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| author | Roger Frank <rfrank@pglaf.org> | 2025-10-14 20:03:31 -0700 |
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| committer | Roger Frank <rfrank@pglaf.org> | 2025-10-14 20:03:31 -0700 |
| commit | a1bcfcbb383d07092bde8fdd3497e93776307f85 (patch) | |
| tree | a0d53552bd9ff1e3bda10a9864654d17b46dc0c5 /35316-h | |
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diff --git a/35316-h/35316-h.htm b/35316-h/35316-h.htm new file mode 100644 index 0000000..10a1c73 --- /dev/null +++ b/35316-h/35316-h.htm @@ -0,0 +1,9838 @@ +<!DOCTYPE html PUBLIC "-//W3C//DTD XHTML 1.0 Strict//EN" + "http://www.w3.org/TR/xhtml1/DTD/xhtml1-strict.dtd"> + +<html xmlns="http://www.w3.org/1999/xhtml"> + <head> + <meta http-equiv="Content-Type" content="text/html;charset=iso-8859-1" /> + <title> + The Project Gutenberg eBook of Dragons of the Air, by H. G. Seeley. + </title> + <style type="text/css"> + + p { margin-top: .75em; + text-align: justify; + text-indent: 1em; + margin-bottom: .75em; + } + h1,h2,h3,h4,h5,h6 { + text-align: center; /* all headings centered */ + margin-top: 2.5em; + clear: both; + } + hr { width: 33%; + margin-top: 2em; + margin-bottom: 2em; + margin-left: auto; + margin-right: auto; + clear: both; + } + + table {margin-left: auto; margin-right: auto;} + + body{margin-left: 10%; + margin-right: 10%; + } + + .pagenum { /* uncomment the next line for invisible page numbers */ + /* visibility: hidden; */ + position: absolute; + left: 92%; + font-size: smaller; + text-align: right; + } /* page numbers */ + + .linenum {position: absolute; top: auto; left: 4%;} /* poetry number */ + .blockquot{margin-left: 15%; margin-right: 15%;} + .sidenote {width: 20%; padding-bottom: .5em; padding-top: .5em; + padding-left: .5em; padding-right: .5em; margin-left: 1em; + float: right; clear: right; margin-top: 1em; + font-size: smaller; color: black; background: #eeeeee; border: dashed 1px;} + + .bb {border-bottom: solid 2px;} + .bl {border-left: solid 2px;} + .bt {border-top: solid 2px;} + .br {border-right: solid 2px;} + .bbox {border: solid 2px;} + .noidt {text-indent: 0em;} + + .center {text-align: center;} + .smcap {font-variant: small-caps;} + .u {text-decoration: underline;} + .ft20 {font-size: 2em;} + .ft30 {font-size: 4em;} + .ft50 {font-size: 5em;} + + .caption {font-weight: bold; text-align: center;} + + .figcenter {margin: auto; text-align: center; margin-bottom: 2em;} + + .figleft {float: left; clear: left; margin-left: 0; margin-bottom: 1em; margin-top: + 1em; margin-right: 1em; padding: 0; text-align: center;} + + .figright {float: right; clear: right; margin-left: 1em; margin-bottom: 1em; + margin-top: 1em; margin-right: 0; padding: 0; text-align: center;} + + .footnotes {border: dashed 1px;} + .footnote {margin-left: 10%; margin-right: 10%; font-size: 0.9em;} + .footnote .label {position: absolute; right: 84%; text-align: right;} + .fnanchor {vertical-align: super; font-size: .8em; text-decoration: none;} + + .poem {margin-left:10%; margin-right:10%; text-align: left;} + .poem br {display: none;} + .poem .stanza {margin: 1em 0em 1em 0em;} + .poem span.i0 {display: block; margin-left: 0em; padding-left: 3em; text-indent: -3em;} + .poem span.i2 {display: block; margin-left: 2em; padding-left: 3em; text-indent: -3em;} + .poem span.i4 {display: block; margin-left: 4em; padding-left: 3em; text-indent: -3em;} + + </style> + </head> +<body> + + +<pre> + +The Project Gutenberg EBook of Dragons of the Air, by H. G. Seeley + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: Dragons of the Air + An Account of Extinct Flying Reptiles + +Author: H. G. Seeley + +Release Date: February 18, 2011 [EBook #35316] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK DRAGONS OF THE AIR *** + + + + +Produced by Chris Curnow and the Online Distributed +Proofreading Team at http://www.pgdp.net (This file was +produced from images generously made available by The +Internet Archive) + + + + + + +</pre> + + + + +<p> </p> +<h2>DRAGONS OF THE AIR</h2> +<p> </p> +<p> </p> + + +<hr style="width: 65%;" /> +<div class="figcenter" style="width: 1024px;"> +<a name="Fig_47" id="Fig_47"></a> +<span class="caption">FIG. 47. RHAMPHORHYNCHUS PHYLLURUS<br /><br /> +SHOWING THE PRESERVATION OF THE WING MEMBRANES</span> +<img src="images/i_005.jpg" width="1024" height="614" alt="FIG. 47. RHAMPHORHYNCHUS PHYLLURUS" title="FIG. 47. RHAMPHORHYNCHUS PHYLLURUS" /> +<span class="caption"><i>From the Lithographic slate of Eichstädt, Bavaria</i></span> +<p style='text-align:right'><i>Frontispiece</i></p> +</div> + + +<hr style="width: 65%;" /> +<h1>DRAGONS OF THE AIR</h1> + +<h3>AN ACCOUNT OF<br /> +EXTINCT FLYING REPTILES</h3> + +<h5>BY</h5> + +<h2>H. G. SEELEY, F.R.S.</h2> + +<h5>PROFESSOR OF GEOLOGY IN KING'S COLLEGE, LONDON; LECTURER ON GEOLOGY<br /> +AND MINERALOGY IN THE ROYAL INDIAN ENGINEERING COLLEGE</h5> + +<h4>WITH EIGHTY ILLUSTRATIONS</h4> + + +<div class='center'> +<table border="0" cellpadding="0" cellspacing="0" summary=""> +<tr><td align='left'>"I AM A BROTHER OF DRAGONS"</td></tr> +<tr><td align='left'></td><td align='left'><i>Job</i> xxx. 29</td></tr> +</table></div> + +<h4>NEW YORK: D. APPLETON & CO.<br /> +LONDON: METHUEN & CO.<br /> +<small>1901</small></h4> + + +<hr style="width: 65%;" /> +<p><span class='pagenum'>[Pg v]</span></p> +<h2>PREFACE</h2> + + +<p>I was a student of law at a time when Sir +Richard Owen was lecturing on Extinct Fossil +Reptiles. The skill of the great master, who built +bones together as a child builds with a box of +bricks, taught me that the laws which determine the +forms of animals were less understood at that time +than the laws which govern the relations of men in +their country. The laws of Nature promised a better +return of new knowledge for reasonable study. A +lecture on Flying Reptiles determined me to attempt +to fathom the mysteries which gave new types of life +to the Earth and afterwards took them away.</p> + +<p>Thus I became the very humble servant of the +Dragons of the Air. Knowing but little about them +I went to Cambridge, and for ten years worked with +the Professor of Geology, the late Rev. Adam Sedgwick, +<small>LL.D., F.R.S.</small>, in gathering their bones from the +so-called Cambridge Coprolite bed, the Cambridge +Greensand. The bones came in thousands, battered +and broken, but instructive as better materials might +<span class='pagenum'>[Pg vi]</span> +not have been. My rooms became filled with remains +of existing birds, lizards, and mammals, which threw +light on the astonishing collection of old bones which +I assisted in bringing together for the University.</p> + +<p>In time I had something to say about Flying +Animals which was new. The story was told in +the theatre of the Royal Institution, in a series +of lectures. Some of them were repeated in several +English towns. There was still much to learn of +foreign forms of flying animals; but at last, with +the aid of the Government grant administered by the +Royal Society, and the chiefs of the great Continental +museums, I saw all the specimens in Europe.</p> + +<p>So I have again written out my lectures, with the +aid of the latest discoveries, and the story of animal +structure has lost nothing in interest as a twice-told +tale. It still presents in epitome the story of life on +the Earth. He who understands whence the Flying +Reptiles came, how they endured, and disappeared +from the Earth, has solved some of the greatest +mysteries of life. I have only contributed something +towards solving the problems.</p> + +<p>In telling my story, chiefly of facts in Nature, an +attempt is made to show how a naturalist does his +work, in the hope that perhaps a few readers will find +happiness in following the workings of the laws of +life. Such an illumination has proved to many +worth seeking, a solid return for labour, which is +<span class='pagenum'>[Pg vii]</span> +not to be marketed on the Exchange, but may be +taken freely without exhausting the treasury of +Nature's truths. Such outlines of knowledge as +here are offered to a larger public, may also, I +believe, be acceptable to students of science and +scientific men.</p> + +<p>The drawings given in illustration of the text have +been made for me by Miss E. B. Seeley.</p> + +<p style='text-align: right'>H. G. S.</p> + +<p><span class="smcap">Kensington</span>, <i>May, 1901</i></p> + +<p><span class='pagenum'>[Pg viii]</span></p> + + +<hr style="width: 65%;" /> +<p><span class='pagenum'>[Pg ix]</span></p> +<h2>CONTENTS</h2> + +<div class='center'> +<table border="0" cellpadding="4" cellspacing="0" summary=""> +<tr><td align='right'></td><td align='right'><small>PAGE</small></td></tr> +<tr><td align='center' colspan='2'><a href="#CHAPTER_I"><b>CHAPTER I.</b></a></td></tr> +<tr><td align='left'>FLYING REPTILES</td><td align='right'><a href="#Page_1">1</a></td></tr> +<tr><td align='center' colspan='2'><a href="#CHAPTER_II"><b>CHAPTER II.</b></a></td></tr> +<tr><td align='left'>HOW A REPTILE IS KNOWN</td><td align='right'><a href="#Page_4">4</a></td></tr> +<tr><td align='center' colspan='2'><a href="#CHAPTER_III"><b>CHAPTER III.</b></a></td></tr> +<tr><td align='left'>A REPTILE IS KNOWN BY ITS BONES</td><td align='right'><a href="#Page_11">11</a></td></tr> +<tr><td align='center' colspan='2'><a href="#CHAPTER_IV"><b>CHAPTER IV.</b></a></td></tr> +<tr><td align='left'>ANIMALS WHICH FLY</td><td align='right'><a href="#Page_15">15</a></td></tr> +<tr><td align='center' colspan='2'><a href="#CHAPTER_V"><b>CHAPTER V.</b></a></td></tr> +<tr><td align='left'>DISCOVERY OF THE PTERODACTYLE</td><td align='right'><a href="#Page_27">27</a></td></tr> +<tr><td align='center' colspan='2'><a href="#CHAPTER_VI"><b>CHAPTER VI.</b></a></td></tr> +<tr><td align='left'>HOW ANIMALS ARE INTERPRETED BY THEIR BONES</td><td align='right'><a href="#Page_37">37</a></td></tr> +<tr><td align='center' colspan='2'><a href="#CHAPTER_VII"><b>CHAPTER VII.</b></a></td></tr> +<tr><td align='left'>INTERPRETATION OF PTERODACTYLES BY THEIR SOFT PARTS</td><td align='right'><a href="#Page_45">45</a></td></tr> +<tr><td align='center' colspan='2'><a href="#CHAPTER_VIII"><b>CHAPTER VIII.