microscopic structure approaches to that of the dentine of such Carboniferous fishes as Diplodus. Hinde has described Conodonts from the Silurian of Canada. 2 Ueber Conodont en \ Munich, 1886. 120 THE CHAIN OF LIFE. Besides the inferior tribes already referred to, the modern seas and rivers present four leading types of fishes : first, the ordinary bony fishes (Teleostians), such as the Cod, Salmon, and Herring ; secondly, the Ganoid fishes, protected with bony plates on the skin, as the Bony-pike 1 and Sturgeon ; thirdly, the Sharks and their allies, the Dog-fishes and Rays : fourthly, the peculiar and at present rare group of semi-reptilian fishes to which the name of Dipnoi has been given, on account of their capacity for breathing both in air and in water. Of these four types the first is altogether modern, and includes the great majority of our present fishes. It does not make its appearance till the Cretaceous age, and then is at once represented by at least three of the modern families, those of the Salmon, Herring, and Perch. The history of the other three groups is precisely the opposite of this. They abound exceedingly at an early period, and dwindle to a much smaller number in the modern time. This is especially the case with the Ganoids and the Dipnoi. It is also remarkable that these groups of old-fashioned fishes 2 are in some respects the highest members of the class, approaching the nearest to the reptiles ; but this accords with a well-known palaeontological law, namely, that the higher members of low groups give way on the intro- duction of more elevated types, while the lower members may continue. Thus the decadence of these higher fish begins with the incoming of the reptiles, just as the decadence of the higher Mollusks and predaceous Crustaceans began with the incoming of the fishes. Further, the modern Ganoids and Dipnoi are mostly fresh-water animals, though the Sharks are largely pelagic. In the Palaeozoic there seem to have been abundance of marine species of all these types ; but though marine, they probably flourished most in bays and estuaries and on shallow banks ; and the existence of these implies continental masses of land. This explains the curious coincidence that the introduction of fishes and of an abundant land flora synchronise, and 1 Lepidosteus. * Palceichthyes of Giinther. APPEARANCE OF VERTEBRATE ANIMALS. 121 that the ocean was still dominated by Invertebrates long after the fishes had become supreme in bays, estuaries, and rivers. The first fishes that we certainly know are the Ganoids and Sharks, which appear near the close of the Upper Silurian, in the English Ludlow for example (Fig. 104). The Ganoids found here all belong to an extinct group, characterised by the covering of the head and anterior part of the body with large bony plates. They are mostly small fishes, and probably fed at the bottom, and used their long or rounded bony snouts for FIG- 104. a, Head shield of an Upper Silurian fish (CyatJtaspis). 6, Spine of a Silurian Shark (Onchus tenui-striatus, Agass.). c, d, Scales of Thecodus, enlarged. grubbing in the mud for food. In this respect they present a singular resemblance to the Trilobites, so that we seem to have here animals of an entirely new type, the Vertebrate, and with bony instead of shelly coverings, taking up the role and, to some extent, the external form of a group about to pass away. Yet I presume that no derivationist would be hardy enough to affirm that the Trilobites could have been the ancestors of these fishes. Nor indeed is any ancestry even hypothetically known for them, for the doubtful Lampreys of the Cambrian Silurian are too remote and uncertain to be used in that way. 122 THE CHAIN OF LIFE. The head-shield copied in outline in Fig. 104, and the restora- tion after Lankester in the frontispiece to this chapter, may serve to represent these curious primitive Ganoids, which are continued in the Devonian fishes represented in Figs. 105, 106. Along with these, and not improbably their enemies, were certain Sharks (Fig. 104), known to us only by the spines which FIG. 105, Cephalaspis Dawsoni (Lankester). Lower Devonian of Gaspe. were attached to their fins as weapons of defence, and by detached bony tubercles which protected their skin. These remains are chiefly interesting as indications that two of the great leading divisions of the class of fishes originated together. In the Devonian age the Ganoids and Sharks, thus introduced in the Silurian, may be said to culminate. The former, more APPEARANCE OF VERTEBRATE ANIMALS. 123 especially, are represented by a great variety of species, some of them nearly allied to their Silurian predecessors (Fig. 106), others of forms and structure not dissimilar to those of the few surviving representatives of the order, or altogether peculiar to the Devonian (Fig. 107). So numerous are these fishes, and of so many genera and species and this not merely in one region, but in widely separated parts of the world that the Devonian has not inaptly been called the reign of Ganoids. FIG. 106. Devonian Placoganoid Fishes (Pterichthys comutus, Cephalaspis Lyelli), from Scotland. As an illustration at once of the very peculiar forms of some of these fishes and of their wide distribution, I figure here along with the British species a Cephalaspis (Fig. 105) found in the Lower Devonian of Gaspe, in the same beds with some of the antique Devonian plants described in the last chapter. A new and interesting light has recently been cast upon some of the most anomalous of the ancient fishes by the study of the now rare and peculiar species of the group of Dipnoi. Two of these, belonging to the genus Lepidosiren, are the 124 THE CHAIN OF LIFE. " Mud-fishes " of the rivers of tropical Africa and America (Fig. 1 08, b.) These creatures have an elongated and elegant form, FIG. 107. Devonian Lepidoganoid Fishes (Diplacanthus and Osteolepis). After Page and Nicholson. and the body is covered with overlapping horny scales like those of ordinary fishes ; but the pectoral and ventral fins are rod-like, FIG. 108. Modern Dipnoi. a, Ceratodus Fosteri. Australia, b, Lepido&iren annectus. Africa. and are supported by simple cartilaginous rays, while the tail- fin forms a fringe around the posterior part of the body., APPEARANCE OF VERTEBRATE ANIMALS. 125 Unlike ordinary fishes, they have lungs as well as gills, and their mouths are armed with sharp, bony, beak-like teeth (Fig. 115), with which they can inflict terrible bites on the small fishes and frogs which furnish them with food. Their most remarkable habit is that of burying themselves in the mud of dried-up ponds, thus forming a sort of water-chamber or " cocoon," in which they remain in a torpid state until the return of the rainy season sets them free. Another example of these Dipnoi is the Barramunda, or Ceratodus of the Australian rivers (Fig. 1080). This fish re- sembles the Lepidosiren in many essential points of structure ; FIG. 109. Anterior part of the palate of Dipterus. Showing the dental plates at Devonian. After Traqoair. but its fins have lateral rays, and are consequently of some breadth, though of peculiar form, and its mouth is armed with flat, pavement-like teeth, wherewith it browses on aquatic grasses. These modern fishes have enabled us to understand several mysterious forms met with in the older rocks. In the first place, they show the meaning of certain flat-toothed fishes, like Dipterus of the Devonian (Fig. 109), Conchodus of the Car- boniferous (Fig. no), and Ceratodus of the Carboniferous and Trias (Figs, in, 112), previously of very doubtful character. These must all have been of similar structure and habits with 126 THE CHAIN OF LIFE. the Barramunda, which is thus the sole survivor, perhaps itself verging on extinction, of a group of herbivorous fishes introduced, it may be, contemporaneously with the first stream affording the requisite vegetable food, and which have continued almost without improvement or deterioration to the present time. FIG. no. Dental plate of Conchodus plicatus (Dn.). Coal-formation of Nova Scotia. Acadian Geology. These fishes are, however, very closely connected with the Ganoids, and there are some of these, with fringed fins and overlapping scales, which, while regarded as true Ganoids, resemble the Dipnoi very closely. FIG. in. Dental plate of Ceratodus Barrandii. Coal-formation of Bohemia. After Fritsch. Again, certain huge fishes, whose remains are found in the Devonian of Ohio, 1 had jaws on the same plan with those of Lepidosiren, but of enormous size and strength (Figs. 113, 114, 1 Dinichthys Terrelli and Z>. Hertzeri (Newberry). APPEARANCE OF VERTEBRATE ANIMALS. 127 115), so that in this and some other points of structure they may be regarded as colossal Mud-fishes, and they must have FIG. ii2. Dental plate of Ceratodus serratus. From the Trias. had the same destructive powers, but on a far grander scale. They were besides clothed with heavy armour of bony scales, having some resemblance to that of those mailed fishes of FIG. 113. Jaws of D;nichthys Hertzeri (Newberry). Laterally compressed ; one-sixth natural size. smaller size already referred to, and indicating that, huge though they were, and formidable in destructive power, they 128 THE CHAIN OF LIFE. also had enemies to be dreaded. These plates serve to ally them with the Ganoids, as their jaws do with Lepidosiren. We are thus enabled to see in the streams, lakes, and bays of the Palaeozoic, harmless fishes, of the type of Ceratodus, feeding on plants, and huge precursors of the Mud-fishes FIG. 114. Lower Jaw of Dinichthys Htrtzeri, One-sixth natural size. darting from the depths, and provided with a dental apparatus more formidable than that of any modern fish, sufficient to pierce the strongest armour of the Ganoids, and to destroy and devour the largest aquatic animals. These huge fishes, armed with shears two or three feet in length, and capable of cutting FIG. 115. Jaws of Lepidosiren. Natural size. After Newberry. asunder scale, flesh, and bone, are the beau ideal of destructive monsters of the deep, far surpassing our modern Sharks ; and if, by means of supplementary lungs, they could breathe in air as well as in water, they would on that account be all the more vigorous and voracious. APPEARANCE OF VERTEBRATE ANIMALS. 129 Newberry has well remarked that while in the Devonian the Ganoids and Dipnoi were the real tyrants of the sea, as well as of the streams, in the Carboniferous they already diminish in size, though still abundant as to numbers, and are more limited to estuaries and fresh waters. Thus their departure from power had already begun, and went on until in modern times the proportion of Ganoids to ordinary fishes is, according to Giinther, nine out of 9,000. The Carboniferous, indeed, very specially abounds in small Ganoids, though there are many large and formidable species. One of these smaller species, a FIG. 116. A small Carboniferous Ganoid (Pal&oniscns (Rhadinichthys) Modulus' Dn.). Lower Carboniferous, New Brunswick, a. Outline, b, c, d, Sculpture of scales' magnified. very beautiful little fish, of fresh-water ponds and streams in the older part of the Carboniferous age, is represented of the natural size in Fig. 116, and is not a restoration, being found preserved entire, though flattened, in a fine bituminous shale, which has perfectly preserved even the most delicate sculpturing of its bony scales The Sharks in the Carboniferous increase in number and importance. Fig. 117 shows a few examples of their teeth and spines. In the Carboniferous, however, there is a great 130 THE CHAIN OF LIFE. preponderance of those species with flat, crushing teeth fitted for grinding shells, 1 which in diminishing numbers continue up to the present time, when they are represented by the Port Jackson Shark and a few other species. The increase toward the modem time of the true Sharks 2 with sharp cutting teeth, is obviously related to the increase of the ordinary fishes which FIG. 117. Teeth and Spines of Carboniferous Sharks. Nova Scotia. Dij>lod-us penetrans. b, Psammodus. c, Cienoptychius cristatus. d, Spine, Gyracanth-us magntficus. One-eighth natural size. Acadian Geology. furnish them with food. Another curious difference, connected probably with the same circumstance, is the fact that in the sharp toothed Sharks of the Carboniferous the two side fangs of each tooth are the largest, or are exclusively developed (Fig. 117, a), while in later periods the central point becomes 1 Cestracionts. 2 Selachians. APPEARANCE OF VERTEBRATE ANIMALS. 131 dominant, or is developed to the exclusion of the others (Figs. 118, 119). The Ganoids and Dipnoi still, however, occupy a very im- portant place through the Mesozoic ages (Fig. 120), and it is only at the close of the Cretaceous that they finally give place to the Teleosts, or common fishes, which, though perhaps more fully specialised in purely ichthyic features, have dropped the reptilian characteristics of their predecessors (Fig. 121). It is interesting to observe that these old-fashioned fishes had culminated before the advent of air-breathing Vertebrates, which appear for the first time in the Carboniferous. It is further to be observed that groups of fishes furnished with means of aiding their gills by rudimentary lungs were especially suited to waters more charged with carbonic acid, and less with free oxygen, than those of more recent times. This remark especially applies to the mephitic and sluggish streams and lagoons of the Carboniferous swamps, where, in the midst of a rank VPp-f FlG - "8. Teeth of Cretaceous Sharks (ptocba and Ptyckodus).-A.for Leidy. tation and reeking masses of decaying organic matter, the half air-breathing fishes and the amphibious reptilian animals met with each other and found equally congenial abodes. Thus, independently of the fact that some of these fishes were probably vegetable feeders, it is not altogether an accident, but a wise adaptation, that caused the culmination of the reptilian fishes and batrachian reptiles to coincide with the enormous development of the lower forms K 2 1 3 2 THE CHAIN OF LIFE. of land-plants in the Devonian and Carboniferous. Another curious illustration of the diminishing necessity for air-breathing to the fishes, is the change of the tail from the unequally-lobed or heterocercal form, which pre- vailed in the Palaeozoic, to the more modern equally-lobed (homocercal) style in the Meso- zoic. The former is better suited to animals which have to rise rapidly to the surface for air, and is still continued in some modern fishes, which for other reasons need to ascend and descend, or to turn them- selves in the water; but the homocercal form is best suited to the ordinary fish, whether Ganoids or Teleosts (Fig. 122). It is curious also to find the beginning of the dominancy of the ordinary fish to coincide with FIG. 119. Tooth of a Tertiary Shark (Carcharodon). FIG. 120. A Liassic Ganoid (Dapediiis). Restored. After Nicholson. that of the broad-leaved exogenous trees in the later Cretaceous, and to precede immediately the appearance of the mammals on the land ; all these changes being related to the purer air, the APPEARANCE OF VERTEBRATE ANIMALS. 133 clearer waters, and the more varied continental profiles of the later geological periods. Thus physical improvement and the changes of animal and vegetable life are linked together by correlations which imply not only design, but prescience, whether we attribute these qualities to a spiritual Creator or to mere atoms and forces. ct FIG. 121. Cretaceous Fishes of the modern or Teleostian type. a, Beryx ' Le^vesiensis. English chalk, b, Portheus molossus (Cope). A large fish from the American Cretaceous. One twenty-eighth natural size. The history of fishes extends further through geological time than that of any other Vertebrates, and is perhaps more com- pletely known to us, in consequence of the greater facilities for the preservation of their remains in aqueous deposits. If we receive Pander's Conodonts as indicating a low type of 134 THE CHAIN OF LIFE. cartilaginous fishes, these must have continued for vast ages without any elevation, and struggling for a bare existence amidst formidable Cuttle-fishes and Crustaceans, before, under more favourable conditions, they suddenly expanded into the high and perfect types of Ganoids and Sharks. If we reject the early Conodonts, then the two last-mentioned types spring together and suddenly into existence, like the armed men from the dragon's teeth of Cadmus. They rapidly attain to numbers and grandeur unexampled in later times, and become the lords of the waters at the time when there was probably no Vertebrate FIG. 122. Modern Ganoids (Folypterus. Africa. Lepidosteus. America). life on the land. As the reptiles establish themselves on the land and in the waters, the Ganoids diminish, but the Sharks hold their own. At length the reign of reptiles is over, but the Ganoids, instead of resuming their pristine numbers, give place to the Teleosts, and become reduced to insignificance ; while the Sharks, profiting by the decadence of the great marine reptiles, remain the tyrants of the seas. This history is strangely unlike a continuous evolution ; but we are anticipat- ing facts which will fall to be discussed in a subsequent chapter. CHAPTER VI. THE FIRST AIR-BREATHERS. WERE our experience limited to the animals whose remains are found in the earlier Palaeozoic rocks, we might be unable to conceive the possibility of an animal capable of living and breathing in the thin and apparently uncongenial medium of air. More especially would this appear doubtful if our experience of the atmosphere presented it to us as loaded with carbonic acid, and less rich in vital air than it is at present. Even the mechanical difficulties of the case might strike us as considerable, in our ignorance of the capabilities of limbs. Still, as time wore on, we should find this problem worked out along three distinct lines of advancement those of the Mol- lusk, the Arthropod, and the Vertebrate, and in each of these with different machinery, related to the previous locomotive and water-breathing apparatus of the type. Respiration under water depends, not on the water itself, but on the small percentage of free oxygen which it contains, and this is utilised for the aeration of the blood of animals, by that wonderful and often extremely beautiful apparatus of deli- cate fibres or laminae penetrated with blood-vessels, which we call a gill. Except those lowest creatures which aerate their blood merely at the general surface of the body, all animals capable of respiration in water are provided with gills in some 138 THE CHAIN OF LIFE. form, though in many of the humbler types, like that of the familiar Oyster, the gills are used for the double purpose of aerating the blood and, by their minute vibrating threads or cilia, drifting food to the mouth. In the great group of radiated animals, the Protozoa, Calen- terata, and JEchinodermata, no air-breathing creature exists, or, in so far as is known, has existed, so that this vast group of animals is limited altogether to the waters ; and this is undoubtedly one mark of its inferiority. In the sub-kingdom of the Mollusks the highest class, that of the Cuttle-fishes and Nautili, has been, singularly enough, rejected as unfit for this promotion, though it was early intro- duced, and attains to a high development of muscular energy and nervous power. The group next in order, that of the Snails and their allies, alone ventures in some of its families to assume the role of air-breathing. As might be expected, in creatures of this stamp the simplest means are employed to effect the result. In the sub-aquatic species the gills are contained in a chamber, where- they are protected and kept supplied with water. In the air-breathing species, this gill-chamber is merely emptied of its contents and converted into an air-sac or func- tional lung. Thus a rude and imperfect method of air-breath- ing is contrived, which scarcely separates the animals that possess it from their aquatic relatives, but which nevertheless gives to us the beautiful and varied groups of the Land-snails and of the air-breathing fresh-water Snails. In the worms and Crustaceans the gills are placed at the sides of the body, and connected with its several segments. But the Crustaceans, like the Cuttle-fishes, though the highest aquatic type, never become air-breathers. It is true some of them, like the Land-crabs, live in the air ; but they retain their gills, and have to carry with them a supply of water to keep these moist. But in order to elevate the Annulose type to the true dignity of air breathing, three new classes had to be introduced, differing THE FIRST AIR-BREATHERS. 