</b></a></td></tr> +<tr><td align='left'>THE PLAN OF THE SKELETON</td><td align='right'><a href="#Page_58">58</a></td></tr> +<tr><td align='center' colspan='2'><a href="#CHAPTER_IX"><b>CHAPTER IX.</b></a></td></tr> +<tr><td align='left'>THE BACKBONE, OR VERTEBRAL COLUMN</td><td align='right'><a href="#Page_78">78</a><span class='pagenum'>[Pg x]</span></td></tr> +<tr><td align='center' colspan='2'><a href="#CHAPTER_X"><b>CHAPTER X.</b></a></td></tr> +<tr><td align='left'>THE HIP-GIRDLE AND HIND LIMB</td><td align='right'><a href="#Page_93">93</a></td></tr> +<tr><td align='center' colspan='2'><a href="#CHAPTER_XI"><b>CHAPTER XI.</b></a></td></tr> +<tr><td align='left'>SHOULDER-GIRDLE AND FORE LIMB</td><td align='right'><a href="#Page_107">107</a></td></tr> +<tr><td align='center' colspan='2'><a href="#CHAPTER_XII"><b>CHAPTER XII.</b></a></td></tr> +<tr><td align='left'>EVIDENCES OF THE ANIMAL'S HABITS FROM ITS REMAINS</td><td align='right'><a href="#Page_134">134</a></td></tr> +<tr><td align='center' colspan='2'><a href="#CHAPTER_XIII"><b>CHAPTER XIII.</b></a></td></tr> +<tr><td align='left'>ANCIENT ORNITHOSAURS FROM THE LIAS</td><td align='right'><a href="#Page_143">143</a></td></tr> +<tr><td align='center' colspan='2'><a href="#CHAPTER_XIV"><b>CHAPTER XIV.</b></a></td></tr> +<tr><td align='left'>ORNITHOSAURS FROM THE MIDDLE SECONDARY ROCKS</td><td align='right'><a href="#Page_153">153</a></td></tr> +<tr><td align='center' colspan='2'><a href="#CHAPTER_XV"><b>CHAPTER XV.</b></a></td></tr> +<tr><td align='left'>ORNITHOSAURS FROM THE UPPER SECONDARY ROCKS</td><td align='right'><a href="#Page_172">172</a></td></tr> +<tr><td align='center' colspan='2'><a href="#CHAPTER_XVI"><b>CHAPTER XVI.</b></a></td></tr> +<tr><td align='left'>CLASSIFICATION OF THE ORNITHOSAURIA</td><td align='right'><a href="#Page_187">187</a></td></tr> +<tr><td align='center' colspan='2'><a href="#CHAPTER_XVII"><b>CHAPTER XVII.</b></a></td></tr> +<tr><td align='left'>FAMILY RELATIONS OF PTERODACTYLES TO ANIMALS WHICH LIVED WITH THEM</td><td align='right'><a href="#Page_196">196</a></td></tr> +<tr><td align='center' colspan='2'><a href="#CHAPTER_XVIII"><b>CHAPTER XVIII.</b></a></td></tr> +<tr><td align='left'>HOW PTERODACTYLES MAY HAVE ORIGINATED</td><td align='right'><a href="#Page_213">213</a></td></tr> +<tr><td align='center' colspan='2'> </td></tr> +<tr><td align='left'><a href="#APPENDIX">APPENDIX</a></td><td align='right'><a href="#Page_231">231</a></td></tr> +<tr><td align='center' colspan='2'> </td></tr> +<tr><td align='left'><a href="#INDEX">INDEX</a></td><td align='right'><a href="#Page_233">233</a></td></tr> +</table></div> + + + +<hr style="width: 65%;" /> +<p><span class='pagenum'>[Pg xi]</span></p> +<h2>LIST OF ILLUSTRATIONS</h2> + + + +<div class='center'> +<table border="0" cellpadding="4" cellspacing="0" summary=""> +<tr><td align='right'><small>FIG.</small></td><td align='right'></td><td align='right'><small>PAGE</small></td></tr> +<tr><td align='right'><a href="#Fig_47">47.</a></td><td align='left'>Wings of Rhamphorhynchus</td><td align='right'><i>Frontispiece</i></td></tr> +<tr><td align='right'><a href="#Fig_1">1.</a></td><td align='left'>Lung of the lung-fish Ceratodus</td><td align='right'><a href="#Page_5">5</a></td></tr> +<tr><td align='right'><a href="#Fig_2">2.</a></td><td align='left'>Attachment of the lower jaw in a Mammal and in a Pterodactyle</td><td align='right'><a href="#Page_12">12</a></td></tr> +<tr><td align='right'><a href="#Fig_3">3.</a></td><td align='left'>Chaldæan Dragon</td><td align='right'><a href="#Page_15">15</a></td></tr> +<tr><td align='right'><a href="#Fig_4">4.</a></td><td align='left'>Winged human figure from the Temple of Ephesus</td><td align='right'><a href="#Page_16">16</a></td></tr> +<tr><td align='right'><a href="#Fig_5">5.</a></td><td align='left'>Flying fish Exocœtus</td><td align='right'><a href="#Page_18">18</a></td></tr> +<tr><td align='right'><a href="#Fig_6">6.</a></td><td align='left'>Flying Frog</td><td align='right'><a href="#Page_19">19</a></td></tr> +<tr><td align='right'><a href="#Fig_7">7.</a></td><td align='left'>Flying Lizard (Draco)</td><td align='right'><a href="#Page_20">20</a></td></tr> +<tr><td align='right'><a href="#Fig_8">8.</a></td><td align='left'>Birds in flight</td><td align='right'><a href="#Page_22">22</a></td></tr> +<tr><td align='right'><a href="#Fig_9">9.</a></td><td align='left'>Flying Squirrel (Pteromys)</td><td align='right'><a href="#Page_24">24</a></td></tr> +<tr><td align='right'><a href="#Fig_10">10.</a></td><td align='left'>Bats, flying and walking</td><td align='right'><a href="#Page_25">25</a></td></tr> +<tr><td align='right'><a href="#Fig_11">11.</a></td><td align='left'>Skeleton of <i>Pterodactylus longirostris</i></td><td align='right'><a href="#Page_28">28</a></td></tr> +<tr><td align='right'><a href="#Fig_12">12.</a></td><td align='left'>The skeleton restored</td><td align='right'><a href="#Page_29">29</a></td></tr> +<tr><td align='right'><a href="#Fig_13">13.</a></td><td align='left'>The animal form restored</td><td align='right'><a href="#Page_30">30</a></td></tr> +<tr><td align='right'><a href="#Fig_14">14.</a></td><td align='left'>Fore limbs in four types of mammals</td><td align='right'><a href="#Page_38">38</a></td></tr> +<tr><td align='right'><a href="#Fig_15">15.</a></td><td align='left'>Pneumatic foramen in Pterodactyle bone</td><td align='right'><a href="#Page_46">46</a></td></tr> +<tr><td align='right'><a href="#Fig_16">16.</a></td><td align='left'>Lungs of the bird Apteryx</td><td align='right'><a href="#Page_48">48</a></td></tr> +<tr><td align='right'><a href="#Fig_17">17.</a></td><td align='left'>Air cells in the body of an Ostrich</td><td align='right'><a href="#Page_49">49</a></td></tr> +<tr><td align='right'><a href="#Fig_18">18.</a></td><td align='left'>Lung of a Chameleon</td><td align='right'><a href="#Page_51">51</a></td></tr> +<tr><td align='right'><a href="#Fig_19">19.</a></td><td align='left'>Brain in Pterodactyle, Mammal, Bird, and Reptiles</td><td align='right'><a href="#Page_53">53</a></td></tr> +<tr><td align='right'><a href="#Fig_20">20.</a></td><td align='left'>Skull of Kingfisher and Rhamphorhynchus</td><td align='right'><a href="#Page_63">63</a></td></tr> +<tr><td align='right'><a href="#Fig_21">21.</a></td><td align='left'>Skull of Heron and Rhamphorhynchus</td><td align='right'><a href="#Page_65">65</a></td></tr> +<tr><td align='right'><a href="#Fig_22">22.</a></td><td align='left'>Palate of Macrocercus and ? Campylognathus</td><td align='right'><a href="#Page_71">71</a></td></tr> +<tr><td align='right'><a href="#Fig_23">23.</a></td><td align='left'>Lower jaw of Echidna and Ornithostoma</td><td align='right'><a href="#Page_76">76</a></td></tr> +<tr><td align='right'><a href="#Fig_24">24.</a></td><td align='left'>First two neck vertebræ of Ornithocheirus</td><td align='right'><a href="#Page_81">81</a><span class='pagenum'>[Pg xii]</span></td></tr> +<tr><td align='right'><a href="#Fig_25">25.</a></td><td align='left'>Middle neck vertebræ of Ornithocheirus</td><td align='right'><a href="#Page_83">83</a></td></tr> +<tr><td align='right'><a href="#Fig_26">26.</a></td><td align='left'>Back vertebra of Ornithocheirus and Crocodile</td><td align='right'><a href="#Page_86">86</a></td></tr> +<tr><td align='right'><a href="#Fig_27">27.</a></td><td align='left'>Sacrum, with hip bones, of Rhamphorhynchus</td><td align='right'><a href="#Page_88">88</a></td></tr> +<tr><td align='right'><a href="#Fig_28">28.</a></td><td align='left'>Extremity of tail of <i>Rhamphorhynchus phyllurus</i></td><td align='right'><a href="#Page_91">91</a></td></tr> +<tr><td align='right'><a href="#Fig_29">29.</a></td><td align='left'>Hip-girdle bones in Apteryx and Rhamphorhynchus</td><td align='right'><a href="#Page_95">95</a></td></tr> +<tr><td align='right'><a href="#Fig_30">30.</a></td><td align='left'>Pelvis with prepubic bone in Pterodactylus</td><td align='right'><a href="#Page_96">96</a></td></tr> +<tr><td align='right'><a href="#Fig_31">31.</a></td><td align='left'>Pelvis with prepubic bones in Rhamphorhynchus</td><td align='right'><a href="#Page_97">97</a></td></tr> +<tr><td align='right'><a href="#Fig_32">32.</a></td><td align='left'>Pelvis of an Alligator seen from below</td><td align='right'><a href="#Page_98">98</a></td></tr> +<tr><td align='right'><a href="#Fig_33">33.</a></td><td align='left'>Femora: Echidna, Ornithocheirus, Ursus</td><td align='right'><a href="#Page_100">100</a></td></tr> +<tr><td align='right'><a href="#Fig_34">34.</a></td><td align='left'>Tibia and fibula: Dimorphodon and Vulture</td><td align='right'><a href="#Page_102">102</a></td></tr> +<tr><td align='right'><a href="#Fig_35">35.</a></td><td align='left'>Metatarsus and digits in three Pterodactyles</td><td align='right'><a href="#Page_104">104</a></td></tr> +<tr><td align='right'><a href="#Fig_36">36.</a></td><td align='left'>Sternum in Cormorant and Rhamphorhynchus</td><td align='right'><a href="#Page_108">108</a></td></tr> +<tr><td align='right'><a href="#Fig_37">37.</a></td><td align='left'>Sternum in Ornithocheirus</td><td align='right'><a href="#Page_109">109</a></td></tr> +<tr><td align='right'><a href="#Fig_38">38.</a></td><td align='left'>Shoulder-girdle bones in a bird and three Pterodactyles</td><td align='right'><a href="#Page_113">113</a></td></tr> +<tr><td align='right'><a href="#Fig_39">39.</a></td><td align='left'>The Notarium from the back of Ornithocheirus</td><td align='right'><a href="#Page_115">115</a></td></tr> +<tr><td align='right'><a href="#Fig_40">40.</a></td><td align='left'>The shoulder-girdle of Ornithocheirus</td><td align='right'><a href="#Page_115">115</a></td></tr> +<tr><td align='right'><a href="#Fig_41">41.</a></td><td align='left'>Humerus of Pigeon and Ornithocheirus</td><td align='right'><a href="#Page_119">119</a></td></tr> +<tr><td align='right'><a href="#Fig_42">42.</a></td><td align='left'>Fore-arm of Golden Eagle and Dimorphodon</td><td align='right'><a href="#Page_120">120</a></td></tr> +<tr><td align='right'><a href="#Fig_43">43.</a></td><td align='left'>Wrist bones of Ornithocheirus</td><td align='right'><a href="#Page_124">124</a></td></tr> +<tr><td align='right'><a href="#Fig_44">44.</a></td><td align='left'>Clawed digits of the hand in two Pterodactyles</td><td align='right'><a href="#Page_125">125</a></td></tr> +<tr><td align='right'><a href="#Fig_45">45.</a></td><td align='left'>Claw from the hand of Ornithocheirus</td><td align='right'><a href="#Page_129">129</a></td></tr> +<tr><td align='right'><a href="#Fig_46">46.</a></td><td align='left'>The hand in Archæopteryx and the Ostrich</td><td align='right'><a href="#Page_130">130</a></td></tr> +<tr><td align='right'><a href="#Fig_48">48.</a></td><td align='left'>Slab of Lias with bones of Dimorphodon</td><td align='right'><i>To face page</i> <a href="#Page_143">143</a></td></tr> +<tr><td align='right'><a href="#Fig_49">49.</a></td><td align='left'>Dimorphodon (restored form) at rest</td><td align='right'><a href="#Page_144">144</a></td></tr> +<tr><td align='right'><a href="#Fig_50">50.</a></td><td align='left'>Dimorphodon (restored form of the animal)</td><td align='right'><i>To face page</i> <a href="#Page_145">145</a></td></tr> +<tr><td align='right'><a href="#Fig_51">51.</a></td><td align='left'>Dimorphodon skeleton, walking as a quadruped</td><td align='right'><i>To face page</i> <a href="#Page_146">146</a></td></tr> +<tr><td align='right'><a href="#Fig_52">52.</a></td><td align='left'>Dimorphodon skeleton as a biped</td><td align='right'><i>To face page</i> <a href="#Page_147">147</a></td></tr> +<tr><td align='right'><a href="#Fig_53">53.</a></td><td align='left'>Lower jaw of Dorygnathus</td><td align='right'><a href="#Page_149">149</a></td></tr> +<tr><td align='right'><a href="#Fig_54">54.</a></td><td align='left'>Dimorphodon (wing membranes spread for flight)</td><td align='right'><i>To face page</i> <a href="#Page_150">150</a></td></tr> +<tr><td align='right'><a href="#Fig_55">55.</a></td><td align='left'>Pelvis of Dimorphodon</td><td align='right'><a href="#Page_151">151</a></td></tr> +<tr><td align='right'><a href="#Fig_56">56.</a></td><td align='left'>Rhamphorhynchus skeleton (restored)</td><td align='right'><a href="#Page_161">161</a></td></tr> +<tr><td align='right'><a href="#Fig_57">57.</a></td><td align='left'>Scaphognathus (restoration of 1875)</td><td align='right'><a href="#Page_163">163</a></td></tr> +<tr><td align='right'><a href="#Fig_58">58.</a></td><td align='left'>Six restorations of Ornithosaurs</td><td align='right'><a href="#Page_164">164</a></td></tr> +<tr><td align='right'><a href="#Fig_59">59.</a></td><td align='left'>Ptenodracon skeleton (restored)</td><td align='right'><a href="#Page_167">167</a></td></tr> +<tr><td align='right'><a href="#Fig_60">60.</a></td><td align='left'><i>Cycnorhamphus suevicus</i> slab with bones</td><td align='right'><i>To face page</i> <a href="#Page_168">168</a></td></tr> +<tr><td align='right'><a href="#Fig_61">61.</a></td><td align='left'><i>Cycnorhamphus suevicus</i> (form of the animal)</td><td align='right'><i>To face page</i> <a href="#Page_169">169</a></td></tr> +<tr><td align='right'><a href="#Fig_62">62.</a></td><td align='left'><i>Cycnorhamphus suevicus</i> skeleton (restored)</td><td align='right'><a href="#Page_170">170</a><span class='pagenum'>[Pg xiii]</span></td></tr> +<tr><td align='right'><a href="#Fig_63">63.</a></td><td align='left'><i>Cycnorhamphus Fraasi</i> (restored skeleton form of the animal)</td><td align='right'><i>To face page</i> <a href="#Page_170">170</a></td></tr> +<tr><td align='right'><a href="#Fig_64">64.</a></td><td align='left'><i>Cycnorhamphus Fraasi</i> (restoration of the form of the body)</td><td align='right'><i>To face page</i> <a href="#Page_171">171</a></td></tr> +<tr><td align='right'><a href="#Fig_65">65.</a></td><td align='left'>Neck vertebra of Doratorhynchus from the Purbeck</td><td align='right'><a href="#Page_173">173</a></td></tr> +<tr><td align='right'><a href="#Fig_66">66.</a></td><td align='left'>Neck bone of Ornithodesmus from the Wealden</td><td align='right'><a href="#Page_173">173</a></td></tr> +<tr><td align='right'><a href="#Fig_67">67.</a></td><td align='left'>Sternum of Ornithodesmus, seen from the front</td><td align='right'><a href="#Page_175">175</a></td></tr> +<tr><td align='right'><a href="#Fig_68">68.</a></td><td align='left'>Sternum of Ornithodesmus, side view, showing the keel</td><td align='right'><a href="#Page_175">175</a></td></tr> +<tr><td align='right'><a href="#Fig_69">69.</a></td><td align='left'>Diagram of known parts of skull of Ornithocheirus</td><td align='right'><a href="#Page_177">177</a></td></tr> +<tr><td align='right'><a href="#Fig_70">70.</a></td><td align='left'>Neck bone of Ornithocheirus</td><td align='right'><a href="#Page_179">179</a></td></tr> +<tr><td align='right'><a href="#Fig_71">71.</a></td><td align='left'>Jaws of Ornithocheirus from the Chalk</td><td align='right'><a href="#Page_180">180</a></td></tr> +<tr><td align='right'><a href="#Fig_72">72.</a></td><td align='left'>Palate of the English Toothless Pterodactyle</td><td align='right'><a href="#Page_181">181</a></td></tr> +<tr><td align='right'><a href="#Fig_73">73.</a></td><td align='left'>Two views of the skull of Ornithostoma (Pteranodon)</td><td align='right'><a href="#Page_182">182</a></td></tr> +<tr><td align='right'><a href="#Fig_74">74.</a></td><td align='left'>Skeleton of Ornithostoma</td><td align='right'><a href="#Page_183">183</a></td></tr> +<tr><td align='right'><a href="#Fig_75">75.</a></td><td align='left'>Comparison of six skulls of Ornithosaurs</td><td align='right'><a href="#Page_192">192</a></td></tr> +<tr><td align='right'><a href="#Fig_76">76.</a></td><td align='left'>Pelvis of Ornithostoma</td><td align='right'><a href="#Page_195">195</a></td></tr> +<tr><td align='right'><a href="#Fig_77">77.</a></td><td align='left'>Skull of Anchisaurus and Dimorphodon</td><td align='right'><a href="#Page_199">199</a></td></tr> +<tr><td align='right'><a href="#Fig_78">78.</a></td><td align='left'>Skull of Ornithosuchus and Dimorphodon</td><td align='right'><a href="#Page_201">201</a></td></tr> +<tr><td align='right'><a href="#Fig_79">79.</a></td><td align='left'>The pelvis in Ornithosaur and Dinosaur</td><td align='right'><a href="#Page_204">204</a></td></tr> +<tr><td align='right'><a href="#Fig_80">80.</a></td><td align='left'>The prepubic bones in Dimorphodon and Iguanodon</td><td align='right'><a href="#Page_206">206</a></td></tr> +</table></div> + +<div class="blockquot"><p class="noidt">These figures are greatly reduced in size, and when two or more bones are shown +in the same figure all are brought to the same size to facilitate the comparison.</p></div> + +<p><span class='pagenum'>[Pg xiv]</span></p> + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_1" id="Page_1">[Pg 1]</a></span></p> +<h1>DRAGONS OF THE AIR</h1> + + + +<hr style="width: 15%;" /> +<h2><a name="CHAPTER_I" id="CHAPTER_I"></a><small>CHAPTER I</small><br /><br /> + +FLYING REPTILES</h2> + + +<p>The history of life on the earth during the +epochs of geological time unfolds no more +wonderful discovery among types of animals which +have become extinct than the family of fossils known +as flying reptiles. Its coming into existence, its +structure, and passing away from the living world +are among the great mysteries of Nature.</p> + +<p>The animals are astonishing in their plan of construction. +In aspect they are unlike birds and beasts +which, in this age, hover over land and sea. They +gather into themselves in the body of a single individual, +structures which, at the present day, are +among the most distinctive characters of certain +mammals, birds, and reptiles.</p> + +<p>The name "flying reptile" expresses this anomaly. +Its invention is due to the genius of the great French +naturalist Cuvier, who was the first to realise that this +extinct animal, entombed in slabs of stone, is one of +the wonders of the world.</p> + +<p>The word "reptile" has impressed the imagination +with unpleasant sound, even when the habits of the<span class='pagenum'><a name="Page_2" id="Page_2">[Pg 2]</a></span> +animals it indicates are unknown. It is familiarly +associated with life which is reputed venomous, and +is creeping and cold. Its common type, the serpent, +in many parts of the world takes a yearly toll of +victims from man and beast, and has become the +representative of silent, active strength, dreaded craft, +and danger.</p> + +<p>Science uses the word "reptile" in a more exact +way, to define the assemblage of cold-blooded animals +which in familiar description are separately named +serpents, lizards, turtles, hatteria, and crocodiles.</p> + +<p>Turtles and the rest of them survive from great +geological antiquity. They present from age to age +diversity of aspect and habit, and in unexpected +differences of outward proportion of the body show +how the laws of life have preserved each animal type. +For the vital organs which constitute each animal +a reptile, and the distinctive bony structures with +which they are associated, remain unaffected, or but +little modified, by the animal's external change in +appearance.</p> + +<p>The creeping reptile is commonly imagined as the +antithesis of the bird. For the bird overcomes the +forces that hold even man to the earth, and enjoys +exalted aerial conditions of life. Therefore the marvel +is shared equally by learned and unlearned, that the +power of flight should have been an endowment of +animals sprung from the breed of serpents, or crocodiles, +enabling them to move through the air as +though they too were of a heaven-born race. The +wonder would not be lessened if the animal were +a degraded representative of a nobler type, or if +it should be demonstrated that even beasts have +advanced in the battle of life. The winged reptile,<span class='pagenum'><a name="Page_3" id="Page_3">[Pg 3]</a></span> +when compared with a bird, is not less astounding +than the poetic conceptions in Milton's <i>Paradise +Lost</i> of degradation which overtakes life that once +was amongst the highest. And on the other hand, +from the point of view of the teaching of Darwin +in the theories of modern science, we are led to ask +whether a flying reptile may not be evidence of the +physical exaltation which raises animals in the scale +of organisation. The dominance upon the earth of +flying reptiles during the great middle period of +geological history will long engage the interest +of those who can realise the complexity of its +structure, or care to unravel the meaning of the +procession of animal forms in successive geological +ages which preceded the coming of man.