139 altogether in their details of structure ; and all three seem to have been placed on the earth about the same time. They are : First, the Myriapods, or Gallyworms and Centipedes ; secondly, the Insects; and thirdly, the Arachnidans, or Spiders and Scorpions. In the Myriapods a system of air-tubes, kept open by elastic spiral fibres, penetrates the body by lateral pores, thus retaining the resemblance to the lateral respiration of the Crustaceans and worms. In the Insects, where this type of structure rises to its highest mechanical perfection, and where the animal is enabled to be not merely an air-breather, but a flier, the same system of lateral pores and internal air-tubes is adopted, and is so extended and ramified as to give a very perfect respi- ration. In the Spiders and Scorpions the system is the same, except that in the latter and a part of the former the whole or a part of the tracheal system becomes expanded into air- chambers simulating true lungs. Among the Vertebrates, the fishes are breathers by gills attached to arches at the sides of the neck. But already in the Devonian we have reason to believe that there were fishes having the swimming-bladder opening into the back of the mouth to receive air, and divided into chambers, so as to. con- stitute an imperfect lung. And here we have not, as in the lower types, an adaptation of the old water-breathing organs, but an entirely new apparatus. In the next grade of Verte- brates we find, as in the Frogs, Water-lizards, etc., that the young are aquatic and breathe by gills, while the adults ac- quire lungs, sometimes retaining their gills also, but in the higher forms parting with them. Thus in the vertebrates alone we have true lungs, distinct structurally from gills; and these lungs attain to their highest perfection in the birds and mammals. The oldest air breathers at present known are Scorpions and insects allied to the modern May-flies, which have been found in the Silurian. Next to these, and more important in number 140 THE CHAIN OF LIFE. and variety, are the insects of the Erian plant beds of New Brunswick. They were discovered by the late lamented Prof. C. F. Hartt in the plant-bearing shales of the Middle Devonian (Fig. 123). The beds containing them hold also a species of Eurypterus, an obscure Trilobite, and a Crustacean allied to the modern Stomapods, 1 besides a shell which may possibly be that of a Land-snail, to be mentioned in the sequel. They are also exceedingly rich in beautifully-preserved remains of FIG. 123. Wings of Devonian Insects. Middle Devonian of New Brunswick. a, F latephemerct antiqua (Scudder). b, Homothetus fossilis (Scudder). c, Lithen- tomum Harttii (Scudder). d, Xenoneura antiquor-unt (Scudder). Devonian plants. The collection made by Prof. Hartt is limited to a few fragments of wings ; but these, in the skilful hands of Mr. Scudder, h,ave proved to be rich in geological interest. One is a gigantic Efhe^tera or May-fly, which must have been five inches in tfre expanse of the wings, which are more complex in their venation than those of its modern allies (Fig. 123, a). Another presents peculiarities between those of the May-flies and Dragon-flies (Fig. 123, b). A third is a Neuropter, not belonging to any known family, but allied to some in the Coal-formation (Fig. 123, c). A fourth (Fig. 1 23, d) 1 Amphipeltis paradoxm of Salter. THE FIRST AIR-BREATHERS. 141 is a small and delicate wing, supposed to have belonged to an animal having some points of resemblance to the modern crickets. Two others are represented by mere fragments of wings, insufficient to determine their affinities with certainty. No other insects of this age have been discovered elsewhere ; but it is to be borne in mind that no other locality rich in Devonian plants has probably been so thoroughly explored. The hard slaty ridges containing these fossils are well exposed on the coast near the city of St. John, and Messrs. Hartt and Matthew of that city, acting, I believe, in concert with and aided by the Natural History Society of the place, not only searched superficially, but removed by blasting large portions of the richest beds, and examined every fragment with the greatest care. Their primary object was fossil plants, of which they obtained magnificent collections ; and it is scarcely possible that the insects could have been found but for the exhaustive methods of exploration employed. It is interesting to observe, respecting these oldest insects, that they all belong to those families which have jaws, and not suctorial apparatus, that they are not of those which undergo a complete metamorphosis, and that their modern congeners pass their larval stage in the water. Thus the waters gave birth to the first insects, and their earliest families were not of those which suck honied juices or the blood of animals, or which pass through a worm-like infancy. These groups belong apparently to much later times. On one of the specimens collected by Messrs. Hartt and Matthew, and placed by them in my hands, is a spiral form which in every particular of external marking resembles a genus of modern West Indian Land-snails. 1 I have hesitated to describe it, as the structure is lost and the form imperfect ; but I cannot help regarding it as an indication that this group of land animals also will be traced back to the Devonian age. 1 Genus Slrophia. I have provisionally named the St. John species Str ophites erianus. 142 THE CHAIN OF LIFE. Ascending from the Devonian to the Carboniferous, we at once find ourselves in the midst of air-breathers of various types. Here are Myriapods, insects of several orders, Spiders, Scorpions, Land-snails, and Batrachian reptiles, and these of many species, and found in many localities widely separated. We can thus people those dark, luxuriant forests, to which we owe our most valuable beds of coal, with many forms of life; and as most of these belong to tribes likely to multiply abundantly where food was plentiful, we can imagine multitudes of Snails and Millepedes feeding on succulent or decaying vegetable matter, swarms of insects flitting through the air in the sunnier spots, while their larvae luxuriated in decaying masses of leaves or wood, or peopled the pools and streams. In like manner, in imagination we can render these old woods vocal with the trill of crickets and with the piping or booming of smaller and larger Batrachians. Let us now, in accordance with our plan, inquire as to the nature of these early air-breathers and the fortunes of their families in the geological history. The Land-snails known as yet in the Carboniferous are limited to five or six species, belonging to four genera, all American and related to existing American forms. The two earliest known are represented in Figs. 124 and I25. 1 One of them is a Pupa, or elongated Land-snail, so similar to modern forms that it does not merit a generic distinction, and is indeed very near to some existing West Indian species. The other is in like manner a member of the modern genus Zonites. These are from the Coal-formation of Nova Scotia, and the Pupa must have been very abundant, as it has been found in considerable numbers in a layer of shale, and in the stumps of erect trees, in beds separated from each other by a thickness of 2,000 feet of strata. The Zonites is much more rare. A second Pupa is found in Nova Scotia, and two species occur in the Coal- field of Illinois. One of these is a Pupa still smaller than 1 The enlarged figure of Pupa vetusta is too much elongated, and the aperture is somewhat conjectural, as it is usually crushed. THE FIRST AIR-BREATHERS. 143 P. vetusta^ and, like some modern species, with a tooth-like process on the inner lip. The other has been placed in a new Km 124. Land-snail (Pupa -vetusia, Dawson). From the Coal -formation. , Natural size, b, Magnified c, Apex, d, Sculpture. Enlarged. genus, 1 but is very near to some of the smaller American Snails Q I FIG. 125. Land-snail (Zonites (Conulus) prisons, Carpenter). From the Coal-formation, a, Shell Enlarged ; the line below shows the natural size, b, Sculpture. Enlarged. till living. Its most special character is a plate extending from 1 Dawsonella of Bradley. 144 THE CHAIN OF LIFE. the inner lip over half the aperture, a contrivance for protec- tion still seen in some modern forms. Thus the Land-snails come on the stage in at least three generic forms, similar to those which still live, but all of small size, indicating perhaps that the conditions were less favourable for such creatures than those of the temperate and warmer climates at present. It may seem a small step in advance for Sea-snails to lose their gills and to become Land-snails, and this without any elevation of their general structure ; but it must be borne in mind that we have here not only the dropping of the gills for an air-sac, but profound changes in teeth, mucous glands, shell, and other par- ticulars, to fit them for new food and new habits. It is also singular that the Land-snails at once appear instead of the intermediate forms of the air-breathing fresh- water snails. These last may, however, yet be found. The Millepedes, like the Land-snails, were first found in the Coal-formation of Nova Scotia, but species have since been discovered not only in Illinois, but also in Great Britain and in Bohemia. In Nova Scotia alone two genera and five dis- tinct species have been found, all in the interior of erect trees, to which these creatures probably resorted for food and shelter (Fig. 126). All the species yet known are allied to the modern Gallyworms, though presenting special features which seem to separate them as a distinct family, 1 and were probably vegetable- feeders. Some of the species have the peculiarity, unknown among their modern successors, of being armed with long spines. 2 The moist, equable climate and exuberant vegeta- tion of the Coal-period would naturally be very favourable to Millepedes, and it is likely that the discoveries made as yet give but a faint idea of their actual abundance. It is not improbable that they subsequently declined, as we know of none between the Carboniferous and the Jurassic, and they do not seem to have improved up to the modern period. The 1 Archiulidce of Scudder. 2 Eupkobeua armige.a (Meek and Worthen), from Illinois. THE FIRST AIR-BREATHERS. 145 Carnivorous Myriapods, however, or Centipedes proper, a higher and essentially distinct type, are not known until much more recent times. The insects of the Carboniferous as yet known, belong to three out of the ten or more orders into which the class is divided. One of these is represented by a number of species of Cockroach, another by May-flies and a Dragon-fly, and another by some weevil-like Beetles. The Cockroach is cha- racterised by Huxley as one of the " oldest, least modified, FIG. 126- Millepedes. From the Coal-formation. , Xylobius sigilla.ria (Dawson). b, Arckiulus xylobioides (Scudder) Anterior segments. Enlarged, c, X. farctta (Scudder). Caudal portion. Enlarged. and in many ways most instructive forms of insects ; " and both he and Rolleston take its anatomy as typical of that of the class. That these creatures should have abounded in the Coal-period we need not wonder, when we consider the habits of those that infest our houses, and when we further bear in mind the number of species, some of them two inches in length, that exist in tropical climates. So* many species of this family have been found in the Coal-formation on both sides of the Atlantic, 1 that we may fairly regard them as con- stituting one of its most characteristic features, and as probably 1 About fifty in all, as I learn from Mr. Scudder. 146 THE CHAIN OF LIFE. the oldest representatives of the order to which they belong l (Fig. 127). There were also in the Coal-period insects allied to the Locusts and to the Mantids, a carnivorous group. One of the latter (Lithomantis], described by Woodward, is a mag- nificent insect, not unlike some modern tropical species. It was found in the Coal-formation of Scotland. A still larger species, probably the largest insect known, has been described by Brongniart. The May-flies (Ephemeridcz) are represented in the Carboniferous by several very large species. That of which the wing is shown in Fig. 128 must have been seven inches in FIG. 127. Wings of Cockroaches. From the Coal-formation. a, Archimulacris Acadicns (Scudder). b, Blattina Bretonensis (Scudder). c, B. Hesri (Scudder). expanse of wings. The habits of the modern May-flies show us how animals of this group, living as larvae in the streams and lakes, must have afforded large supplies of food to fishes, and when mature must have emerged from the waters in count- less myriads, filling the air for the brief term of their existence in the perfect state. The May-flies represent another insect order. 2 The Coal-measures of Saarbruck have afforded several species allied to the white ants (Termites], insects which must have found abundant scope for their activity in the dead trees Orthoptera. 2 Neuroptera. THE FIRST AIR-BREATHERS. 147 of the carboniferous forests. The occurrence of beetles, 1 espe- cially of the weevil family, which have as yet been found only in Europe, might have been expected, considering the habits and modern distribution of this group. It has been asserted that moths 2 have been found in the Carboniferous ; but the proof of this, so far as known to me, is the occurrence of leaves, noticed by Sternberg, with markings similar to those made by the larvae of minute leaf-mining moths. This, however, is uncertain evi- FIG. 128. Wing of May-fly ( H aplopklebinm Barnesii, Scudder). From the Coal-formation. dence. If we consider the orders of insects not found in the Coal-formation, we can perceive good reasons for the absence of some of them. Those containing the lice and fleas, and other minute and parasitic insects, we can scarcely expect to find. The bees and wasps, and the butterflies and moths, are little likely to have been present where there were scarcely any flowering plants ; but such groups as those of the two-winged flies, the plant-bugs and the ants, we might have expected, but 1 Coleoptera. Tinea. L 2 148 THE CHAIN OF LIFE. for the fact of their being highly specialised forms, and for that reason likely to have appeared later. 1 There are, indeed, as yet no haustellate or suctorial insects known in this early period. Plausible theories of the phylogeny of insects are not wanting ; but they do not well suit the known facts as to their first appearance ; and perhaps we may venture without FIG. 129. A Jurassic Sphinx-moth (Sphinx Snelleri, Weyenburgh). much blame to apply to the insects of the Coal-period the remark made by Wollaston with reference to the rich insect fauna of the isolated rock of St. Helena : " To a mind which, like my own, can accept the doctrine of creative acts as not necessarily * unphilosophical,' the mysteries [of the existence of these species in an island so remote from other lands], how- 1 One highly specialised Carboniferous insect recently found is the Frotophasma of Brongniart, a relative of the modern "Walking-sticks." THE FIRST AIR-BREATHERS. 