</p> + +<p>The outer vesture of an animal counts for little in +estimating the value of ties which bind orders of +animals together, which are included in the larger +classes of life. The kindred relationship which makes +the snake of the same class as the tortoise is determined +by the soft vital organs—brain, heart, lungs—which +are the essentials of an animal's existence and +control its way of life. The wonder which science +weaves into the meaning of the word "reptile," "bird," +or "mammal," is partly in exhibiting minor changes +of character in those organs and other soft parts, but +far more in showing that while they endure unchanged, +the hard parts of the skeleton are modified +in many ways. For the bones of the reptile orders +stretch their affinities in one direction towards the +skeletons of salamanders and fishes; and extend +them also at the same time in other directions, +towards birds and mammals. This mystery we may +hope to partly unravel.</p> + + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_4" id="Page_4">[Pg 4]</a></span></p> +<h2><a name="CHAPTER_II" id="CHAPTER_II"></a><small>CHAPTER II</small><br /><br /> + +HOW A REPTILE IS KNOWN</h2> + +<h4>DEFINITION OF REPTILES BY THEIR +VITAL ORGANS</h4> + + +<p>The relations of reptiles to other animals may be +stated so as to make evident the characters and +affinities which bind them together. Early in the +nineteenth century naturalists included with the Reptilia +the tribe of salamanders and frogs which are +named Amphibia. The two groups have been separated +from each other because the young of Amphibia +pass through a tadpole stage of development. They +then breathe by gills, like fishes, taking oxygen from +the air which is suspended in water, before lungs are +acquired which afterwards enable the animals to take +oxygen directly from the air. The amphibian sometimes +sheds the gills, and leaves the water to live on +land. Sometimes gills and lungs are retained through +life in the same individual. This amphibian condition +of lung and gill being present at the same +time is paralleled by a few fishes which still exist, +like the Australian <i>Ceratodus</i>, the lung-fish, an ancient +type of fish which belongs to early days in geological +time.</p> + +<p>This metamorphosis has been held to separate the<span class='pagenum'><a name="Page_5" id="Page_5">[Pg 5]</a></span> +amphibian type from the reptile because no existing +reptile develops gills or undergoes a metamorphosis. +Yet the character may not be more important +as a ground for classification +than the community of gills and lungs +in the fish and amphibian is ground for +putting them together in one natural +group. For although no gills are found +in reptiles, birds, or mammals, the +embryo of each in an early stage of +development appears to possess gill-arches, +and gill-clefts between them, +through which gills might have been +developed, even in the higher vertebrates, +if the conditions of life had +been favourable to such modification +of structure. In their bones Reptiles +and Amphibia have much in common. +Nearly all true reptiles lay eggs, which +are defined like those of birds by comparatively +large size, and are contained +in shells. This condition is not usual +in amphibians or fishes. When hatched +the young reptile is completely formed, +the image of its parent, and has no need +to grow a covering to its skin like some +birds, or shed its tail like some tadpoles. +The reptile is like the bird in freedom +from important changes of form after the egg is +hatched; and the only structure shed by both is the +little horn upon the nose, with which the embryo +breaks the shell and emerges a reptile or a bird, +growing to maturity with small subsequent variations +in the proportions of the body.</p> + + +<div class="figcenter" style="width: 157px;"> +<a name="Fig_1" id="Fig_1"></a> +<span class="caption">FIG. 1 LUNG OF THE FISH CERATODUS</span> +<img src="images/i_022.jpg" width="157" height="640" alt="FIG. 1 LUNG OF THE FISH CERATODUS" title="FIG. 1" /> +</div> +<p class="center">Partly laid open to show its chambered structure<br /> +(After Günther)</p> + + +<p><span class='pagenum'><a name="Page_6" id="Page_6">[Pg 6]</a></span></p> + +<h4>THE REPTILE SKIN</h4> + +<p>Between one class of animals and another the +differences in the condition of the skin are more +or less distinctive. In a few amphibians there are +some bones in the skin on the under side of the +body, though the skin is usually naked, and in frogs +is said to transmit air to the blood, so as to exercise +a respiratory function of a minor kind. This naked +condition, so unlike the armoured skin of the true +Reptilia, appears to have been paralleled by a number +of extinct groups of fossils of the Secondary rocks, +such as Ichthyosaurs and Plesiosaurs, which were +aquatic, and probably also by some Dinosauria, which +were terrestrial.</p> + +<p>Living reptiles are usually defended with some +kind of protection to the skin. Among snakes and +lizards the skin has commonly a covering of overlapping +scales, usually of horny or bony texture. +The tortoise and turtle tribe shut up the animal in a +true box of bone, which is cased with an armour of +horny plates. Crocodiles have a thick skin embedding +a less continuous coat of mail. Thus the +skin of a reptile does not at first suggest anything +which might become an organ of flight; and its +dermal appendages, or scales, may seem further removed +from the feathers which ensure flying powers +to the bird than from the naked skin of a frog.</p> + + +<h4>THE REPTILE BRAIN</h4> + +<p>Although the mode of development of the young +and the covering of the skin are conspicuous among +important characters by which animals are classified, +the brain is an organ of some importance, although<span class='pagenum'><a name="Page_7" id="Page_7">[Pg 7]</a></span> +of greater weight in the higher Vertebrata than in its +lower groups. Reptiles have links in the mode of +arrangement of the parts of their brains with fishes +and amphibians. The regions of that organ are commonly +arranged in pairs of nervous masses, known +as (1) the olfactory lobes, (2) the cerebrum, behind +which is the minute pineal body, followed by (3) the +pair of optic lobes, and hindermost of all (4) the +single mass termed the cerebellum. These parts of +the brain are extended in longitudinal order, one +behind the other in all three groups. The olfactory +lobes of the brain in Fishes may be as large as +the cerebrum; but among Reptiles and Amphibians +they are relatively smaller, and they assume more of +the condition found in mammals like the Hare or +Mole, being altogether subordinate in size. And the +cerebral masses begin to be wider and higher than +the other parts of the brain, though they do not extend +forward above the olfactory lobes, as is often seen in +Mammals. In Crocodiles the cerebral hemispheres +have a tendency to a broad circular form. Among +Chelonian reptiles that region of the brain is more +remarkable for height. Lizards and Ophidians both +have this part of the brain somewhat pear-shaped, +pointed in front, and elongated. The amphibian +brain only differs from the lizard type in degree; and +differences between lizards' and amphibian brains are +less noticeable than between the other orders of +reptiles. The reptilian brain is easily distinguished +from that of all other animals by the position and +proportions of its regions (see <a href="#Fig_19">Fig. 19, p. 53</a>).</p> + +<p>Birds have the parts of the brain formed and +arranged in a way that is equally distinctive. The +cerebral lobes are relatively large and convex, and<span class='pagenum'><a name="Page_8" id="Page_8">[Pg 8]</a></span> +deserve the descriptive name "hemispheres." They +are always smooth, as among the lower Mammals, +and extend backward so as to abut against the hind +brain, termed the cerebellum. This junction is +brought about in a peculiar way. The cerebral +hemispheres in a bird do not extend backward to +override the optic lobes, and hide them, as occurs +among adult mammals, but they extend back between +the optic lobes, so as to force them apart and +push them aside, downward and backward, till they +extend laterally beyond the junction of the cerebrum +with the cerebellum. The brain of a Bird is never +reptilian; but in the young Mammal the brain has +a very reptilian aspect, because both have their parts +primarily arranged in a line. Therefore the brain +appears to determine the boundary between bird +and reptile exactly.</p> + + +<h4>REPTILIAN BREATHING ORGANS</h4> + +<p>The breathing organs of Birds and Reptiles which +are associated with these different types of brain are +not quite the same. The Frog has a cellular lung +which, in the details of the minute sacs which branch +and cluster at the terminations of the tubes, is not +unlike the condition in a Mammal. In a mammal +respiration is aided by the bellows-like action of the +muscles connected with the ribs, which encase the +cavity where the lungs are placed, and this structure +is absent in the Frog and its allies. The Frog, on the +other hand, has to swallow air in much the same way +as man swallows water. The air is similarly grasped +by the muscles, and conveyed by them downward to +the lungs. Therefore a Frog keeps its mouth shut,<span class='pagenum'><a name="Page_9" id="Page_9">[Pg 9]</a></span> +and the animal dies from want of air if its mouth is +open for a few minutes.</p> + +<p>Crocodiles commonly lie in the sun with their +mouths widely open. The lungs in both Crocodiles +and Turtles are moderately dense, traversed by great +bronchial tubes, but do not differ essentially in plan +from those of a Frog, though the great branches of +the bronchial tubes are stronger, and the air chambers +into which the lung is divided are somewhat smaller. +The New Zealand Hatteria has the lungs of this +cellular type, though rather resembling the amphibian +than the Crocodile. The lungs during life in all +these animals attain considerable size, the maximum +dimensions being found in the terrestrial tortoises, +which owe much of their elevated bulk to the dimensions +of the air cells which form the lungs.</p> + +<p>The lungs of Serpents and Lizards are formed on +a different plan. In both those groups of reptiles +the dense cellular tissue is limited to the part of the +lung which is nearest to the throat. This network +of blood vessels and air cells extends about the +principal bronchial tube much as in other animals, +but as it extends backward the blood vessels become +few until the <i>tubular</i> lung appears in its hinder part, +as it extends down the body, almost as simple in +structure as the air bladder of a fish. Among Serpents +only one of these tubular lungs is commonly +present, and the structure has a less efficient appearance +as a breathing organ than the single lung of the +fish <i>Ceratodus</i> (<a href="#Fig_1">Fig. 1</a>). The Chameleons are a group of +lizards which differ in many ways from most of their +nearest kindred, and the lungs, while conforming in +general plan to the lizard type in being dense at the +throat, and a tubular bladder in the body, give off<span class='pagenum'><a name="Page_10" id="Page_10">[Pg 10]</a></span> +on both sides a number of short lateral branches +like the fingers of a glove (<a href="#Fig_18">Fig. 18, p. 51</a>).</p> + +<p>Thus the breathing organs of reptiles present two +or three distinct types which have caused Serpents +and Lizards to be associated in one group by most +naturalists who have studied their anatomy; while +Crocodiles and Chelonians represent a type of lung +which is quite different, and in those groups has +much in common. These characters of the breathing +organs contribute to separate the cold-blooded +armoured reptiles from the warm-blooded birds +clothed with feathers, as well as from the warm-blooded +mammals which suckle their young; for both +these higher groups have denser and more elastic +spongy lung tissue.</p> + +<p>It will be seen hereafter that many birds in the +most active development of their breathing organs +substantially revert to the condition of the Serpent +or Chameleon in a somewhat modified way. Because, +instead of having one great bronchial tube expanded +to form a vast reservoir of air which can be discharged +from the lung in which the reptile has +accumulated it, the bird has the lateral branches +of the bronchial tubes prolonged so as to pierce the +walls of the lung, when its covering membrane expands +to form many air cells, which fill much of the +cavity of the bird's body (see <a href="#Fig_16">Fig. 16</a>). Thus the bird +appears to combine the characters of such a lung as +that of a Crocodile, with a condition which has some +analogy with the lung of a Chameleon. It is this link +of structure of the breathing organs between reptiles +and birds that constitutes one of the chief interests +of flying reptiles, for they prove to have possessed +air cells prolonged from the lungs, which extended +into the bones.</p> + + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_11" id="Page_11">[Pg 11]</a></span></p> +<h2><a name="CHAPTER_III" id="CHAPTER_III"></a><small>CHAPTER III</small><br /><br /> + +A REPTILE IS KNOWN BY ITS +BONES</h2> + + +<p>Such are a few illustrations of ways in which +reptiles resemble other animals, and differ from +them, in the organs by means of which the classification +of animals is made. But such an idea is +incomplete without noticing that the bony framework +of the body associated with such vital organs also +shows in its chief parts that reptiles are easily recognised +by their bones. I will therefore briefly state +how reptiles are defined in some regions of the +skeleton, for in tracing the history of reptile life +the bones are the principal remains of animals +preserved in the rocks; and the soft organs which +have perished can only be inferred to have been +present from the persistence of durable characteristic +parts of the skeleton, which are associated with those +soft organs in animals which exist at the present day, +and are unknown in other animals in which the +skeleton is different.</p> + + +<h4>THE HANG OF THE LOWER JAW</h4> + +<p>The manner in which the lower jaw is connected +with the skull yields one of the most easily recog<span class='pagenum'><a name="Page_12" id="Page_12">[Pg 12]</a></span>nised +differences between the great groups of vertebrate +animals.</p> + +<p><i>In Mammals.</i>—In every mammal—such as the Dog +or Sheep—the lower jaw, which is formed of one bone +on each side, joins directly on to the head of the +animal, and moves upon a bone of the skull which +is named the temporal bone. This character is +sufficient to prove, by the law of association of soft +and hard parts of the body, that such an animal had +warm blood and suckled its young.</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_2" id="Fig_2"></a> +<span class="caption">FIG. 2</span> +<img src="images/i_029.jpg" width="640" height="191" alt="FIG. 2" title="FIG. 2" /> +<p class="center">Comparison to show the articulation with the lower jaw in a mammal and +<i>Pterodactylus Kochi</i>.<br /> +The quadrate bone is lettered Q in this Pterodactyle, and comes between the skull and<br /> +the lower jaw like the quadrate bone in a bird and in lizards.</p> +</div> + + +<p><i>In Birds.</i>—In birds a great difference is found in +this region of the head. The temporal bone, which +it will be more convenient to name the squamosal +bone, from its squamous or scale-like form, is still +a part of the brain case, and assists in covering the +brain itself, exactly as among mammals. But the +lower jaw is now made up of five or six bones. And +between the hindermost and the squamosal there is +an intervening bar of bone, unknown among mammalia, +which moves upon the skull by a joint, just as +the lower jaw moves upon it. This movable bone +unites with parts of the palate and the face, and is +known as the quadrate bone. Its presence proves +that the animal possessing it laid eggs, and if the<span class='pagenum'><a name="Page_13" id="Page_13">[Pg 13]</a></span> +face bones join its outer border just above the lower +jaw, it proves that the animal possessed hot blood.</p> + +<p><i>In Reptiles.</i>—All reptiles are also regarded as +possessing the quadrate bone. But the squamosal +bone with which it always unites is in less close +union with the brain case, and never covers the brain +itself. Serpents show an extreme divergence in this +condition from birds, for the squamosal bone appears +to be a loose external plate of bone which rests +upon the compact brain case and gives attachment +to the quadrate bone which is as free as in a bird. +Among Lizards the quadrate bone is usually almost +as free. In the other division of existing Reptilia, +including Crocodiles, the New Zealand lizard-like +reptile Hatteria, called Tuatera, and Turtles, the +squamosal and quadrate bones are firmly united with +the bones of the brain case, face, and palate, so that +the quadrate bone has no movement; and the same +condition appears in amphibians, such as Toads and +Frogs. With these conditions of the quadrate bone +are associated cold blood, terrestrial life, and young +developed from eggs.</p> + +<p><i>In Fishes.</i>—Bony fishes, and all others in which +separate bones build up the skull, differ from Reptiles +and Birds much as those animals differ from +Mammals. The union of the lower jaw with the +skull becomes complicated by the presence of additional +bones. The quadrate bone still forms a pulley +articulation upon which the lower jaw works, but +between it and the squamosal bone is the characteristic +bone of the fish known as the hyomandibular, +commonly connected with opercular bones and +metapterygoid which intervene, and help to unite +the quadrate with the brain case. In the Cartila<span class='pagenum'><a name="Page_14" id="Page_14">[Pg 14]</a></span>ginous +fishes there is only one bone connecting the +jaws with the skull on each side. This appears to +prove that just as the structure of the arch of bones +suspending the jaw may be complicated by the +mysterious process called segmentation, which separates +a bone into portions, so simplification and +variation may result because the primitive divisions +of the material cease to be made which exists before +bones are formed.</p> + +<p>The principal regions of the skull and skeleton all +vary in the chief groups of animals with backbones; +so that the Reptile may be recognised among fossils, +even in extinct groups of animals and occasionally +restored from a fragment, to the aspect which characterised +it while it lived.</p> + + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_15" id="Page_15">[Pg 15]</a></span></p> +<h2><a name="CHAPTER_IV" id="CHAPTER_IV"></a><small>CHAPTER IV</small><br /><br /> + +ANIMALS WHICH FLY</h2> + + +<p>The nature of a reptile is now sufficiently intelligible +for something to be said concerning +flight, and structures by means of which some animals +lift themselves in the air. It is not without interest +to remember that, from the earliest periods in human +records, representations have been made of animals +which were furnished with wings, yet walked upon +four feet, and in their +typical aspect have the +head shaped like that of +a bird. They are commonly +named Dragons.</p> + + +<h4>FLYING DRAGONS</h4> + +<div class="figcenter" style="width: 461px;"> +<a name="Fig_3" id="Fig_3"></a> +<span class="caption">FIG. 3 From <i>The Battle between Bel and the Dragon</i></span> +<img src="images/i_032.jpg" width="461" height="640" alt="FIG. 3" title="FIG. 3" /> +</div> + +<p>The effigy of the +dragon survives to the +present day in the figure +over which St. George +triumphs, on the reverse +of the British sovereign. +In the luxuriant imaginations +of ancient Eastern +peoples, dating back +<span class='pagenum'><a name="Page_16" id="Page_16">[Pg 16]</a></span> +to prehistoric ages, perhaps 5000 <small>B.C.