149 ever great, become at least conceivable ; but those which are not able to do this may, perhaps, succeed in elaborating some special theory of their own, which, even if it does not satisfy all the requirements of the problem, may at least prove convincing to themselves." The suctorial insects make their first certain appearance in the Jurassic; and the magnificent Sphinx Moth in Fig. 129 is an example of the magnitude and perfection to which that tribe attained in the age of the Solenhofen slate ; though FIG. 130. An Eocene Butterfly (Prodryas persephone, Scudder. From Colorado. Weyenburgh, who describes it, fancies that he sees evidence that it may, unlike any modern moths, have been provided with a sting. The most perfect and beautiful fossil butterfly known to me is that represented in Fig. 130, from a photo- graph kindly given to me by Mr. Scudder. It is from the Tertiary rocks of Western America, and is laid out in stone as neatly as if prepared by an entomologist, while its preserva- tion is so perfect that even the microscopic scales on the wings can be made out. It belongs to one of the highest types* of 150 THE CHAIN OF LIFE. modern butterflies, that to which the Vanessce. belong, but with some points of structure pointing to the lower group of the " Skippers " (Hesperiadcz). Scudder remarks that while the fore-wings resemble those of the former group, the hind-wings look more like those of the latter ; and this seems to be a common character of two or three others of the few fossil species known, none of which are older than the Tertiary. We know too little of the spiders and scorpions of the Carboniferous to say more than that they closely resemble modern forms. Two of the scorpions are represented in Figs. 131 and 132; and the only spider certainly known, which is FIG. 131. Abdominal part of a Carboniferous Scorpion. 1 from Silesia, is said to belong to the group of the hunting or trap-door spiders (Lycosa)? The Batrachians of the Coal are its most characteristic and remarkable air-breathers, especially so as the precursors of the reptiles of the Mesozoic age. Cope in a recent summary enumerates no less than thirty-nine genera and about one hundred species ; and to these have to be added at least a dozen more recently discovered in Europe ; though it was only in 1841 that the first indications of such creatures were found, and were then regarded by geologists with the same scepticism which some of them still apply to Eozoon. The 1 This was first described as part of the larva of a Dragon-fly. It is now recognised as belonging to a Scorpion. - Protolvcosa (Roemer). THE FIRST AIR-BREATHERS. 151 first trace ever observed of batrachians in the Carboniferous consisted of a series of small but well-marked footprints found by the late Sir W. E. Logan in the Lower Carboniferous shales of Horton Bluff, in Nova Scotia. In that year this pains- taking geologist had examined the coal-fields of Pennsylvania and Nova Scotia, with the view of following up his important discovery of the Stigmarm, or roots of Sigillaria, as accom- paniments of the coal-underclays. On his return he read a paper, detailing his observations, before the Geological Society of London. In this he mentioned the footprints in question ; FIG. 132. Carboniferous Scorpion (Eoscorpius carbonari-us, Meek and Worthen). Illinois. but the paper was published only in abstract, and the import- ance of the discovery was overlooked for a time, the anatomists evidently being shy to acknowledge the validity of the evidence for a fact so unexpected. Fig. 133 is a representation of another slab subsequently found in beds of the same age in Nova Scotia, and which may serve to indicate the nature of Sir William's discovery. In consequence of the neglect of this first hint by the London geologists, the discovery of bones of a batrachian by von Dechen at Saarbruck in 1844, and that of footprints by King in Pennsylvania in the same year, are usually represented as the first facts of this kind. My own earliest discovery of reptilian bones in Nova Scotia was made in 1844, 152 THE CHAIN OF LIFE. though not published till some time afterward, and was fol- lowed up by further collections in company with Sir Charles Lyell in 1851, at which time also the earliest land-snail was found, and in the following year the first millepede. Since that time the progress of discovery has been astonishingly rapid, and has extended over most of the principal coal-areas on both sides of the Atlantic. FIG. 133. Footprints of one of the oldest known Batrachians, probably a species of Dendrerpeton. From the Lower Carboniferous of Parrsboro', Nova Scotia. Upper figure natural size. We may, for convenience, call these animals reptiles, but they are regarded as belonging to that lower grade of reptilian animals, the Amphibians or Batrachians, which includes the modern frogs and newts and water- lizards. 1 Still it would be doing great injustice to the carboniferous reptiles not to say, 1 Menopoma, Rlenobranchus, &c.~ THE FIRST AIR-BREATHERS. 153 that while related to this low type, they presented a much greater range of organisation than it shows at present, evincing a capability to fill most of the places now occupied by the true reptiles. Some of them were aquatic, and with limbs rudimentary or little developed, but many of them walked on the land, and were powerful and predaceous creatures. They had large and complex teeth, they were protected by external bony plates, and some of them had in addition a beautiful covering of horny plates and spines, and ornamental lappets. Many had well-developed ribs, indicating a condition of respi- ration much in advance of that in the ribless batrachians. Some of them attained to size and strength rivalling those of the modern alligators, while some of the smallest species exhibit characters approaching in some respects to the lizards. Perhaps the most fish-like of these animals are those first dis- covered by von Dechen (Archegosaurus, Fig. 134). Their long heads, short neeks, supports for gills, feeble limbs and long flat tail, show that they were aquatic creatures presenting many points of resemblance to the Ganoid fishes which must have been their companions. Yet they show what no fish can exhibit, fore and hind limbs with proper toes y and the complete series of bones that appear in our own arms and legs, while they must have had true lungs and breathed through nostrils. So different are they from the fish in details, that a single limb bone, a vertebra, a rib, or a fragment of a skull bone, suffices to distinguish them. Much has been said recently of the genesis of limbs ; and here, as far as now known, we have the first true limbs ; but it is scarcely too much to say that the feet of Archegosaurus differ more from the fins of any car- boniferous fish than they do from the human hand; while it is certain that the feet which made the impressions represented in Fig. 133, on the lowest beds of the Carboniferous, or that from the upper coal-formation represented in Fig. 139, were not less typical or perfectly formed feet than those of modern lizards. 154 THE CHAIN OF LIFE. Leaving these fish-like forms, we find the remainder of the carboniferous reptiles to diverge from them along three lines. The first leads to snake-like creatures, destitute of limbs, and which must have been functionally the representatives of FIG. -i^.Archegosaurus Decheni. Head and anterior limb reduced. Coal-field of Saarbruck. the serpents in the Palaeozoic, though batrachian in their affinities (Fig. 135). They are found both in Europe and America ; and Huxley describes one from Ireland more than FIG. 135. Ptyonius. A Snake-like Amphibian. Coal-measures of Ohio. After Cope. twenty-one inches long, and with over one hundred vertebrae. 1 Some extraordinary traces are found on the sandstones of the coal- formation, 2 which appear to indicate that there may have 1 Ophiderpeton Broivnriggii. 2 Diplichnites. THE FIRST AIR-BREATHERS. 155 been species of this type much larger than any represented by skeletons, and with bodies perhaps six inches in diameter. It is not unlikely that they had the habits of the modern water- snakes. A second line leads upward to large crocodile-like creatures, with formidable teeth, strong bony armour, and well-developed limbs (Labyrinthodontia^ Figs. 136, 137). Some of them must have attained a length of ten feet. They were lizard-like in form, could walk well, as is seen from the footprints of some of the species which present a considerable stride, and moved . 136. A large Carboniferous Labyrinthodont ( Baphetes planiceps, Owen). a, Anterior part of the skull, viewed from beneath. One^sixth natural size, b, One of the largest teeth, natural size. over mud without the belly touching the ground. Their tails were long, and probably useful in swimming. Their heads were flat and massive, and their teeth were strengthened by a remarkable folding inward of the outer plate of enamel (Fig. 137 ). The belly was protected by bony plates and closely imbricated scales. In some of the species at least the upper parts were clothed with horny scales, and the throat and sides were ornamented with pendent scaly fringes or lappets. 1 Their general aspect and mode of life must have resembled 1 These are known in some of the smaller species, but not as yet in the larger. 1 5 6 THE CHAIN OF LIFE. those of modern alligators ; and in the vast swamps of the Coal-period, full of ponds and sluggish streams swarming with fish, they must have found a most suitable abode. While rigid anatomy may ally these animals rather with the batrachians than the true reptiles, it is evident that their great size, their capacity for walking with the body borne well above the ground, their bony and scaly armour, their powerful teeth and their capacious chests, with well-developed ribs, indicate conditions FKJ. 137. Baphetes planiceps (Owen). a, Fragment of maxillary bone showing sculpture, four outer teeth, and one inner tooth. Natural size, b, Section of inner tooth. Magnified, c, Dermal scale. Natural size. of respiration and general vitality quite comparable with those of the highest modem members of the class Reptilia. The third line of progress leads to some slender and beau- tiful creatures (Microsauria), chiefly known to us by remains found in erect trees, and which resembled in form and habits the smaller modern lizards. They have simple teeth, a well- developed brain-case, limbs of some length, and bony and THE FIRST AIR-BREATHERS. 157 scaly armour, the latter in some cases highly ornate. 1 They were probably the most thoroughly terrestrial, and the most active of the coal batrachians, if indeed they were not strictly intermediate between them and the lizards proper. Fig. 138 FIG. 138. A lizard-like Amphibian (Hylonomus aciedentatus). a, Maxillary bone ; enlarged. d, Section of tooth ; magnified. b, Mandible ; enlarged. e, Scale ; natural size and magnified. c, Teeth ; magnified, showing front and side f, Pelvic bone (?) ; natural size. view of ordinary tooth and grooved g, Rib ; natural size, anterior tooth. ft, Scapular bone (?) ; natural size. i, Palate ; natural size. shows some fragments of one of these animals ; and the animal represented in Fig. 139, recently figured by Fritsch, probably belongs to this group. 1 Hylonomus. See Fig. facing this chapter. i 5 8 THE CHAIN OF LIFE. FIG. 139. Stelliosaiints longicostatus (Fritsch). Upper Coal-formation of Bohemia. The Labyrinthodonts of the Carboniferous continue upward into the Permian, where they meet with the true reptiles ; and in the earlier Mesozoic some of the largest and most typical THE FIRST AIR-BREATHERS. 159 examples are found. 1 But here their reign ceases, and they give place to reptiles of more elevated type, whose history we must consider in the next chapter. Nothing can be more remarkable than the apparently sudden and simultaneous incoming of the batrachian reptiles in the Coal-formation. As if at a given signal, they came up like the frogs of Egypt everywhere and in all varieties of form. If, as evolutionists suppose, they were developed from fishes, this must have been by some sudden change, occurring at once all over the world, unless indeed some great and unknown gap separates the Devonian from the Carboniferous a supposition which seems quite contrary to fact or unless in some region yet unexplored this change was proceeding, and at a particular time its products spread themselves over the world a supposi- tion equally improbable. In short, the hypothesis of evolution, as applied to these animals, is surrounded with geological improbabilities. A remarkable picture of the conditions of Palaeozoic land life is presented by the occurrence of remains of reptiles, millepedes and land-snails in such erect trees as that repre- sented in Fig. 140. In the now celebrated section of the South Joggins in Nova Scotia, trees of this kind occur at more than sixty different levels ; but only in one of these have they as yet been found to be rich in animal remains. Fortu- nately this bed is so well exposed and so abundant in trees, that I have myself, within a few years, removed from it about twenty of them, the greater number affording remains of land animals. The history of one of these trees may be shortly stated thus. It was a Sigillaria, perhaps two feet in diameter, and its stem had a dense and imperishable outer bark, a soft cellular inner bark liable to rapid decay, and a slender woody axis not very durable. It grew on the surface of a swamp, now represented by a bed of coal. By inundations and by subsidence, this 1 Mastodonsaurus or Labyrinthodon. i6o THE CHAIN OF LIFE. swamp was exposed to the invasion of muddy and sandy sediment, and this went on accumulating until the stem of the tree was buried to the height of about nine feet, before which time it was no doubt killed. After a time the top decayed and fell, leaving the buried stump imbedded in the sandy soil, which had now become dry, ,or nearly so. The trunk decayed, its inner bark and axis rotting away and falling in shreds into the bottom of the cylindrical hole, about nine feet deep, once occupied by the stem, and now kept open like a shaft or well by the hard resisting outer bank. The ground around this opening became clothed with ferns and reed -like Calamites, partly masking and concealing it. And now millepedes and land FIG. 140. Section showing the position of an erect Sigillaria, containing remains of land animals. 1. Underclay, with rootlets of Stigmaria, resting on gray shale, with two thin coaly seams. 2. Gray sandstone, with erect trees, Calamites, and other stems: 9 feet. 3. Coal, with erect tree on its surface : 6 inches. 4. Underclay with Stigmaria rootlets. a, Calamites. c, Stigmaria roots. b, Stem of plant undetermined. d, Erect trunk, 9 feet high. snails made the buried trunk a home, or fell into it in their wanderings ; and small reptiles sporting around, in pursuit of prey, or themselves pursued, stumbled into the open pitfall, and were unable to extricate themselves, though I have found in some of the layers in these trees trails which show that these THE FIRST AIR-BREATHERS. 161 imprisoned reptiles had wearily wandered round and round, in the vain search for means of exit, till they died of exhaustion and famine. The bones of these dead reptiles, shells of land- snails and crusts of millepedes, accumulated in these natural coffins, and became mixed with vegetable debris falling into them, and with thin layers of mud washed in by the rains ; and this process continued so long that a layer of six inches to a foot in thickness, full of bones, was sometimes produced. At length a new change supervened, the area was again inundated and drifted over with sand, and the hollow trunk was filled to the top and buried under many feet of sediment, never to be re-opened till, after the whole had been hardened into sand- stone and elevated to form a part of the modern coast, when FIG. 1403:. Section of base of erect Sigillaria, containing remains of land animals. Mineral charcoal. b, Dark-coloured sandstone, with plants, bones, &c. c, Gray sandstone, with Calamites and Cordaites. the old tree and its forest companions which had shared the same fate with it, are made to yield up their treasures to the geologist. This history is no fancy picture. It represents the results of long and careful study of the beds holding these erect trees, and of the laborious extraction of great numbers of them, and the breaking-up of their contents into thin flakes, to be carefully examined with the lens under a bright light in search of the relics they contained. Fig. n in Chap. I. represents the extraction of one of these trees, which happened to be partially exposed by the wasting of the cliff; but many others had to be laboriously mined out of the rock by blasting with gunpowder. It is evident that the combination of circumstances referred to If 162 THE CHAIN OF LIFE. above could not often occur ; and it is therefore not wonderful that only in one place and one bed has evidence of it been found, and that even in this some of the trees have been filled up at once by sand and clay, or so crushed by falling in or lateral pressure, that they could receive no animal remains. In one respect this is a striking evidence of the imperfection of the geological record, since, but for what may be called a fortunate accident, many of the most interesting inhabitants of the coal forests might have been altogether unknown to us. On the other hand, it shows how strange and unexpected are the ways in which the relics of the old world have been pre- served for our inspection, and that there is probably scarcely any animal or plant that has ever lived of which some fragment does not exist, did we know where to look for it. It may be well to remark, in closing this chapter, how many new forms of life, air-breathing and otherwise, make their first appearance in the Carboniferous, and have continued to prevail until now. Here we find the first specimens of Amphibians, Spiders, Myriapods, Orthopterous and Coleopterous Insects, and of the Crabs among ten-footed Crustaceans. In the latter group Woodward has recently described the oldest known crab, from the Coal-formation of Belgium. M 2 CHAPTER VII. THE EMPIRE OF THE GREAT REPTILES. HAD we lived in the Carboniferous period, we might have supposed that the line of the great Labyrinthodont liatrachians would have been continued onward and elevated, perhaps, in the direction of the Mammalia, to which some features of their structure point. But we should have been mistaken in this. The Labyrinthodonts, it is true, extend into the Trias ; but there is perhaps a sign of their coming degra- dation in the appearance in the Permian of the first known mud-eel, a humble Batrachian form allied to the Newts and Water-lizards. 1 Their special peculiarities are dropped in the Mesozoic in favour of those of certain small and feeble lizard- like animals, appearing first in the Carboniferous, and more manifestly in the Permian, and which are the true forerunners, though they can scarcely be the ancestors, of the magnificent reptilian species of the Mesozoic, which have caused this period to be called " the age of reptiles." The leading reptilian animal from the European Permian has long been the Proterosaurus, from the copper slates of Thuringia (Fig. 141), a reptile of lizard-like form, with well- developed limbs, and attaining a length of three or four feet. It resembles more nearly those large modern lizards known as Beinertii of Geinitz. 1 66 THE CHAIN OF LIFE. "Monitors," than any other existing form. The fore-limb represented in the figure foreshadows very closely the bones of the human arm and hand. Besides this we find in the Permian certain lizards (Theriodonts of Owen) which present the remarkable and advanced peculiarity already predicted by some Carboniferous Microsauria, 1 of having distinct canine teeth, producing a division into incisors, canines, and molars, in the manner of the Carnivorous quadrupeds, which they seem also to have resembled in some other parts of their skele- tons. It is not impossible that the foot- prints in the Permian sandstones of Scotland, which have been referred to tortoises, were those of animals of this type. Cope has recently described from the Permian of Texas a number of reptiles which have the complex dentition of the Theriodonts, and others which simulate that of Herbivorous mammals, by the possession of flat grinding teeth supposed to be adapted to vegetable food. 2 The teeth of all these Permian reptiles were set in sockets, also an advanced peculiarity. Thus already in the Permian, before the final decadence of the Carboniferous flora, and while the Palaeozoic invertebrates still lingered in the sea, the age of reptiles dawned, and gave promise of its future greatness by the as- sumption on the part of reptilian species of structures now limited to the Mammalia. But the great Mesozoic reptiles were not fully enthroned, till the Permian, an unsettled and disturbed age, characterised by great earth movements, had passed away, and until that period of continental elevation, with local deserts and desiccation, and much volcanic action, which we call the Trias, had also passed. 1 Hyleopeton. 2 Diadictes and Bolasatirus (Cope). FIG. 141. Arm of Proterosaurus Speneri. Re- duced. Permian. . THE EMPIRE OF THE GREAT REPTILES. 167 Then in the Jurassic and early Cretaceous the reptiles culmi- nated, and presented features of magnitude and structural complexity unrivalled in later times. At the same time the Labyrinthodonts disappear, or are degraded into the humble stations which the modern Batrachians now occupy. To understand the reptiles of this age, it will be necessary to notice the subdivisions of their modern representatives. The true reptiles now existing constitute the following orders : i, the Turtles and Tortoises (Chelonia) ; 2, the Snakes (Ophi- dia) ; 3, the Lizards (Lacertilia) 4, the Crocodiles and Alligators (Crocodilia). All of these, except the snakes, are well represented among Mesozoic fossils ; but we have in this middle age of the earth's geological history to add to them from five to seven orders now altogether extinct, and these not FIG. 142. Skeleton of Ichthyosaurus. Lias. England. of low and inferior organisation, but including species far in advance of any now existing both in elevation and magnitude, and constituting the veritable aristocracy of the reptile race. It will best serve our purpose here to consider chiefly these perished orders and their history, and then to notice very shortly those that now survive. The first of the extinct orders is that of the great Sea-lizards, 1 of which the now familiar Ichthyosaurus and Plesiosaurus of the English seas, to be seen in all museums and text-books, are the types (Figs. 142, 1420 and 142^). These were marine animals of large size, but not fishes or amphibians. They were true air-breathing reptiles, but with paddles for swimming instead of feet, and some of them with long flattened tails for steering and propulsion. They bore, in short, precisely 1 Enaliosauria, including Ichthyopterygia and Sauropterygia. 1 58 THE CHAIN OF LIFE. the same relation to the other members of the class Reptilia which the Whales and Porpoises bear to the ordinary quadru- peds. Some of these animals are believed to have been fifty or sixty feet in length, thus rivalling the Whales, while others FIG. 1420. Head of Pliosaurus. Jurassic. Much reduced. were of smaller dimensions, like the Porpoises and Dolphins. Some, like the Ichthyosaurus and Pliosaurus (Fig. 1420), were strongly built and powerful swimmers, and able to destroy the largest fishes, while others, like Plesiosaurus, had the body short and compact, the head small, and the neck long and FIG. 142^. Paddle of Flesiosaurus Oxoniensis. Jurassic. After Phillips. One-tenth natural size. flexible, and probably preyed on small animals near the borders of the waters. Catalogues of British fossils alone include about thirty species of Enaliosaurs, which haunted the coasts of Mesozoic Europe, a wonderful fact, when we consider the absence of these creatures from the modern seas, and the probability that only a fraction of the species are yet known to us. - THE EMPIRE OF THE GREAT REPTILES. 169 Another remarkable group is that to which Cope has given the name of Pythonomorpha, and which he regards as allied to the serpents, or as gigantic sea-serpents provided with swimming paddles, but which Owen considers more nearly connected with the lizards. In either case they constitute a group by themselves, remarkable not only on account of their anatomical affinities with animals so unlike them in general port, but also for their enormously extended length and formid- able dentition (Fig. 143). Such animals as the Mososaurus of Maestricht and Clidastes of Western America may have exceeded in length the largest Ichthyosaurs and the most bulky of living Cetaceans, though their slender forms and numerous vertebrae remind one of the semi-fabulous sea- serpent, rather than of any known animal of our modern age. They were characteristic of the Later Mesozoic, more especially of the Cretaceous period, and must have been formidable enemies to the fishes of their time. Owen has formed two orders 1 for the reception of some remarkable extinct reptiles of this age, found especially in South Africa and India, but also in Europe and America. The first includes large lizard-like animals having horny jaws like those of turtles, and in some of the species with great defensive tusks (Fig. 144). Their mode of life is not well known, but they may have been peaceable and harmless vege- table feeders. The second has been already referred to, in connection with the Permian, where it first appears, though it is continued in the Trias (Fig. 145). The resemblance of the skulls of these creatures to those of Carnivorous mammals is very striking, and nothing can be more singular than their early appearance and their decadence before the advent of those Tertiary mammals which in more modern times occupy their place. Perhaps the most extraordinary of all the Mesozoic modifi- cations of the reptilian type was that of the flying reptiles, or 1 Anomodontia and Theriodontia. * 170 THE CHAIN OF LIFE. FIG. 144. An Anomodont Reptile of the Trias (Dicynodon lacerticeps, Owen). Reduced. FiG. 145. ATheriodont Reptile of the Trias (Lycosanrns). After Owen. Reduced. FiG. 146. Skeleton of Fterodochylus crassirostris* Jurassic of Solenhofen. Reduced. THE EMPIRE OF THE GREAT REPTILES. 171 Pterodactyls. These were, in short, lizards modified for flight, somewhat in the same manner with the bats among the mam- mals. If the bat may be likened to a flying shrew-mouse, a Pterodactyl may in like manner be compared to a flying lizard ; but the modification in the latter case is by much the more remarkable, inasmuch as the lizard is a cold-blooded animal, and far less likely to be endowed with the active circulation and muscular power necessary to flight than is the mouse. In point of fact, there can be no doubt that the Pterodactyls must have been provided with some approach to a mammalian or ornithic heart, as they certainly were with great breast-muscles FIG. 147. Restoration of Rhamphorhyncus Bucklandi. Jurassic of England. After Phillips. a, One of the teeth. Natural size. attached to a keel in the breast-bone for working their large membranous wings. These wings were also somewhat original in their construction. They were not furnished with pinions? like those of the bird, but with a membrane like that of the bat, and this, instead of being stretched over four enormously lengthened fingers, as in that quadruped, was supported on a single elongated finger, corresponding, singularly enough, to the little finger, which usually inconspicuous member consti- tuted in some of these strange creatures a limb longer Jthan the whole body (Figs. 146, 147.) The other fingers of the hand were left free for walking or grasping. They are thus believed to have been able to walk as well as to fly, and even in case of 172 THE CHAIN OF LIFE. need, to swim ; while they could probably perch like birds on rocks and trees. Their heads, though very lightly framed, were large and reptilian in aspect, and furnished with sharp teeth, and sometimes probably with a beak as well. Few creatures of the old world are of more hideous and sinister aspect. Yet some of them must have been as light and graceful on the wing as swallows or sea-gulls. There are many species, most of them small, but some of those in the later Mesozoic attained to so great a size that the expanse of their wings must have ex- ceeded twenty feet, making them veritable flying dragons, FIG. 148. A Jurassic Bird ( Archceopteryx macroura). After Owen. probably formidable to all the smaller animals of their time. Though these animals were strictly reptiles, they combined in their structures contrivances for aerial locomotion now distri- buted between the bats and the birds. They had bat-like wings and bird-like chests. Some had horny beaks. All had hollow limb bones, and air cavities to give lightness to the skull. Their brains approach to those of birds, and, as already stated, their respiration and circulation must have been of a high order. These facts are very suggestive, and perhaps in no point is the imagination or the faith of the devout evolutionist more severely . THE EMPIRE OF THE GREAT REPTILES. 173 tested than in realising the spontaneous assumption of these characters oy reptiles, and their subsequent distribution between the very dissimilar types in which they are now continued. The approximation of the winged reptiles to the birds is further increased by the facts that in the Jurassic and Creta- ceous periods there were birds having reptilian tails and probably toothed jaws (Archczopteryx macroura, Fig. 148). The species just named, while in its limbs, trunk, and feathers a veritable perching bird, resembles a reptile in its head and tail. In the Cretaceous of Western America, Marsh has re- cently discovered two distinct types of toothed birds, one having the teeth in regular sockets, the other having them implanted in a groove in the jaw. One of these birds FIG. 149. Jaw of a Cretaceous Toothed Bird (Ichthyornis dispar). After Marsh. Natural size. (Ichthyo>-nis dispar, Fig. 149) was larger than a pigeon, with powerful wings constructed like those of ordinary birds. It had also the curious and old-fashioned peculiarity of bicon- cave vertebrae, like those of fishes and some reptiles. Another (Hesperornis regalis) stood fiv 7 e or six feet high, and had rudi- mentary wings like those of the Penguins. These toothed birds extend into the Eocene Tertiary, where the Odontopteryx of Owen has been known for some time. In the Eocene, however, this toothed bird is associated with others of ordinary types, allied closely to the Ostriches, the Pelicans, the Ibis, the Woodpeckers, the Hawks, the Owls, the Vultures, and the ordi- nary perching birds. In the Later Mesozoic, indeed, some reptiles became so bird-like that they nearly approached the earliest birds; but this was a final and futile effort of the reptile to obtain in the air that supremacy which it had long 174 THE CHAIN OF LIFE. enjoyed in earth and water ; and its failure was immediately succeeded in the Eocene by the appearance of a cloud of true birds, representing all the existing orders of the class. We may close our notice of the winged reptiles of the Meso- zoic by quoting from Phillips his summary of the characters of Rhamphorhyncus (Fig. I47) 1 : "Gifted with ample means of flight, able at least to perch on rocks and scuffle along the shore, perhaps competent to dive, though not so well as a palmiped bird, many fishes must have yielded to the cruel beak and sharp teeth of the Rhamphorhyncus. If we ask to FIG. 150. Jaw of Bathygnathus borealis (Leidy). A Triassic IMnosaur from Prince Edward Island. a, Cross section of second tooth, natural size, b, Fifth tooth, natural size. which of the many families of birds the analogy of structure and probable way of life would lead us to assimilate Rham- phorhyncus, the answer must point to the swimming races, with long wings, clawed feet, hooked beak, and habits of violence and voracity ; and for preference, the shortness of the legs and other circumstances may be held to claim for the Stonesfield fossil a more than fanciful similitude to the groups of Cormor- ants and other marine divers which constitute an effective part of the picturesque army of robbers of the sea." 1 Geology of Oxford, p. 227. - THE EMPIRE OF THE GREAT REPTILES. 175 Lastly, the reptiles, in this age of their imperial sway, cul- minated in the Dinosaurians, animals far above any modern Reptilia in the perfection of their organisation, and many of them of gigantic size. Just as the Pterosaurs filled the place now occupied by the birds, so the Dinosaurs filled that repre- sented by the mammals, or rather they took up a place holding some close relations with both the birds and the mammals. FIG. 151. Hadrosaurus Foiilkii (Cope). An Herbivorous Dinosaur, 28 feet long. After Hawkins's restoration. There were thus reptilian animals which on the one hand were the elephants and lions of their time, and on the other bore a grotesque resemblance to creatures so unlike these as the Ostriches, in so far as their anatomical structure was concerned while it is evident that their whole organisation places them in the highest position possible within the reptilian class. Some 1 76 THE CHAIN OF LIFE. of them must have been herbivorous, and probably slow in movement and quiet in nature. Others were carnivorous and of terrible energy, while furnished with the most destructive weapons (Figs. 152, 153). Many had the power of erecting themselves on their hind-feet and walking as bipeds ; and to adapt them to this end their hinder limbs were very large and strong, and they had long pillar- like tails, while their fore- feet were comparatively small, and used perhaps mainly for prehension (Figs. 151, 154). The size of some of these creatures was stupendous. The FIG. 152. Jaws of Megalosaurus. After Phillips. One-tenth natural size Hadrosaurus of New Jersey, an Herbivorous species (Fig. 151), when erected on its hind limbs and tail, must have stood more than twenty feet in height. Megalosaurus and Iguanodon, of the English Jurassic and Wealden, must have been of still more gigantic size. The former was a carnivorous animal, its head (Fig. 152) four or five feet in length, armed with teeth, sabre- shaped, sharp and crenate on the edges (Fig. 153), its hind limbs of enormous power, so that if our imagination does not fail us in the attempt to realise such a wonder, we may even THE EMPIRE OF THE GREAT REPTILES. 177 suppose this huge animal, much larger than the largest ele- phant, springing like a tiger on its prey, a miracle of terrible strength and ferocity, before which no living thing could stand.. Its companion, Iguanodon, was, on the contrary, a harmless herbivorous creature, using its great strength and stature as a means of obtaining leaves and fruits for food, and perhaps falling a prey to the larger Carnivorous Dinosaurs its contem- poraries. A still more bulky animal was the Cefevsaurus, so FIG. 153. Tooth of Megalosaurus. Natural size- a, Cross section, b, Crenellation of edges. Enlarged. admirably described by Phillips. Its thigh-bone measures more than five feet in length and a foot in diameter ; and it must have stood* ten feet high when on all fours, while its length must have reached forty or fifty feet. It seems from the forms of its bones to have been able to walk on land, but probably spent most of its time in the water, where it may be compared to a huge reptilian hippopotamus. Very recently some bones found in rocks, possibly of Wealden age, in N i;8 THE CHAIN OF LIFE. Western America, and described by Cope and by Marsh, indicate that even Ceteosaurus had not attained to the maxi- mum of Diriosaurian dimensions. These new animals have vertebrae twenty inches in length and from twelve inches to thirteen inches in the diameter of their bodies, while their lateral processes stretched three and a half feet. The shoulder- blade of one species is five feet in length, and its thigh-bone is six feet long. From these measurements Cope concludes that, unlike most other Dinosaurs, it had the fore-feet larger in proportion than the hind-feet, so as to have somewhat the appearance of a large giraffe. The bones of the back have a remarkable cavernous structure, which Cope interprets as indi- cating air cavities, to give lightness, as in the case of the bones of birds; but Owen suggests that the cavities were filled with cartilage, and that the animals were aquatic in their habits. Evidently in point of size the Dinosaurs had a better claim than even Behemoth to be called the " chief of the ways of God." Some of them, however, were of small size, and probably active and bird-like in their movements. One of these is the animal represented in Fig. 154, a species from the lithographic limestone of Solenhofen. 