</small>, the dragons +present an astonishing constancy of form. In after-times +they underwent a curious evolution, as the conception +of Babylon and Egypt is traced through +Assyria to Greece. The Wings, which had been associated +at first with the fore limb of the typical dragon, +become characteristic of the Lion, and of the poet's +winged Horse, and finally of the Human figure itself, +<span class='pagenum'><a name="Page_17" id="Page_17">[Pg 17]</a></span> +carved on the great columns of the Greek temples of +Ephesus. These flying animals are historically descendants +of the same common stock with the dragons +of China and Japan, which still preserve the aspect +of reptiles. Their interest is chiefly in evidence of +a latent spirit of evolution in days too remote for its +meaning to be now understood, which has carried the +winged forms higher and ever higher in grade of +organisation, till their wings ceased to be associated +with feelings of terror. The Hebrew cherubim are +regarded by H. E. Ryle, Bishop of Exeter, as probably +Dragons, and the figure of the conventional +angel is the human form of the Dragon.</p> + +<div class="figcenter" style="width: 480px;"> +<a name="Fig_4" id="Fig_4"></a> +<span class="caption">FIG. 4. FIGURE FROM THE TEMPLE OF EPHESUS</span> +<img src="images/i_033.jpg" width="480" height="636" alt="FIG. 4. FIGURE FROM THE TEMPLE OF EPHESUS" title="" /> +</div> + + +<h4>ORGANS OF FLIGHT</h4> + +<p>Turning from this reference to the realm of mythology +to existing nature, the power of flight is +popularly associated with all the chief types of +vertebrate animals—fishes, frogs, lizards, birds, and +mammals. Many of the animals ill deserve the +name of flyers, and most are exceptions to different +conditions of existence which control their kindred, +but it is convenient to examine for a little the nature +of the structures by which this movement in the air, +which is not always flight, is made possible. Certain +fishes, like the lung-fish Ceratodus, of Queensland, +and the mud-fish Lepidosiren, are capable of leaving +the water and living on land, and for a time breathe +air. But neither these fishes nor Periophthalmus, +which runs with rapid movement of its fins and +carries the body more or less out of water, or the +climbing perch, Anabas, carried out of water over +the country by Indian jugglers, ever put on the +slightest approach to wings.</p> +<p><span class='pagenum'><a name="Page_18" id="Page_18">[Pg 18]</a></span></p> + +<h4>FLYING FISHES</h4> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_5" id="Fig_5"></a> +<span class="caption">FIG. 5. THE FLYING FISH EXOCŒTUS</span> +<p class="center">With the fins extended moving through the air</p> +<img src="images/i_035.jpg" width="640" height="365" alt="FIG. 5." title="FIG. 5." /> +</div> + +<p>The flight of fishes is a kind of parachute support +not unlike that by which a folded paper is made to +travel in the air. It is chiefly seen in the numerous +species of a genus Exocœtus, allied to the gar-pike +(Belone), which is common in tropical seas, and +usually from a foot to eighteen inches long. They +emerge from the water, and for a time support themselves +in the air by means of the greatly developed +breast fins, which sometimes extend backward to the +tail fin. Although these fins appear to correspond +to the fore limbs of other animals, they may not +be moved at the will of the fish like the wing of a +bird. When the flying fishes are seen in shoals in +the vicinity of ships, those fins remain extended, so +that the fish is said sometimes to travel 200 yards +at a speed of fifteen miles an hour, rising twenty feet +or more above the surface of the sea, travelling in +a straight line, though sometimes influenced by the +wind. Here the organ, which is at once a fin and a<span class='pagenum'><a name="Page_19" id="Page_19">[Pg 19]</a></span> +wing, consists of a number of thin long rods, or rays, +which are connected by membrane, and vary in +length to form an outline not unlike the wing of a +bird which tapers to a point. The interest of these +animals is chiefly in the fact that flight is separated +from the condition of having lungs with which it +is associated in birds, for although the flying fish has +an air bladder, there is no duct to connect it with +the throat.</p> + + +<h4>FLYING FROGS</h4> + +<div class="figcenter" style="width: 480px;"> +<a name="Fig_6" id="Fig_6"></a> +<span class="caption">FIG. 6. THE FLYING FROG (RHACOPHORUS)</span> +<img src="images/i_036.jpg" width="480" height="548" alt="FIG. 6." title="FIG. 6." /> +<p class="center">The membranes of the foot and +hand extend between the metatarsal +and metacarpal bones, as well as the +bones of the digits.</p> +</div> + + +<p>Among amphibians the organs of flight are also +of a parachute kind, but of a different nature. They +are seen in certain frogs which +frequent trees, and are limited +to membranes which extend +between the diverging digits +of the hand and foot, forming +webs as fully developed as in +the foot of a swimming bird. +As these frogs leap, the membranes +are expanded and help +to support the weight of the +body, so that the animal descends +more easily as it moves +from branch to branch. There +is no evidence that the bones +of the digits ever became elongated like the fin rays +of the flying fish or the wing bones of a Bat; but +the web suggests the basis of such a wing, and the +possibilities under which wings may first originate, +by elongation of the bones of a webbed hand like +that of a Flying Frog.</p> +<p><span class='pagenum'><a name="Page_20" id="Page_20">[Pg 20]</a></span></p> + +<h4>FLYING LIZARDS</h4> + + +<p>The Reptilia in their several orders are remarkable +for absence of any modification of the arms which +might suggest a capacity for acquiring wings, as +being latent in their organisation. Crocodiles, Tortoises, +and Serpents are alike of the earth, and not +of the air. But among Lizards there are small groups +of animals in which a limited capacity for +movement through the air is developed. +It is best known in the family of small +lizards named Dragons, represented typically +by the species <i>Draco volans</i> found in the Oriental +region of the East Indies and Malay Archipelago.</p> + +<div class="figcenter" style="width: 620px;"> +<a name="Fig_7" id="Fig_7"></a> +<span class="caption">FIG. 7. THE FLYING DRAGON, DRACO</span> +<p class="center">Forming a parachute by means of the extended ribs</p> +<img src="images/i_037.jpg" width="620" height="480" alt="FIG. 7." title="FIG. 7." /> +</div> + + +<p>The organ of flight is produced in an unexpected +way, by means of the ribs instead of the limbs. The +ribs extend outward as far as the arms can stretch, +and the first five or six are prolonged beyond the +body so as to spread a fold of skin on each side +between the arm and the leg. The membrane admits<span class='pagenum'><a name="Page_21" id="Page_21">[Pg 21]</a></span> +of some movement with the ribs. This arrangement +forms a parachute, which enables the animal to move +rapidly among branches of trees, extending the structure +at will, so that it is used with rapidity too quick +to be followed by the eye, as it leaps through considerable +distances.</p> + +<p>A less singular aid to movement in the air is found +in some of the lizards termed Geckos. The so-called +Flying Gecko (<i>Platydactylus homalocephalus</i>) has a +fringe unconnected with ribs, which extends laterally +on the sides of the body and tail, as well as at the +back and front of the fore and hind limbs, and between +the digits, where the web is sometimes almost as +well developed as among Tree Frogs. This is essentially +a lateral horizontal frill, extending round the +body. Its chief interest is in the circumstance that it +includes a membrane which extends between the wrist +bones and the shoulder on the front of the arm. That +is the only part of the fringe which represents the wing +membrane of a bird. The fossil flying reptiles have +not only that membrane, but the lateral membranes +at the sides of the body and behind the arms.</p> + +<p>Other lizards have the skin developed in the +direction of the circumference of the body. In the +Australian Chlamydosaurus it forms an immense +frill round the neck like a mediæval collar. But +though such an adornment might break a fall, it +could not be regarded as an organ of flight.</p> + + +<h4>FLYING BIRDS</h4> + + +<p>The wings of birds, when they are developed so as +to minister to flight, are all made upon one plan; but +as examples of the variation which the organs contributing +to make the fore limb manifest, I may +<span class='pagenum'><a name="Page_22" id="Page_22">[Pg 22]</a></span> +instance the short swimming limb of the Penguin, +the practically useless rudiment of a wing found in +the Ostrich or Kiwi, and the fully developed wing of +the Pigeon. The wings of birds obtain an extensive +surface to support the animal by muscular movements +of three modifications of structure. First, the bones +of the fore limb are so shaped that they cannot, in +existing birds, be applied to the ground for support +and be used like the limbs of quadrupeds, and are +therefore folded up at the sides of the body, and<span class='pagenum'><a name="Page_23" id="Page_23">[Pg 23]</a></span> +carried in an unused or useless state so long as the +animal hops on the ground or walks, balancing its +weight on the hind legs. Secondly, there are two +small folds of skin, less conspicuous than those on +the arms of Geckos; one is between the wrist bones +and the shoulder, and the smaller hinder membrane +is between the upper arm and the body. These +membranous expansions are insignificant, and would +in themselves be inadequate to support the body or +materially assist its movements. Thirdly, the bird +develops appendages to the skin which are familiarly +known as feathers, and the large feathers which make +the wing are attached to the skin covering the lower +arm bone named the ulna, and the other bones which +represent the wrist and hand. The area and form of +the bird's wing are due to individual appendages to +the skin, which are unknown in any other group of +animals. Between the extended wing of the Albatross, +measuring eleven feet in spread, and the condition +in the Kiwi of New Zealand, in which the +wing is vanishing, there is every possible variation in +size and form. As a rule, the larger the animal the +smaller is the wing area. The problem of the origin +of the bird's wing is not to be explained by study of +existing animals; for the rowing organ of the Penguin, +which in itself would never suggest flight, +becomes an organ of flight in other birds by the +growth upon it of suitable feathers. Anyone who +has seen the birds named Divers feeding under water, +swimming rapidly with their wings, might never +suspect that they were also organs of aerial flight. +The Ostrich is even more interesting, for it has not +developed flight, and still retains at the extremities +of two of the digits the slender claws of a limb +which was originally no wing at all, but the support +of a four-footed animal (<a href="#Fig_46">Fig. 46, p. 130</a>).