1 Nothing in the life of the Mesozoic has so seized on the imagi- nation of evolutionists as the links of connection between birds and reptiles, which has even been introduced by Huxley into the classification of animals, by his grouping these heretofore very distinct classes in one gigantic and comprehensive class of Sauropsida. It is necessary, therefore, to glance at these connections, and if possible to arrive at some conception of their true value. The links which connect the reptiles and the birds are twofold. First, that between the Dinosaurs and the ostrich tribe, 2 and, secondly, that between the Pterodactyls and 1 Cope has proposed the names Camerosaurus, Amphiccelius, &c., for these problematical animals. Marsh names them Titanosaurus ; Atlanto- saurus, &c., while Owen holds that some of them at least are identical with his genus Chondrosteosaurus. Seeley and Hulke adopt the name Ornithopsis, and support Cope's view of their nature. 2 Ratitce. THE EMPIRE OF THE GREAT REPTILES. 179 their allies, and the peculiar Mesozoic birds, such as Archa- opteryx. The first would serve to account for the few excep- tional Struthious birds of the modern world. The second would account for the Passerine and other more ordinary birds ; and thus, according to evolution, the now somewhat homo- geneous class of birds would have a double, or more probably multiple, origin from several lines of reptilian ancestors. This, no doubt, greatly complicates the links of connection, whether these be supposed to indicate derivation or not. FIG. 154. Compsognathus. One of the smaller Dinosaurs. After Wagner. If we inquire as to the first connection above stated, we may define it briefly in the words of Prof. Phillips, with reference to Megalosaurus, which "was not a ground-crawler, like the alligator, but moving with free steps, chiefly, if not solely, on the hind limbs, and claiming a curious analogy, if not some degree of affinity, with the ostrich." 1 But the 1 Woodward in a recent paper refers to a still more curious resemblance of the Dinosaurs to the biped lizard of Australia (Chlamydosaurus], which runs on its hind limbs, and even perches on trees. N 2 180 THE CHAIN OF LIFE. question arises, Was this resemblance merely that of two oviparous bipeds, or anything more? and when we set off, against the resemblance in haunch bones and hind limbs, the entire dissimilarity in head, in fore limbs, in vertebrae, in tail, and probably in external covering, we are disposed to agree with Huxley in his statement, with respect to the Struthious birds, that their " total amount of approximation to the reptilian type is but small ; and the gap between reptiles and birds is but very slightly narrowed by their existence." There is therefore here a great gap, even in the linking together of the types, independently of any question of derivation. The second line of connection appears at first sight more promising. Archaeopteryx has a reptilian tail, and claws on the wing ; and, as it had toothed jaws, like some of the birds in the Cretaceous, must have altogether made a much nearer approach to a reptile than any modern bird does. The re- markable "fish-bird" (Ichthyornis) of Marsh is also very reptilian in some of its characters. But when we compare these reptilian birds with the Pterodactyls and their allies, a vast gap at once becomes apparent. Disregarding the ex- ternal clothing, we find the wing in the two groups entirely dissimilar in details of construction, and this dissimilarity extends to the hind limbs as well, so that the Pterodactyls resemble bats rather than birds. Without committing ourselves to any doctrine of develop- ment, we might have rejoiced if our geological discoveries had established a continuous chain, or two continuous chains, of being between the reptiles and the birds ; but this end is evi- dently still far from being attained, though some approximation has undoubtedly been made. To quote again the admission of Huxley : " Birds are no more modified reptiles than reptiles are modified birds, the reptilian and ornithic types being both in reality somewhat different superstructures, raised upon one and the same ground-plan " that ground-plan being the idea of the air-breathing oviparous vertebrate, and the reptile THE EMPIRE OF THE GREAT REPTILES. 181 representing the less specialized and less ornate building. As yet the origin of that idea, and the mode of carrying it out to completion, remain unknown, except to the Architect and Builder, who may reveal them to earnest seekers for truth in His own good time. As to links of connection with the Mammalia, these are still more obscure. In the Mesozoic the mammals are represented as yet only by a few small species allied to the pouched (Mar- supial) and insectivorous quadrupeds of Australia, and these are closely linked with some of the smaller carnivorous Mam- malia of the early Tertiary ; but neither approach very closely to any known reptilian types. Nor have we yet any connecting links between the great marine reptiles and the Cetaceans and Sirenians which in the Tertiary take their place in the sea. It is an interesting fact, to come before us in our next chapter, that the great land reptiles of the Mesozoic survived long enough to become contemporary with 'the introduction and first luxuriance of the modern types of vegetation in the later Cretaceous. It would be natural to suppose that access to these great supplies of better food would have stimulated the increase and development of the herbivorous species, and would have indirectly had the same effect on those that were carnivorous ; but the opposite result seems to have followed, and in the next period the reptiles altogether gave place to the mammals, unless, indeed, they were themselves by some mysterious and comparatively rapid process transformed into Mammalia, to suit them to the better conditions of an improved world. So far as yet known, the reign of reptiles was world- wide in its time ; and the imagination is taxed to conceive of a state of things in which the seas swarmed with great reptiles on every coast, when the land was trodden by colossal reptilian bipeds and quadrupeds, in comparison with some of which our elephants are pygmies, and when the air was filled with the grotesque and formidable Pterodactyls. Yet this is no fancy 182 THE CHAIN OF LIFE. picture. It represents a time which actually existed, when that comparatively low, brutal, and insensate type of existence represented by the modern crocodiles and alligators was supreme in the world. The duration of these creatures was long, and in watching the progress of creation, they would have seemed the permanent inhabitants of the earth. Yet all have perished, and their modern successors, except a few large species existing in the warmer climates, have become subject to the more recently introduced Mammalia. How did the ancient reptile aristocracy perish? We are ignorant of the details of the catastrophe, but their final dis- appearance and replacement by the more modern fauna was connected with a great continental subsidence in the Creta- ceous age, and with changes of climate and conditions pre- ceding and subsequent to it. Yet the struggle for continued dominion was hard and protracted ; and toward its close some of the champions of the reign of reptiles were the greatest and most magnificent examples of the type ; as if they had risen in their might to defy approaching ruin. Thus some of the most stupendous forms appear in the later Cretaceous, after the great subsidence had made progress and almost at- tained its consummation. Like the antediluvian giants, they were undismayed even when the land began to sink beneath their feet ; and for them there was no ark of deliverance. LOWER CRETACEOUS LEAVES. REDUCED IN SIZE. After Lesquereux. a, A ralia Saporteana. b, Sassafras araliopsis. c, Quercus primordialis. d, Fagi4s polyclada. e, Salix protecefolia, f, Laurtts proteafolia. CHAPTER VIII. THE FIRST FORESTS OF MODERN TYPE. FOR a long time it was believed by geologists that a great and mysterious gap separated the Upper Cretaceous from the oldest Tertiary formations ; and in Western Europe, in so far as physical conditions and animal life are concerned, the severance seemed nearly complete. Oceanic deposits, like the Upper Chalk, are succeeded by beds of littoral and estuarine characters. The last and some of the greatest of the Me- sozoic Saurians have their burial-places in the Upper Cre- taceous, and appear no more on earth. The wonderful shell-fishes of the Ammonite group, and the cuttle-fishes of the Belemnite type, share the same fate. With the earliest deposits of the Eocene Tertiary came in multitudes of large Mammalia heretofore unknown, and the Cetaceans appear in the sea instead of the great marine lizards ; while shells, corals, and crustaceans of modem types swarm in the waters. Thus it is true that a great and apparently somewhat abrupt change takes place at the close of the Cretaceous, and ter- minates for ever the reptilian age. Even in regions like Western America, where physically the later Cretaceous shades gradually into the earlier Tertiary, so that there have been doubts as to the limits of these several periods, the same great change in animal life occurs. But a link of connection has at length been found in the history of the vegetable kingdom. The modern flora came 186 THE CHAIN OF LIFE. in with its full force in the later Cretaceous, before the end of the reptilian age, and continued onward to the present time. Thus the plant takes precedence of the animal, and the preparation was made for the mammalian life of the Eocene by the introduction of the modern flora in the Cre- taceous period. In like manner it is possible that the great graphite deposits of the Laurentian indicate a vegetation which preceded the swarming marine life of the Cambrian ; and it is not improbable that the Palaeozoic land flora existed long before the first land animals. Thus the plant, as in the old Mosaic record, ever appears on the day before the animal, in each stage of the development of the world. In Chapter iv. we traced the history of the old and rich vegetation of the Coal period. But this vegetation con- sisted principally of cryptogams and those lowest phaeno- gams, of the pine and cycad groups, which have naked seeds. In the modern flora we may arrange the several groups of plants, somewhat naturally, as follows : Series /., CRYPTOGAMS : Class i, Thallophytes, sea- weeds, lichens, fungi. ,, 2, Anophytes, mosses, &c. ,, 3, Acrogens, ferns, lycopods, horsetails. Series //., PH^ENOGAMS : Class 4, GymnosptrmSj pines, cycads, &c. 5, Endogens, palms, grasses, &c. ,, 6, Exogens, oaks, maples, &c. With reference to the history of these groups the record stands as follows : In the Palaeozoic age classes 3 and 4 culminated, and constituted the great mass of the arboreal vegetation. On entering the Mesozoic, No. 3 THE FIRST FORESTS OF MODERN TYPE. 187 becomes somewhat diminished, but No. 4 continues with unabated prevalence, so that the Mesozoic has sometimes been characterized as emphatically the age of Gymnosperms. With these appear some Endogens, allied to the modern Yuccas and Screw pines and Arums. But in the lower Mesozoic rocks we have no representatives of the broad- leaved Exogens (Angiosperms), which constitute the great mass of ordinary forest vegetation ; and it is only in the Cretaceous that we find them appearing in force, and that the monotonous vegetation of the older style was replaced by the more beautiful and varied forms of our modern woods. In Europe, in the lower part of the Upper Cretaceous of Bohemia (Cenomaruan), have been found some leaves which indicate the beginning of this change. These have been referred to Caesalpinias or Brasilettos, pod-bearing trees of India and tropical America, Aralias or Ginsengs, Magnolias, Laurels, an Ivy, and a peculiar and uncertain genus (Credneria). With these are noble palms, both of the types with pinnate and palmate leaves, and trees allied to the Giant Sequoias of California, and to the Araucarian pines of the' southern hemi- sphere. (See Frontispiece to this Chapter.) These ancient Cretaceous forests of Eastern Europe are compared by Saporta with those which now live in the warmer portions of China or in South America truly a marvellous change from the sombre and uniform vegetation by which they seem to have been im- mediately preceded. A still further development of modern vegetation takes place in the next or highest member of the Cretaceous, the Maestricht beds (Senonian\ where we find a crowd of modern types. On this great change Count Saporta remarks with truth that there seem to have been periods of pause and of activity in the introduction of plants. The Jurassic period was one of inactivity ; and a new and vigorous evolution, as he regards it, is introduced in the middle of the Cretaceous. This new and grand elevation of the vegetable kingdom in 1 88 THE CHAIN OF LIFE. the Cretaceous age was not local merely. In Moravia, in the Hartz, in Belgium and France, even in Greenland, the same great renewing of the face of the earth was in progress. In America it was proceeding on a grand scale, and seems to have set in earlier than in Europe. 1 In the Dakota group of the West, one of the lower members of the Cretaceous, and cover- ing a vast area, a rich angiospermous flora has been discovered by Hayden, and described by Lesquereux and Newberry, and beds of coal have been formed from its remains. In Vancouver's Island in British Columbia, Cretaceous coal measures occur, comparable in value and in the excellence of the fuel they afford with those of the true coal formation. Some of the beds of coal are eight feet in thickness, and the shales associated with them abound in leaves of exogenous trees generally similar to those still living in America. In these beds are also found mineralized trunks, which present under the microscope the familiar structures of our oaks, birches, and other modern trees. Thus all over the northern hemi- sphere the elevation of the land out of the waters of the great Cretaceous subsidence was signalized by a development of noble and exuberant forest vegetation, of the types still extant. The following list of families found in the Cretaceous, after Saporta, will show the botanist how fully our modern Exogens are represented : APETAL^E. GAMOPETAL^E. POLYPETAL^E. Myricacea. Apocynacecz. Araliacece. Cupulifercz. Ericacecz. Hamameliacecz. Betulacecz. Ebenacece, Helleborinea. Salicacecz. Myrsinecz. Magnoliacece. Morecs. Tiliacefz. Proteacecz. Celastracecz. Lauracece. Anacardiacecz, Myrtacecz. 1 A poplar occurs in Greenland, in beds held to be Lower Cretaceous. THE FIRST FORESTS OF MODERN TYPE. 189 Of the plants in this list, some, like the oaks, birches, willows, and heaths, are common and familiar members of the flora of the northern hemisphere to-day, and even of the European flora. Some, like the Magnolias, Myricas, and witch-hazels, are characteristically American, and a few, like the Proteaceae, are now confined to the southern hemisphere. Some of these families have dwindled since the Cretaceous time, so as to be represented by very few species, or at least have not advanced, while others have multiplied and prospered ; and on the whole the flora of the northern hemisphere seems to have been as rich in this early beginning of our modern forests as it is at the present day. Lesquereux's results, with reference to the American flora of the Dakota group, are very similar, and present some surprising features of resemblance to modern American forests, though he remarks that these Cretaceous trees are generally characterized by the even or unserrated edges of their leaves ; and the same remark seems to apply to the oldest Cretaceous leaves of Europe. A very singular feature of the Cretaceous flora is the number of species of some genera now represented by few or even a single species; and this is the more remarkable when we consider how few species, comparatively, of the older flora, are known to us. For example, Lesquereux, though aware of the great variability of the modern Sassafras of America, recog- nizes eight species of this genus in the Dakota Cretaceous, one of which seems to be that still living in America, so that it has continued unchanged, while the others have perished (Fig. 155). Thus this genus culminates at once in the Cre- taceous, but continues still in one of its species. Again, the tulip-tree, Liriodendron, one of the most beautiful, unique, and invariable of American trees, is represented by one sole species in the present world. There seem to be no less than four in the Dakota beds, besides others in the Cretaceous of New Jersey, and one species is found in the Tertiary of Greenland as well as in that of Europe (Fig. 156). There are 190 THE CHAIN OF LIFE. probably four or five species of plane-tree (Platanus} now extant, of which but one occurs in America, unless P. Mexicana, the Mexican plane-tree, is a good species as distinct from the ordinary, more northern, form. There are seven species, accord- ing to Lesquereux, in the Cretaceous of Dakota alone. This sort of evolution backward, or from many species to few, would FIG. 155. Sassafras cretaceum (Newberry). probably be greatly increased, had we fuller knowledge of the Cretaceous flora, as there are several genera already repre- sented by as many species as they can boast in modern times. We have already seen that this abrupt and sudden culmination of genera and families, and their subsequent decadence, is no rare thing in geology, and it connects itself with that idea of periods of creative activity which we have already had occasion to notice. THE FIRST FORESTS OF MODERN TYPE. 191 I have dwelt principally on the phsenogamous plants of the Cretaceous, as presenting the most noteworthy and new FIG. 156. Liriodendrtm primavum (Newberry). A Cretaceons Tulip-tree. features of the time ; but we must not forget that though cryp- togams were deposed from the high position they held in the Palaeozoic, they still existed; and there are more especially FIG. 157. Onoclea sensibilis. Eocene. After Newberry. many interesting species of ferns and equisetums in the Cre- aceous and Eocene rocks. These are, however, of modern 192 THE CHAIN OF LIFE. types ; and it is remarkable that some of them appear to have continued without even specific change from the later Cre- taceous up to the present time. A striking illustration of this is afforded by two ferns discovered side by side in the oldest Eocene beds 1 of the plains west of Red River, and described in Dr. G. M. Dawson's report on the 49th parallel. One of these is the well-known and very common Onoclea sensibilis (Fig. 157), or sensitive fern of Eastern America. 2 FIG. 158. Davallia. tenuifolia. Eocene. After Dawson. Natural size and enlarged. This species came into existence at latest at the close of the Cretaceous, and has apparently been continued in America up to the present time. In Europe, where it does not now live, it occurs as a fossil in Eocene beds in the Isle of Mull. The other is Davallia tenuifolia (Fig. 158), a delicate little plant belonging to a genus not now represented in America, and to a species at present found only in Asia. Yet this species also 1 By some regarded as Upper Cretaceous. 2 First recognized in American Eocene by Newberry. i THE FIRST FORESTS OF MODERN TYPE. 193 lived in America in early Eocene times, but has since been banished, though its former companion, the Onodea, still holds its ground. Such cases of specific persistence along with great changes of habitat are very instructive as to the permanence of species. Count Saporta, whose just remarks on the marvellously sudden incoming of the Cretaceous flora we have already referred to, also notices the fact that the families and genera represented in this flora are a most miscellaneous and uncon- nected assemblage, showing either the simultaneous appearance of many dissimilar types, or requiring us to believe in the existence of these and of intermediate forms for a very long period before that in which they are first found. This may, however, be placed in connection with the appearance of an exogenous tree (Syringoxylon) in the Devonian, referred to in a previous chapter. It would be a strange and now little suspected case of imperfection of the record, if it should be found that trees of this type were lurking in exceptional corners through all the vast periods between the Devonian and the Cretaceous, to burst forth in unwonted variety and luxuriance in the latter period. The new Cretaceous flora appears first in beds which had been recently elevated from the ocean of the great Cretaceous subsidence ; and when it first flourished, in temperate regions at least, the continents were of small dimensions, and broken up into groups of islands. Farther, America would seem to have had precedence of the Eastern Continent, and the Arctic of the Temperate regions. Thus on the elevation of the later Cretaceous land, plants previously established in the far north spread themselves southward, over newly-raised lands, radiating from the polar regions into Europe, Asia, and America. This seems the only way of accounting for the similarity of the plants in these distant countries. The new flora of the Upper Cretaceous in its journey southward met with a climate probably warmer than the present, yet not so warm as to prevent trees o 194 THE CHAIN OF LIFE. similar to those now living in the same latitudes from flourishing. Let us now trace this flora through the succeeding ages, in which I shall follow pretty closely some general statements made by Count De Saporta in memoirs recently published. At the beginning of the Eocene we find a humid and warm climate in Europe, with great forests of oaks, chestnuts, laurels, giant pines, and other genera, some of them still European, others Asiatic or American, and many of them survivors of the FIG. 159. Eocene Leaves. From Aix. , Quercus antecedens (Saporta). b, Diospyros pyrifolia (Saporta). c, Myrica Mathesonii (Saporta). Cretaceous (Figs. 159 to 162) ; and at the same period similar forests overspread those great plains of North America which were rising from out the Cretaceous sea, and there vast swampy beds were formed of vegetable debris^ giving origin to beds of brown coal, some of them eighteen feet in thick- ness. Then came in Europe and Asia that great subsidence under the sea, during which the Nummuline limestones were deposited, and when the old continent was resolved again into an archipelago of islands, perhaps closely connected THE FIRST FORESTS OF MODERN TYPE. 195 with more southern lands. This led to a great increase of southern forms of plants, which does not seem to have occurred to the same extent in America, where the flora FIG. 160. An Ancient Clover (Trifolinm talceogceum, Saporta). Eocene. Aix. FIG. 161. An Eocene Maple (Acer sei tianus, Saporta). Aix. FIG. 162. A European Magnoli of the Eocene (M. diante, Saporta). Aix. is more continuous, though showing a warmer climate in the older than in the newer Eocene. At this period Palms, Screw- pines, Proteaceous shrubs, Myrtles, Acacias, and other plants o 2 J 9 6 THE CHAIN OF LIFE. of the character of those of more southern climates were dominant in Europe (Fig. 163). The well-known beds of Bournemouth, in the south of England, 1 contain a rich flora of the Eocene age, perhaps of its middle period, and reminding us of the forests of sub-tropical India or Australia. Gradual elevation of the land favoured for a time the extension of these plants, and the warmth of the climate FIG. 163. Flower and Leaf of Botnbax sepultiflorum. Eocene of Aix. After Saporta. A European representative of the Silk-cotton-tree of the East Indies and Tropical America. allowed them to extend even into Arctic latitudes. But in the middle of the Eocene another subsidence occurred, which exterminated much of the Eocene flora, and was perhaps accompanied with a reduction of temperature, in which the more northern lands became covered with great forests of trees allied to the Pines. In England a remarkable deposit of 1 Described by La Harpe and Gaudin, and recently by Gardner. THE FIRST FORESTS OF MODERN TYPE. 197 this age is that of Bovey Tracey, in Devonshire, where beds of clay and brown coal have aiforded a rich flora of American and southern types. The Sequoia shown in Fig. 164 abounds at this place, and is a near relation to the celebrated " big trees " of California ; the Cinnamomum in Fig. 165 is a type equally foreign from modern England. It is a curious feature of the Bovey deposit that immediately above these Eocene beds, holding a rich flora of warm temperate character, are gla- cial clays with leaves of Arctic willows and of the dwarf birch, FIG. 164. Branch and Fruit of Sequoia Couttsice (Heer). Eocene. England. indicating a climate much more severe than that of the British Islands at present. 1 In the Miocene period the land again rose, and the northern flora spread itself southward equally over Europe, Asia, and America, so that the Miocene flora of all these regions is very similar ; and this Miocene flora continues substantially to this day in Eastern America and Eastern Asia, except that it has 1 Recent discoveries have since the publication of the first edition removed the Bovey Tracey beds from the Miocene to the Eocene. See Reports of Mr. Starkie Gardner to the British Association. I 9 8 THE CHAIN OF LIFE. been greatly reduced in number of species by the intervention of the cold glacial period ; but in Europe and Western America it has been largely replaced by other apparently more modern species. A striking result of recent discoveries is the fact that in Cretaceous and Eocene times a very warm climate prevailed FIG. 165. Cinnamomum Sckeuchzeri (Heer). Eocene. England. in the extreme Arctic regions, and trees of temperate latitudes grew there freely. In the recent Arctic expedition, Captain Fielden found in latitude 81 40', within 600 miles of the Pole, a bed of lignite from twenty-five to thirty feet in thick- ness, associated with remains of plants such as now grow only in temperate latitudes. THE FIRST FORESTS OF MODERN TYPE. 199 " From the character of the plant-remains, Dr. Heer infers that the lignite of this locality represents an ancient peat-moss, which must have been of wide extent, with reeds, sedges, birches, poplar, and certain conifers growing on its banks ; while the higher and drier ground in the neighbourhood probably supported a growth of pines and firs, with elms and hazel-bushes. The remains of water-lilies suggest the existence of a fresh-water lake in the old peat-moss, which must have remained unfrozen during a great part of the year." It is to be observed with reference to the age of these beds, that as the Later Cretaceous and Eocene flora of Europe and America migrated from the north, the plants found in the beds of that age in the temperate latitudes may really be some- what older in the Arctic regions, a fact which produces some uncertainty as to their actual age. The warmth required for the growth of luxuriant forests near the Pole might be secured by a different distribution of land and water, and of the oceanic currents, but the require- ments of plants as to light seem more difficult to meet, and it has been doubted whether species similar to those which are accustomed in modern times to regular alternations of day and night could submit to the long Arctic winter darkness. It is known, however, that in conservatories in Northern Russia plants supplied with heat and moisture can endure in winter great deprivation of light, and at Disco, in Greenland, roses and fuchsias flourish as house plants. 1 These facts show that if there were sufficient light and heat in summer, a great number of the plants of temperate latitudes could endure extreme cold and much deprivation of light in winter. ]t may be well here to inform the reader that some confusion as to the succession of the Cretaceous and Tertiary floras in America has arisen from the fact that the plants which are evidently Eocene in Greenland and America have been until lately incorrectly regarded as Miocene in Europe. In the 1 Lyell, Principles ; Brown, Ftorula Discoana. 200 THE CHAIN OF LIFE. Western States, the Dakota group of Lesquereux is overlain by 2000 feet of Cretaceous beds, containing the marine shells characteristic of that age, but no plants. But in Vancouver's Island these same Upper Cretaceous beds contain an abundant flora, which some botanists have called Tertiary for the rea- son already mentioned. Above the 2000 feet of marine beds overlying the Dakota group is the Lower Lignite group of Lesquereux, holding many fossil plants, including Palms and other evidences of a warmer climate than that of the Cretaceous, and which, constitute a Lower Eocene flora corresponding in some respects to that of Europe. This is succeeded by an Upper Lignite group, also Eocene, but representing a more temperate climate, and therefore resembling more nearly the Cretaceous flora. This is nearly identical with the so-called Miocene of Greenland, Alaska, and Mackenzie River, which the facts collected by the Canadian geologists have shown to be really Eocene. 1 But the Canadian reports containing these facts are comparatively little known in Europe, hence incorrect ideas as to the succession of these floras have been handed from one writer to another, To those who adopt extreme views as to the refrigeration of the northern hemisphere in so-called glacial times, there is great difficulty in accounting for the continued existence of the early Tertiary flora ; but if we adopt moderate views as to this, and demand merely a great subsidence, with much reduction of mean temperature, we may suppose that the plants previously existing were preserved on insular spots, whence they were ready to recolonize the land on its emergence from the sea. It seems certain, however, that our continents never regained, after the Glacial period, the exuberance of plant life which they presented in the Miocene and earlier Pliocene ; and we shall find that this statement applies to the world of animals as well as to that of plants. This reduction was more extreme in Europe than in Eastern Asia and Eastern 1 G. M. Dawson, Report on ^gth Parallel ; Reports on British Columbia. - THE FIRST FORESTS OF MODERN TYPE. 201 America, and the fact is thus accounted for in a recent lecture by Prof. Asa Gray : " I conceive that three things have conspired to this loss. First, Europe, hardly extending south of latitude 40, is all within the limits generally assigned to severe glacial action. Second, its mountains trend east and west, from the Pyrenees to the Carpathians and the Caucasus beyond, near its southern border ; and they had glaciers of their own, which must have begun their operations, and poured down the northward flanks, while the plains were still covered with forest, on the retreat from the great ice-wave coming from the north. Attacked both on front and rear, much of the forest must have perished then and there. Third, across the line of retreat of those which may have flanked the mountain-ranges, or were stationed south of them, stretched the Mediterranean, an impassable barrier. Some hardy trees may have eked out their existence on the northern shore of the Mediterranean and the Atlantic coast. But we doubt not, Taxodium and Sequoias, Magnolias and Liquidam- bars, and even Hickories and the like, were among the missing. Escape by the east, and rehabilitation from that quarter until a very late period, were apparently prevented by the prolongation of the Mediterranean to the Caspian, and thence to the Siberian ocean. If we accept the supposition of Nordenskiold, that, anterior to the Glacial period, Europe was 'bounded on the south by an ocean extending from the Atlantic over the present deserts of Sahara and Central Asia to the Pacific,' all chance of these American types having escaped from or re-entered Europe from the south and east is excluded. Europe may thus be conceived to have been for a time somewhat in the condition in which Greenland is now, and indeed to have been connected with Greenland in this or in earlier times. 1 Such a junction, cutting off access of the Gulf Stream to the Polar Sea, 1 Gray's reasoning is based on the extreme view of the Glacial period now prevalent in America, contrary, as it appears to me, to the actual facts ; but with limitations it holds good on more moderate views as well. 202 THE CHAIN OF LIFE. would, as some think, other things remaining as they are, almost of itself give glaciation to Europe. Greenland may be referred to, by way of comparison, as a country which, having undergone extreme glaciation, bears the marks of it in the extreme poverty of its flora, and in the absence of the plants to which its southern portion, extending six degrees below the Arctic Circle, might be entitled. It ought to have trees, and might support them. But since destruction by glaciation no way has been opened for their return. Europe fared much better, but suffered in its degree in a similar way. " Turning for a moment to the American continent for a con- trast, we find the land unbroken and open down to the tropic, and the mountains running north and south. The trees, when touched on the north by the on-coming refrigeration, had only to move their southern border southward, along an open way, as far as the exigency required ; and there was no impediment to their due return. Then the more southern latitude of the United States gave great advantage over Europe. On the Atlantic border, proper glaciation was felt only in the northern part, down to about latitude 40. In the interior of the country, owing doubtless to greater dryness and summer heat, the limit receded greatly northward in the Mississippi Valley, and gave only local glaciers to the Rocky Mountains ; and no volcanic' outbreaks or violent changes of any kind have here occurred since the types of our present vegetation came to the land, So our lines have been cast in pleasant places, and the goodly heritage of forest-trees is one of the consequences. " The still greater richness of North-east Asia in arboreal vege- tation may find explanation in the prevalence of particularly favourable conditions, both ante-glacial and recent. The trees of the Miocene circumpolar forest appear to have found there a secure home ; and the Japanese islands, to which most of these trees belong, must be remarkably adapted to them. The situa- tion of these islands analogous to that of Great Britain, but with the advantage of lower latitude and greater sunshine THE FIRST FORESTS OF FOREIGN TYPE. 203 their ample extent north and south, their diversified configura- tion, their proximity to the great Pacific gulf- stream, by which a vast body of warm water sweeps along their accentuated shores, and the comparatively equable diffusion of rain through- out the year, all probably conspire to the preservation and development of an originally ample inheritance." The comparative paucity in species of the west coast of America, though the Sequoias and some other forms which have perished elsewhere are retained there, is admitted to be exceptional, and not easily explained, except by the supposi- tion of peculiar local conditions affecting the comparatively narrow strip of land between the Rocky Mountains and coast ranges, and the Pacific. To such widely-distributed and varied and complex pheno- mena as those which have been discussed in the present chapter, it is impossible to do justice in the space at our command. Details in relation to them will be found in the publications of Heer, of Saporta, and of Lesquereux, and are well worthy of study by botanists, to whom alone they can be made fully intelligible. In general, with reference to now prevalent theories of derivation, they present two very dissimilar aspects. No difficulty can be greater to the evolutionist than to account for the simultaneous appearance of so many modern generic forms in the Cretaceous ; and the fact of many of the genera presenting more and more species the farther we trace them back is a strange anomaly of evolution. On the other hand, the number of species continuing un- changed from the Eocene to the Modern, the others only slightly modified, and the representative species occurring in the floras of the old and new continents, appear to many to give great support to