</p> + +<div class="figcenter" style="width: 480px;"> +<a name="Fig_8" id="Fig_8"></a> +<span class="caption">FIG. 8. POSITION OF BIRDS IN FLIGHT</span> +<img src="images/i_039.jpg" width="480" height="481" alt="FIG. 8. POSITION OF BIRDS IN FLIGHT" title="FIG. 8." /> +</div> + +<p><span class='pagenum'><a name="Page_24" id="Page_24">[Pg 24]</a></span></p> +<h4>FLYING MAMMALS</h4> + +<p>Flight is also developed among mammals. The +Insectivora include several interesting examples of +animals which are capable of a certain motion through +the air. In the tropical forests of the Malay Archipelago +are animals known as Flying Squirrels, Flying +Opossums, Flying Lemurs, Flying +Foxes, in which the skin extends +outward laterally from the sides +of the body so as to connect the +fore limbs with the hind limbs, +and is also prolonged backward +from the hind limbs to the tail. +The four digits are never elongated; +the bones of the fore limb +are neither longer nor larger than +those of the hind limb, and the +foot terminates in five little claws +as in other four-footed animals. +This condition is adapted for the arboreal life which +those animals live, leaping from branch to branch, +feeding on fruits and leaves, and in some cases +upon insects. These mammals may be compared +with the Flying Geckos among reptiles in their +parachute-like support by extension of the skin, +which gives them one of the conditions of support +which contribute to constitute flight.</p> + +<div class="figcenter" style="width: 423px;"> +<a name="Fig_9" id="Fig_9"></a> +<span class="caption">FIG. 9. FLYING SQUIRREL (PTEROMYS)</span> +<img src="images/i_041.jpg" width="423" height="640" alt="FIG. 9." title="FIG. 9." /> +</div> + +<p><i>Bats.</i>—One entire order of mammals—the Bats—not +only possess true wings, but are capable of flight +which is sustained, and in some cases powerful. The +wings are clothed with short hair like the rest of the +<span class='pagenum'><a name="Page_25" id="Page_25">[Pg 25]</a></span> +body, and thus the instrument of flight is unlike that +of a bird. The flight of a Bat differs from that +of all other animals in being dependent upon a +modification of the bones of the fore limb, which, +without interfering with the animal's movements as +a quadruped, secures an extension of the wing which +is not inferior in area to that which the bird obtains +by elongation of the bones of the arm and fore-arm +and its feathers. The distinctive peculiarity of the +Bat's wing is in the circumstance that four of the +digits of the hand have their bones prolonged to +a length which is often equal to the combined length +of the arm and fore-arm. The bones of the digits<span class='pagenum'><a name="Page_26" id="Page_26">[Pg 26]</a></span> +diverge like the ribs of an umbrella, and between +them is the wing membrane, which extends from the +sides of the body outward, unites the fore limb with +the hind limb, and is prolonged down the tail as +in the Flying Foxes. Bats have a small membrane +in front of the bones of the arm and fore-arm +stretching between the shoulder and the wrist, which +corresponds with the wing membrane of a bird; but +the remainder of the membranes in Bats' wings are +absent in birds, because their function is performed +by feathers which give the wing its area. The +elongated digits of the Bat's wing are folded together +and carried at the sides of the body as though they +were a few quill pens attached to its wrist, where the +one digit, which is applied to the ground in walking, +terminates in a claw.</p> + +<div class="figcenter" style="width: 551px;"> +<a name="Fig_10" id="Fig_10"></a> +<span class="caption">FIG. 10 NEW ZEALAND BAT FLYING. BARBASTELLE WALKING</span> +<img src="images/i_042.jpg" width="551" height="480" alt="FIG. 10" title="FIG. 10" /> +</div> + +<p>The organs which support animals in the air are +thus seen to be more or less dissimilar in each of the +great groups of animals. They fall into three chief +types: first, the parachute; secondly, the wing due +to the feathers appended to the skin; and thirdly, the +wing formed of membrane, supported by enormous +elongation of the small bones of the back of the +hand and fingers. The two types of true wings are +limited to birds and bats; and no living reptile +approximates to developing such an organ of flight +as a wing. Judged, therefore, by the method of comparing +the anatomical structures of one animal with +another, which is termed "comparative anatomy," the +existence of flying reptiles might be pronounced +impossible. But in the light which the revelations +of geology afford, our convictions become tempered +with modesty; and we learn that with Nature nothing +is impossible in development of animal structure.</p> + + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_27" id="Page_27">[Pg 27]</a></span></p> +<h2><a name="CHAPTER_V" id="CHAPTER_V"></a><small>CHAPTER V</small><br /><br /> + +DISCOVERY OF THE +PTERODACTYLE</h2> + + +<p>Late in the eighteenth century, in 1784, a small +fossil animal with wings began to be known +through the writings of Collini, as found in the white +lithographic limestone of Solenhofen in Bavaria, and +was regarded by him as a former inhabitant of the +sea. The foremost naturalist of the time, the citizen +Cuvier—for it was in the days of the French Republic—in +1801, in lucid language, interpreted the animal as +a genus of Saurians. That word, so familiar at the +present day, was used in the first half of the century +to include Lizards and Crocodiles; and described +animals akin to reptiles which were manifestly related +neither to Serpents nor Turtles. But the term +saurian is no longer in favour, and has faded from +science, and is interesting only in ancient history of +progress. The lizards soon became classed in close +alliance with snakes. And the crocodiles, with the +Hatteria, were united with chelonians. Most modern +naturalists who use the term saurian still make it +an equivalent of lizard, or an animal of the lizard +kind.</p> +<p><span class='pagenum'><a name="Page_28" id="Page_28">[Pg 28]</a></span></p> + +<h4>CUVIER</h4> + +<div class="figcenter" style="width: 480px;"> +<a name="Fig_11" id="Fig_11"></a> +<span class="caption">FIG. 11. <i>PTERODACTYLUS LONGIROSTRIS</i> (Cuvier)</span> +<img src="images/i_045.jpg" width="480" height="486" alt="FIG. 11." title="FIG. 11." /> +<p class="center">The remains are preserved with the neck arched over the back, and the jaws +opened upward</p> +</div> + + +<p>Cuvier defined this fossil from Solenhofen as distinguished +by the extreme elongation of the fourth +digit of the hand, and from that character invented +for the animal the name Pterodactyle. He tells us +that its flight was not due to prolongation of the ribs, +as among the living lizards named Dragons; or to a +wing formed without the digits being distinguishable +from each other, as among Birds; nor with only one +digit free from the wing, as among Bats; but by<span class='pagenum'><a name="Page_29" id="Page_29">[Pg 29]</a></span> +having the wing supported mainly by a single greatly +elongated digit, while all the others are short and +terminate in claws. Cuvier described the amazing +animal in detail, part by part; and such has been the +influence of his clear words and fame as a great +anatomist that nearly every writer in after-years, +in French and in English, repeated Cuvier's conclusion, +maintained to the end, that the animal is a +saurian.</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_12" id="Fig_12"></a> +<span class="caption">FIG. 12. THE SKELETON OF <i>PTERODACTYLUS LONGIROSTRIS</i></span> +<img src="images/i_046.jpg" width="640" height="404" alt="FIG. 12." title="FIG. 12." /> +<p class="center">Reconstructed from the scattered bones in Fig. 14, showing the limbs +on the left side</p> +</div> + +<p>Long before fashion determined, as an article of +educated belief, that fossil animals exist chiefly to +bridge over the gaps between those which still survive, +the scientific men of Germany were inclined to +see in the Pterodactyle such an intermediate type +of life. At first Sömmerring and Wagler would +have placed the Pterodactyle between mammals +and birds.</p> +<p><span class='pagenum'><a name="Page_30" id="Page_30">[Pg 30]</a></span></p> + +<h4>GOLDFUSS</h4> + +<div class="figcenter" style="width: 541px;"> +<a name="Fig_13" id="Fig_13"></a> +<span class="caption">FIG. 13. THE <i>PTERODACTYLUS LONGIROSTRIS</i> RESTORED +FROM THE REMAINS IN FIG. 11</span> +<img src="images/i_047.jpg" width="541" height="480" alt="FIG. 13." title="FIG. 13." /> +<p class="center">Showing positions of the wing membranes with the animal at rest</p> +</div> + +<p>But the accomplished naturalist Goldfuss, who +described another fine skeleton of a Pterodactyle +in 1831, saw in this flying animal an indication of +the course taken by Nature in changing the reptilian +organisation to that of birds and mammals. It is +the first flash of light on a dark problem, and its +brilliance of inference has never been equalled. Its +effects were seen when Prince Charles Bonaparte, +the eminent ornithologist, in Italy, suggested for the +group the name Ornithosauria; when the profound +anatomist de Blainville, in France, placed the short-tailed +animal in a class between Reptiles and Birds +named Pterodactylia; and Andreas Wagner, of +Munich, who had more Pterodactyles to judge from<span class='pagenum'><a name="Page_31" id="Page_31">[Pg 31]</a></span> +than his predecessors, saw in the fossil animal a +saurian in transition to a bird.