the doctrine of gradual transformation of species. Farther facts and farther comprehension of the dif- ference between species and races will be necessary to the settlement of these questions. In the meantime it would appear that the Jurassic flora rapidly gave place, at a particular 204 THE CHAIN OF LIFE. point of geological time, to that of the modern world, and this not merely in one locality, but over the whole northern hemisphere ; and there are apparently similar facts in the southern hemisphere as well. It farther appears that each genus was at first represented by many species, and that as time went on these were gradually reduced to a few best suited to survive; and that the changes of climate and level which occurred distributed these over different parts of the con- tinents in a way at first sight very anomalous, but which Prof. Gray somewhat quaintly represents as follows : "It is as if Nature, when she had enough species of a genus to go round the four floral regions (Europe, East Asia, West America, and East America), dealt them fairly one at least to each quarter of our zone ; but when she had only two of some peculiar kind, gave one to us, and the other to Japan, Mantchuria, or the Himalayas ; and when she had only one, divided it between the two partners on the opposite sides of the table." Lastly, it seems very probable that many so-called species are nothing more than varietal forms, which may very well be modified descendants of Miocene or Eocene plants now figuring in our lists under different names. C/3 '-3 P j ! ? I .1 - CHAPTER IX. THE REIGN OF MAMMALS. THE incoming of that highest order of animals in which man himself, in so far as his physical nature is concerned, takes his place, presents some features which, though not un- paralleled in the history of other forms of life, are still very striking. The modern Mammalia are somewhat sharply divided into three very unequal groups. First, those which present in their full perfection the property of producing fully developed young, which is one of the distinctive characters of the class. These are the Placental Mammals. Secondly, those in which the young are produced in a very imperfect condition, and are usually nourished for a time in a marsupium or pouch. These are hence called Marsupials. They are for the most part con- fined to Australasia, though a few occur in America ; and are decidedly inferior in rank to the ordinary mammals. Thirdly, those in which there is a bird-like bill, and also certain bird- like or reptilian peculiarities of skeleton and of the alimentary canal. These are the Monotremes, represented by a very few species in Australia and New Guinea. In geological history, so far as the facts are at present known, the second group, that of the Marsupials, antedated the others by a vast lapse of time. The Marsupials appear in the Trias, near the beginning of the Mesozoic period. The Placentals 208 THE CHAIN OF LIFE. are not found until we reach the beginning of the Tertiary. The Monotremes would seem to be a comparatively modern degraded type. Thus the Marsupials existed throughout the reptilian age, and this in those countries of the northern hemisphere in which they are not now found. The Mesozoic Marsupials were, it is true, of small size, but there were probably numerous species, and though unable to cope with the great reptiles that swarmed by the shores and on the plains, they may have found abundant scope in the upland and interior regions of the continents. The Upper Trias of Germany has afforded to Professor Pleininger two teeth of a small mammal, to which the name of Microlestes antiquus has been given, under the impression that it was carnivorous, though it now seems more likely that it was a vegetable feeder. In rocks of nearly the same age in America, Emmons found a jaw-bone of another species (Dro- matherium sylvestre), which has been supposed to be a near ally of the existing Myrmecobius fasriatus of Australia (Figs. 166, 167). In the Stonesfield slate, a member of the English Jurassic, several other species have been found (Fig. 168), and a still larger number in the freshwater beds of the Upper Purbeck. Marsh has obtained many others from the Jurassic of America. None appear to have yet been found in the Cretaceous, but they reappear in the Eocene Tertiary, and continue to the modern time. Their absence in the Cretaceous is probably a mere accident, and they present an illustration of a very permanent type little changed since its first introduction. Lyell enumerates in all thirty-three species from the Mesozoic, all of them of small size, and all more or less nearly related to existing Australian Marsupials, though differing much among themselves, and including both carnivorous and herbivorous forms (Fig. 169). Marsh has recently suggested a somewhat new interpretation of these interesting mammalian remains. 1 He considers them divisible into two groups / one allied to 1 Geological Magazine, July, 1887. THE REIGN OF MAMMALS. 209 the modern Insectivora (Moles, Shrews, Hedgehogs, &c.), but of generalized forms. For these he constitutes a new order (Pantotheria, Marsh). The other group is less numerous and is Marsupial (Allotheria, Marsh). The jaws in Figs. 166 and 1 68 belong to the former group, that in Fig. 169 to the FIG. 166. Jaw of Dromatherium sylvestre (Emmons). From the Trias of North Carolina. latter. We should thus have both placental and Marsupial mammals in the Mesozoic. Marsh remarks that the descent of these different types from a common ancestry would require us to trace mammals back into the Palaeozoic^ that is, on the doctrine of gradual evolution. FIG. 167. Myrmecobius fasciatus. A modern Australian marsupial, allied to Mesozo c species. So soon as the palaeontologist passes from the Upper Cretaceous to the Eocene, he finds himself in the domain of the placental mammals, which appear in numerous and large species, and this, not merely in one region, but in every part of the world in which these deposits are known to exist. p 210 THE CHAIN OF LIFE. Indeed, the recent discoveries in America and in the east of Europe have almost thrown into the shade those researches of Cuvier in the Paris basin which first brought this important fact to light. The Eocene mammals, like the Carboniferous FIG. 168 Jaw, and enlarged molar of Phascolotherium Bucklandi. Stonesfield slate. England. After Phillips. amphibians, the Mesozoic reptiles, and the Cretaceous forests, appear to spring full-grown from the earth, and this at nearly the same time in every part of the northern hemisphere. It has been suggested that they may have come in gradually without our knowledge in the Cretaceous period; but if so, we should have found some of their remains along with those FIG. 169. Flagiauiax Becktesii. Jaw, and pre-molar enlarged, showing flat surface, with ridges. Purbeck. of the Upper Cretaceous plants. But the prevalence of the great reptiles up to the close of the Cretaceous would seem to render the co-existence of large mammals unlikely. It has further been supposed that geological changes in the southern THE REIGN OF MAMMALS. 211 and northern hemispheres may have alternated with each ether, so that there may be in the former Cretaceous beds in which the remains of ancestors of the Eocene mammals may be found. But we do not as yet know of such deposits. We may be content, therefore, to suppose that at the close of the Cretaceous there was established somewhere a sort of Eden for the first placental mammals, in which they were introduced and could live unharmed by the decaying monsters of the reptilian age, until the time came when they could increase and multiply and replenish the earth. The nearest approach to FIG. 170. Restoration of Palteotherinm magnum. Eocene. After Cuvier and Owen. such a centre of mammalian life is perhaps to be found in those great American lake basins embedded in the mountains of the West, which have been so well described by Hayden and Newberry, and which have yielded so many animal remains to the researches of Leidy, Marsh, and Cope. The typical deposits of the Early Eocene have long been those of the Basin of Paris, where thick and highly fossilifer- ous deposits of this age rest on the more or less denuded sur- face of the Upper Chalk, and have afforded a rich harvest ot p 2 212 THE CHAIN OF LIFE. remains of about fifty species of placental quadrupeds, whose bones have been found in the gypsum quarries of Montmartre. The great majority belong to the Ungulates, or hoofed animals, and the most abundant genera are those called by Cuvier Palaotherium (Fig. 170) and Anop father ium, of which there are several species, and which have affinities with the modern Tapirs on the one hand, and with the Horse on the other. Of the Unguiculate or clawed orders there are carnivorous forms allied to the Hyaena and the Fox, a Bat and a Squirrel ; and the Marsupials are represented by an Opossum. Lyell describes a bed of clay associated with the gypsum, in which are numerous footprints, probably produced on the margin of a lake. Many of these might be referred to the Palaeothere and its allies ; but there are others belonging to quadrupeds yet unknown, and there are also tracks of tortoises, crocodiles, and lizards, and of a large wading bird. Such a bed, perhaps deposited on the margin of a salt lake, resorted to as a " lick " by herbivorous animals, and by the carnivorous species which preyed on them, is well fitted, by the thronging life which it indicates, to teach how little we can know of the actual number and variety of the old inhabitants of the earth. In England, Eocene beds of the age of those of Paris, occupy the valley of the Thames and the Isle of Wight and neighbour- ing parts of Hants. They have afforded mammalian fossils similar to those of Paris, though less abundantly, but they are rich in remains of marine animals and of land plants. Instead of describing the well-known animals of the French and English Tertiaries, from these Eocene deposits upwards, I shall shortly sketch the succession in America, as worked out by Marsh and Cope, with the aid of the admirable summary given by Gaudry of the present state of knowledge with refer- ence to the sequence of mammalian life from its appearance in the Early Eocene up to the present time. 1 Eocene mammals, especially those gigantic whale-like 1 Les Enchainements du Monde Animal. THE REIGN OF MAMMALS. 213 creatures called Zeuglodon (Fig. 180), have been found in Eastern North America, but the most remarkable discoveries have been made in the Western Territories, where vast numbers of bones are imbedded in certain ancient and wide-spread lacustrine beds. It may be well to premise here that though the division into Eocene, Miocene, and Pliocene is recognised in America as well as in Europe, the limits of these groups may not precisely correspond with those in the Old World. Still we have this certain point of departure, that the Eocene' begins where the peculiar animals of the Cretaceous end, and that the drying up of the later Cretaceous sea and the esta- blishment of the Eocene land were probably nearly con- temporaneous in both continents. It is true, however, in animals as in plants, that in the successive periods of the Tertiary, America presents an older aspect than Europe, just as its modern fauna still contains such old forms as the opossum. It would seem that as the mountain-ranges and table-lands of Western America en:erged from the Cretaceous waters, they became clothed with Eocene forests and inhabited by Eocene mammals. But the waters, dammed up by surrounding ridges, formed large lake basins, which were drained only by the slow excavation of " canons " as the land rose still higher. In the successive deposits formed in these lakes both by ordinary deposition of silt and by paroxysmal showers of volcanic ashes were entombed great numbers of the animals which fed on their banks. It appears that these deposits, which in some places are estimated at not less than 8000 feet in thickness, hold the remains of three successive faunas, differing materially from each other, and representing the Lower, Middle, and Upper Eocene. On the flanks of the elevated region supporting the beds formed in the Eocene lakes, are other later lake basins of Miocene age, also abounding in animal remains. East of the Rocky Mountains, and also on the Pacific coast, are still later Pliocene deposits 214 THE CHAIN OF LIFE. holding other and more modern Mammalia. The vast area of these formations and the complete sequence which they show are scarcely equalled elsewhere. As in the Paris basin, the large Ungulates constitute the most Fi ;. 171 Coryphodon Ha.ma.tus. A Lower Eocene Perissodactyl skull, greatly reduced, showing small size of brain, a. After Marsh. conspicuous feature. The great group is now usually divided into those that are odd-toed (Perissodactyl) and those that are even-toed (Artiodactyl). Though; these are apparently arbitrary characters, they .correspond with other more funda- THE REIGN OF MAMMALS. 215 mental differences. The first includes such modern animals as the Rhinoceros, Tapir, and Horse. The second includes two somewhat distinct assemblages that with mammillated teeth, of which the Hog and Hippopotamus are types (Bunodonts), and that with crescental plates of enamel in the teeth, of which the Ruminants like the Deer, Ox and Camel, are examples (Selenodonts). The most characteristic animals of the lowest Eocene belong to the genus Coryphodon (Figs. 171, 172), which so abounded in Eocene America that bones of about 150 individuals were found by the Wheeler Expedition in one year in the Eocene FIG. 172. Fore-foot of Coryphodon. Greatly reduced. After Alarsh. beds of New Mexico. These animals in their dentition approached the American tapirs, except that they had great canines like the bear, while their feet resembled those of the elephant, and some of them attained the dimensions of the ox. Coryphodon is thus, as might be expected in a primal placental mammal, a creature of somewhat generalised type. Another point in which it resembles some at least of its early Tertiary contemporaries is the small size of the brain, especially in those parts of it supposed to minister to the intelligence and higher instincts (Fig. 171,0). It is certainly remarkable that as Tertiary time went on the successive groups of mammals were gifted with brains of larger and larger size, fitting them 2i6' THE CHAIN OF LIFE. for higher functions, and ultimately for associating with man. Animals thus low in development of brain were probably slow and sluggish and stubbornly ferocious, and dependent on brute force for subsistence and defence ; and they would have been altogether unsuitable for domestication had they lived to the present time. In the Middle Eocene, the place of Coryphodon was taken FIG. 173. Skull of an Upper Eocene Pefissodactyl (Dinoceras mirabilis\ showing three pairs of horn-bases. Greatly reduced. After Marsh. by Dinoceras and allied forms. Some of the species nearly equalled the elephant in size, but had shorter and stouter limbs, each supported on five great toes the most perfect possible sort of pedestal foot (Figs. 172, 174). They were heavily armed with immense canines on the upper jaws, and two or even three pairs of horns or hard protuberances on the head (Fig. 173). Creatures so supported and so armed, and THE REIGN OF MAMMALS. 217 living where food was plentiful, might well dispense with any great degree of intelligence, and their development of brain is consequently little better than that of Coryphodon. These FIG. 174. Fore-foot of Dinoceras. Greatly reduced. After Marsh great and characteristic Eocene families have no known successors ; and in the Miocene age their place is taken by a very different group, that of which Brontotherium is the "^ - ^ *^7^^ FIG. 175. Skull of Brcntotheri-itm itigens (Marsh). Greatly reduced. A Miocene Perissodactyl. ype (Fig. 175). They are creatures of huge size, with a pair of horn-cores on the nose, and feet with four toes in front and three behind, resembling in form those of the rhinoceros. 218 THE CHAIN OF LIFE. While these gigantic Perissodactyles have no successors as yet known to us, another and less conspicuous Eocene type can be traced onward to modern times by a chain of successors which the imagination of evolutionists has converted into a veritable genetic series, to which they appeal as a ''demon- stration " of the process of descent with specific modifications. In the Lower Eocene are found the remains of a diminutive ungulate (Eohippus)^ of the stature of a moderately-sized dog. It has four toes and a rudiment of a fifth in front, and three toes behind ; and has teeth slightly resembling those of the horse, but more simple and shorter in the crown. In this creature it has been supposed that we have a direct ancestor r FIG. 176. Series of Equine Feet. After Marsh. ic. 1>, Miohippus, Miocene, c. Protohippns Upper Pliocene, e, Eguus, Post-Pliocene and Modern. , Orohippus, Eocene. l>, Miohippus, Miocene, c. Protohippns, Lower Pliocene. ci, PliohippiiS) of the modern horse. A very similar genus (Orohippus), lacking only the fifth rudimentary toe, replaces Eohippus in the Middle Eocene. Mesohippus of the Lower Miocene is as large as a sheep, and has only three toes on the fore-foot and a splint bone, while its teeth assume a more equine character (Fig. 176). In the Upper Miocene Miohippus continues the line, while Protohippus of the Lower Pliocene is still more equine and as large as an ass, and corresponds with the European Hipparion in having the middle toe of each foot alone long enough to reach the ground. In the Upper Pliocene true horses appear with only a single toe, and splint bones instead of the others. In America, though the horse was THE REIGN OF MAMMALS. 219 unknown at the time of the discovery of the continent, several species occur in the Tertiary and Post-Pliocene, showing that the genus existed there up to a comparatively late period ; and when re-introduced it has thriven and run wild in the more temperate regions. What cause could have led to its extinction in Post-Glacial times is as yet a mystery. This genealogy of the horse, independently of its evolutionist application, is very interesting. It shows that some Eocene types were suited to continuance, and even adapted for extension, while others were destined to become altogether extinct at an early date. It shows farther that the power of continuance resided not so much in the gigantic and prominent species as in smaller forms. It is to be observed, however, that Gaudry and other orthodox evolutionists in Europe deduce the horse, not from Eohippus, but from Palaotherium, and that it is equally im- possible to verify either phytogeny, since the mere sequence of more or less closely allied species in time does not prove continuous derivation. Nor indeed are we certain that one- toed horses like those now living did not exist on the dry plains in Eocene times, since the inhabitants of these plains are probably unknown to us. An amusing illustration of the probable reason of the disappearance of the missing links has recently been given by a writer not very favourable to the new philosophy. The several consecutive species may be represented by coins. We may suppose, for example, sixpences to have been coined first, then seven penny and eightpenny pieces, and so on up to a shilling, then pieces representing thirteen, fourteen and fifteen pence, and so on up to a half- crown or crown ; but all the intervening denominations between the sixpence and the shilling, and between the shilling and the half-crown, were found practically of little use. Hence few were coined, and they soon became obsolete. Thus the antiquary would find only a few denominations, and those connecting them would be seldom or never found. It is plain that if we could suppose that nations constructed their 220 THE CHAIN OF LIFE, coinage after this unthinking and empirical fashion, and that if we were justified in ascribing a similar procedure to the Creator, it might help to account for the facts as we find them, otherwise we should rather suppose that in both cases something like plan and calculation determined the selection of the species produced, whether of coins or animals. But Chance is a blind goddess, and if we instal her as creator, we must expect the work to proceed by a series of abortive experiments. The Perissodactyls are not numerous at present. The three groups represented by the Horse, Rhinoceros, and Tapir constitute the whole ; and the two latter forms can be traced back to predecessors in Eocene times, even more closely resembling them than those supposed to be ancestors of the horse resemble that animal. But the few species now living have thus a vast surplusage of possible ancestors. Many species and genera are dropped without any modern repre- sentatives, so that the tendency has been to a gradual elimina- tion of surplus types, until only a few isolated and somewhat specialised forms remain at present. Yet this process of elimination is not necessarily an evolution or survival of the fittest, in the sense of modern derivationists. It rather implies that in certain past states of the earth the conditions of life afforded scope for many forms not now required, or replaced by other types more suited to the advanced and specialised nature of the world. On the other hand, the Artiodactyls have gained in numbers and importance, in comparison with their odd- toed comrades ; and this, though an odd number, namely five, was the typical number with which the earliest quadrupedal forms began life far back in the Palaeozoic. The typical Artiodactyls are those that cleave the hoof, and many of which also chew the cud ; and they are of all others, the horse perhaps excepted, those that are most valuable to man. The lower type (Bunodont), to which the hog belongs, is the older; and many hog-like THE REIGN OF MAMMALS. 221 animals occur from the earlier Tertiary upwards. In the Upper Eocene, even-toed species appear with an approach at least to the crescent-shaped teeth of the modern deer and oxen. Some of the species are obviously forerunners of the modern antelopes and deer, though as yet destitute of horns or antlers. Others, like Oreodon, are of more hog-like aspect, though believed to have been ruminants (Fig. 177). These are characteristic of the Middle Miocene, at which stage true deer appear in Europe (Dtcroceras), though they are not known in America until the Pliocene period. The earliest deer have small and simple antlers, these ornaments becoming larger and more elaborate in approaching the modern era. The FIG. 177. Oreodon major. A generalised Miocene ruminant, with affinities to the Deer, Camel, and Hog. Greatly reduced. After Leidy. hollow-horned ruminants appear for the first time in America in the Lower Pliocene ; and no ancestry has so far been attempted to be traced for them. The antelopes of this group, as well as the gigantic Sivatherium of India, 1 allied to the modern prong-horned antelope of North America, were pro- minent in the Old World in the Miocene. A very noteworthy and specially American group of mammals is that of the Edentates, the Sloths and Ant-eaters, a group which a priori we should have supposed would have been one of the earliest in time. They appear, however, first in the Miocene, without even any suggested ancestry, and are repre- 1 See Frontispiece to this Chapter. 222 THE CHAIN OF LIFE. sented from the first by large species, though they attain their grandest stature in the Megatherium and Mylodon of the Post- Pliocene (Figs. 178, 179), which were sloths of so gigantic size that they must have pulled down trees to feed on their leaves, FIG. 178. Lower Jaw of Megatherium, Greatly reduced. Post- Pliocene of South America. After Owen. unless, indeed, there were trees equally colossal for them to climb. But before the modern time, like the American horses, the larger herbivorous forms suddenly disappear, and are now FIG. 179. Ungual Phalanx and Claw-core of Megatherium. Greatly reduced. represented only by a few diminutive South American species, which can scarcely, by any stretch of imagination, be supposed to be descendants of their gigantic predecessors. The history of these animals, like those of the great Tertiary marsupials of THE REIGN OF MAMMALS. 223 Australia and the many Miocene elephants of India, affords a remarkable illustration of the persistence of similar groups of creatures in successive ages in the same region, along with diminution in magnitude and number of species toward the modern times. The Whale-tribe (Cetaceans) at once in the earliest Eocene takes the place of the great Sea-lizards of the Cretaceous ; and the oldest of the whales are in their dentition more perfect FIG. i So. Tooth of Eocene Whale (Ze2tglodon cetioides). One-half natural size. than any of their successors, since their teeth are each implanted by two roots, and have serrated crowns, like those of the Seals. The great Eocene whales of the Southern Atlantic (Zeuglodon) (Fig. 1 80), which have these characters, attained the length of seventy feet, and are undoubtedly the first of the whales in rank as well as in time. This is perhaps one of the most difficult facts to be explained on the theory of evolution. Allied to the whales is the small and peculiar group of the 224 THE CHAIN OF LIFE. Sea-cows or Dugongs (Sirenians). These creatures, highly specialised and very distinct from all others, appear in the Early Tertiary in forms very similar to those which now exist, and probably in much more numerous species, and they pursue the even tenor of their way down to modern times without perceptible elevation or degradation. " We have questioned," says Gaudry, when speaking of the Tertiary Cetaceans, " these strange and gigantic sovereigns of the Tertiary oceans as to their progenitors they leave us without reply." Their silence is the more significant as one can scarcely suppose these animals to have been nurtured in any limited or secluded space in the early stages of their "development. The true Seals, which are more elevated than the Whales, and very different in type, appear much later, and without any probable ancestry. The Elephants, two or three species of which constitute in the modern world the sole representatives of an order, are a remnant of an ancient race once vastly more numerous. They appear in Europe and Asia in the Miocene, when they were represented by three distinct genera (Elephas, Mastodon, and D mother iuni}. The second genus (Fig. 181) differs from the proper Elephants in having tuberculated teeth, indicating a more swinish habit, and probably a more fierce disposition. The third (Fig. 182) is remarkable for the immense size of some of its species, far exceeding the modern Elephants, and has the farther peculiarity of a pair of descending tusks on the lower jaw, constituting a strong and heavy grubbing-hoe, with which it could probably dig deeply for roots. So important was the group in Miocene times that seven elephants are already known from this formation in India alone, besides three species of Mastodon. Four or five Miocene Mastodons are known in Europe, besides two Dinotheria; and the true Elephants appear there in the Pliocene, and continue to the beginning of the Modern. The elephantine animals are not known in America till the Pliocene, but in that and the Pleistocene, and perhaps up to the human period, the western continent, now altogether THE REIGN OF MAMMALS. 225 226 THE CHAIN OF LIFE. destitute of elephants, possessed several species both of Elephas and Mastodon, which extended, as in Siberia, even into the Arctic regions ; and, as we know from specimens preserved in a frozen state in the latter region, some of the species were so protected by dense fur as to be able to endure extreme cold. The candid Gaudry closes his summary of the history and affinities of the elephantine animals with the words : " How- ever, the sum of the differences compared with that of the FIG. 182. Head of Dinotherium gigctnteum. Greatly reduced. Miocene of Europe. FIG. 183. Wing of Vespertilio aquensis. An Eocene Bat. After Gaudry. resemblances is too great to permit us to indicate any relation of descent between the proboscidians and the animals of other orders known to us at present." So these greatest of all the animals of the land, with their strangely specialised forms and almost human sagacity, stand alone, without father or mother, without descent. The Rodents, or gnawing animals, appear in the Early Eocene on both continents in familiar forms allied to our Squirrels and THE REIGN OF MAMMALS. . 227 Rats. Porcupines and Beavers are added in the Miocene. This group seems thus to have continued much as it was ; and it is still perhaps represented by as many species as at any previous time. Many of the ancient forms were, however, much larger than any modern species, and some of these larger forms l present singular points of approach to very distinct types, as, for example, to that of the Bears ; but these large and composite species are long since extinct. The insectivorous mammals have much the same history with the Rodents. Such highly specialised and abnormal forms as the Bats might be supposed to be modern. But, strange to say, they appear with fully developed wings both in Europe and America in the Eocene (Fig. 183). Gaudry thinks that it is " natural to suppose " that there must have been species existing previously with shorter fingers and rudimentary wings ; but there are no facts to support this supposition, which is the more questionable since the sup- posed rudimentary wings would be useless, and perhaps harm- ful to their possessors. Besides, if from the Eocene to the present the Bats have remained the same, how long would it take to develop an animal with ordinary feet, like those of a shrew, into a bat ? The Early Eocene was not altogether a time of peace in the animal world. The old carnivorous Saurians were dead and buried, but their place was taken by carnivorous mammals, allied to our modern Tigers, Hyaenas, Foxes, and Weasels. The Carnivora, however, were subordinate in the Eocene, and, as already remarked, some of them appear to be intermediate between -marsupial and placental forms a fact which evolu- tionists have noticed with much satisfaction. They appear to attain to their culmination in the Miocene, when their powers seem to be proportionate to those of the great and well-armed quadrupeds they had to deal with. To this age belongs the introduction of the terrible " Cymetar-toothed Tiger" (Mach- 1 For example, Tillotherium of the American Eocene, which was as large as a tapir 3 and in form resembled a bear. Q 2 228 THE CHAIN OF LIFE. airodus, Fig. 184). Its huge tusk-like canines and powerful limbs seem to fit it more than any other of the cat family for destructive efficiency. Yet ordinary cat-like animals were contemporary with it, and have survived it, since Machairodus disappears in the Post-Pliocene, though in previous periods it had been very widely distributed on both continents. It is a curious fact, perhaps of more significance in various ways than we yet understand, that the Dog-bear (Arctocyon\ of the oldest FIG. 184. Skull of a Cymetar-toothed Tiger ( 'Machairodus cultridens). Pliocene, France. Reduced. French Eocene, believed to be the oldest placental mammal known, though technically placed among the Carnivora, has a kind of dentition indicating that, like the modern Bears, it was really omnivorous; and its skull shows some peculiarities tending to those of the Marsupials. Much interest attaches to the first appearance of the order of Apes (Quadrumana\ or, if we take the somewhat deceptive classification favoured by some modern zoologists, the Primates^ THE REIGN OF MAMMALS. 229 including the apes and man. They begin in the Eocene, both in Europe and America, with the lowest tribe, that of the Lemurs, now confined to the island of Madagascar and parts of Africa and Southern Asia, and which may, Gaudry thinks, be modified Marsupials, though he admits that this is hard to understand. He mentions the resemblance of the teeth of monkeys to those of some hog-like animals, a resemblance, however, merely marking a similarity of food, and suggests on this ground that some of the primitive ancestors of the hog FIG. 185. Lower Jaw of Dryopithecus Fontani. An Anthropoid Ape of the Middle Miocene of France. Natural size. may have also given rise to the Monkeys. In the Miocene of Europe and Asia we have true Apes ; and one of these, which rivals man in stature (Dryopithccus), belongs to the group of the gibbons, or long-armed apes, one of the higher families of the modern Qiiadrumana (Fig. 185). This animal presents, indeed, the nearest approach to man made by any Tertiary mammal. Still the differences are great, as, for instance, in the much larger size of the canines and premolars. Yet so much con- fidence has Gaudry in the resemblances, that he even ventures 230 THE CHAIN OF LIFE. to suggest that certain flint chips found in the Miocene of Thenay, and which have been supposed to indicate human workmanship, may have been chipped by the hands of Dryopithecus. Should this view be adopted by evolutionists, it will at least have the effect of preventing flint chips from being received as evidences of the antiquity of man. It is scarcely necessary to sum up this review of the history of the Tertiary mammals. Much that has been said may be modified or changed by future discoveries ; but the great facts of the late appearance of the placental mammals, of their rapid introduction, with their ordinal differentiation nearly complete over all the continents, of the speedy culmination and early decadence of many types, and of the unchanged permanence of others, must in the main be sustained. It is not too much to say that to account for these facts the evolu- tionist must abandon the idea of gradual change, and adopt that of "critical periods" when sudden changes occurred. The history becomes inexplicable, unless with Mivart, Le Conte, and Saporta, we admit "periods of rapid evolution" alternating with others of stagnation or retrogression ; and if we admit these, we practically fall back on the old idea of creation ; only it may perhaps be " Creation by Law." CHAPTER X. THE ADVENT OF MAN. HITHERTO we have met with no trace of man or of his works. Yet (here have been in our upward progress from the dawn of life mute prophecies of his advent. Man is in his bodily frame a vertebrate animal and a mammal; and when first the Amphibians were introduced in the Palaeozoic, the framework of man's body was already sketched out and its principles settled. Those great reptilian lords, the biped Saurians of the Mesozoic, already foreshadowed his erect posture, though their limbs may have been more ornithic than mammalian. The gradual advance in the brain-development of the Tertiary mammals presaged a coming time when mind would obtain the mastery over claw and tooth and horn ; and in the Miocene ages there was already some hint of the precise style of structure in which this new creative idea would be realised. Yet it might have been impossible to imagine beforehand the vast changes which this new idea would inau- gurate. In the lower animals such intelligence as they possess is so tied to the physical organisation that it manifests itself as a mechanical unvarying instinct. Man bursts this bond, and in doing so revolutionises the whole scheme of nature. Old things are now put to new uses, the face of nature is changed, varied arts are introduced, and thought enters into the domain of 234 THE CHAIN OF LIFE. general and abstract truth. Objects are arranged, classified, understood, and while in some respects the whole creation is made to groan under the tyrannous inventions of man, yet these are the inventions of imagination and design. They are the triumph, not of brute force, but of will and intelligence. That man was not in all the earlier ages of the world, except in these prophecies of his coming, geology assures us. That he is, we know. How he came to be, is, independently of Divine revelation, an impenetrable mystery one which it is doubtful if in all its bearings science will ever be competent to solve. Yet there are legitimate scientific questions of great interest relat- ing to the time and manner of his appearance, and to the con- dition of his earlier existence and subsequent history, which belong to geology, and in which so gr