</p> + + +<h4>VON MEYER</h4> + +<p>But the German interpretation is not uniform, +and Hermann von Meyer, the banker-naturalist of +Frankfurt a./M., who made himself conversant with +all that his predecessors knew, and enlarged knowledge +of the Pterodactyles on the most critical facts +of structure, continued to regard them as true reptiles, +but flying reptiles. Such is the influence of von +Meyer that all parts of the world have shown a +disposition to reflect his opinions, especially as they +practically coincide with the earlier teaching of +Cuvier. Owen and Huxley in England, Cope and +Marsh in America, Gaudry in France, and Zittel in +Germany have all placed the Pterodactyles as flying +reptiles. Their judgment is emphatic. But there is +weight of competent opinion to endorse the evolutionary +teaching of Goldfuss that they rise above +reptiles. To form an independent opinion the modern +student must examine the animals, weigh their characters +bone by bone, familiarise himself, if possible, +with some of the rocks in which they are found; +to comprehend the conditions under which the fossils +are preserved, which have added not a little to the +interest in Pterodactyles, and to the difficulty of +interpretation.</p> + + +<h4>GEOLOGICAL HISTORY OF PTERODACTYLES IN +GERMANY</h4> + +<p>We may briefly recapitulate the geological history. +Those remains of Ornithosaurs which have been mentioned, +with a multitude of others which are the glory<span class='pagenum'><a name="Page_32" id="Page_32">[Pg 32]</a></span> +of the museums of Munich, Stuttgart, Tübingen, +Heidelberg, Bonn, Haarlem, and London, have all +been found in working the lithographic stone of +Bavaria. The whitish yellow limestone forms low, +flat-topped hills, now isolated from each other by +natural denudation, which has removed the intervening +rock. The stone is found at some distance +north of the Danube, in a line due north of Augsburg, +in the country about Pappenheim, and especially at +the villages of Solenhofen, Eichstädt, Kelheim, and +Nusplingen. These beds belong to the rocks which +are named White Jura limestone in Germany, which +is of about the same geological age as the Kimeridge +clay in England. Much of it divides into very thin +layers, and in these planes of separation the fossils +are found. They include the <i>Ammonites lithographicus</i> +and a multitude of marine shells, king +crabs and other Crustacea, sea-urchins, and other +fossils, showing that the deposit was formed in the +sea. The preservation of jelly-fish, which so soon +disappear when left dry on the beach, shows that the +ancient calcareous mud had unusual power of preserving +fossils. Into this sea, with its fishes great +and small, came land plants from off the land, dragonflies +and other insects, tortoises and lizards, Pterodactyles +with their flying organs, and birds still +clothed with feathers. Sometimes the wing membranes +of the flying reptiles are found fully stretched +by the wing finger, as in examples to be seen at +Munich and in the Yale Museum in Newhaven, in +America. At Haarlem there is an example in which +the wing membrane appears to be folded much as in +the wing of a Bat, when the animal hangs suspended, +with the flying membrane bent into a few wide undulations.<span class='pagenum'><a name="Page_33" id="Page_33">[Pg 33]</a></span></p> + +<p>The Solenhofen Slate belongs to about the middle +period of the history of flying reptiles, for they +range through the Secondary epochs of geological +time. Remains are recorded in Germany from the +Keuper beds at the top of the Trias, which is the +bottom division of the Secondary strata; and I believe +I have seen fragments of their bones from +the somewhat older Muschelkalk of Germany.</p> + + +<h4>THEIR HISTORY IN ENGLAND</h4> + +<p>In England the remains are found for the first time +in the Lower Lias of Lyme Regis, in Dorset, and the +Upper Lias of Whitby, in Yorkshire. In Würtemberg +they occur on the same horizons. They reappear in +England, in every subsequent age, when the conditions +of the strata and their fossils give evidence of +near proximity to land. In the Stonesfield Slate of +Stonesfield, in Oxfordshire, the bones are found +isolated, but indicate animals of some size, though +not so large as the rare bones of reputed true +birds which appear to have left their remains in +the same deposit.</p> + +<p>At least two Pterodactyles are found in the Oxford +clay, known from more or less fragmentary remains or +isolated bones; just as they occur in the Kimeridge +Clay, Purbeck Limestone, Wealden sandstones, and +especially in newer Secondary rocks, named Gault, +Upper Greensand, and Chalk, in the south-east of +England.</p> + +<p>Owing to exceptional facilities for collecting, in +consequence of the Cambridge Greensand being +excavated for the valuable mineral phosphate of +lime it contains, more than a thousand bones are +preserved, more or less broken and battered, in the<span class='pagenum'><a name="Page_34" id="Page_34">[Pg 34]</a></span> +Woodwardian Museum of the University of Cambridge +alone. To give some idea of their abundance, +it may be stated that they were mostly gathered +during two or three years, as a matter of business, +by an intelligent foreman of washers of the nodules +of phosphate of lime, which, in commerce, are named +coprolites. He soon learned to distinguish Pterodactyle +bones from other fossils by their texture, and +learned the anatomical names of bones from specimens +in the University Museum. This workman, +Mr. Pond, employed by Mr. William Farren, brought +together not only the best of the remains at Cambridge, +but most of those in the museums at York +and in London, and the thousands of less perfect +specimens in public and private collections which +passed through the present writer's hands in endeavours +to secure for the University useful illustrations +of the animal's structure. These fragments, +among which there are few entire bones, are valuable, +for they have afforded opportunities of examining +the articular ends of bones in every aspect, which +is not possible when similar organic remains are embedded +in rock in their natural connexions.</p> + +<p>In England Flying Reptiles disappear with the +Chalk. In that period they were widely distributed, +being found in Bohemia, in Brazil, and Kansas in the +United States, as well as in Kent and other parts of +England. They attained their largest dimensions in +this period of geological time. One imperfect fragment +of a bone from the Laramie rocks of Canada +was described, I believe, by Cope, though not identified +by him as Ornithosaurian, and is probably newer +than other remains.</p> +<p><span class='pagenum'><a name="Page_35" id="Page_35">[Pg 35]</a></span></p> + +<h4>ASPECT OF PTERODACTYLES</h4> + +<p>If this series of animals could all be brought +together they would vary greatly in aspect and +stature, as well as in structure. Some have the head +enormously long, in others it is large and deep, +characters which are shared by extinct reptiles which +do not fly, and to which some birds may approximate; +while in a few the head is small and compact, +no more conspicuous, relatively, than the head of +a Sparrow. The neck may be slender like that of +a Heron, or strong like that of an Eagle; the back is +always short, and the tail may be inconspicuous, or +as long as the back and neck together. These flying +reptiles frequently have the proportions of the limbs +similar to those of a Bat, with fore legs strong and +hind legs relatively small; while in some the limbs +are as long, proportionately, and graceful as those of +a Deer. With these differences in proportions of the +body are associated great differences in the relative +length of the wing and spread of the wing membranes.</p> + + +<h4>DIMENSIONS OF THE ANIMALS</h4> + +<p>The dimensions of the animals have probably +varied in all periods of geological time. The +smallest, in the Lithographic Slate, are smaller than +Sparrows, while associated with them are others in +which the drumstick bone of the leg is eight inches +long. In the Cambridge Greensand and Chalk imperfect +specimens occur, showing that the upper arm +bones are larger than those of an Ox. The shaft is +one and a half inches in diameter and the ends three +inches wide. Such remains may indicate Pterodactyles +not inferior in size to the extinct Moas of<span class='pagenum'><a name="Page_36" id="Page_36">[Pg 36]</a></span> +New Zealand, but with immensely larger heads, +animals far larger than birds of flight.</p> + +<p>The late Sir Richard Owen, on first seeing these +fragmentary remains, said "the flying reptile with +outstretched pinions must have appeared like the +soaring Roc of Arabian romance, but with the features +of leathern wings with crooked claws superinduced, +and gaping mouth with threatening teeth." +Eventually we shall obtain more exact ideas of their +aspect, when the structures of the several regions of +the body have been examined. The great dimensions +of the stretch of wing, often computed at +twenty feet in the larger examples, might lead to +expectations of great weight of body, if it were not +known that an albatross, with wings spreading +eleven feet, only weighs about seventeen pounds.</p> + + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_37" id="Page_37">[Pg 37]</a></span></p> +<h2><a name="CHAPTER_VI" id="CHAPTER_VI"></a><small>CHAPTER VI</small><br /><br /> + +HOW ANIMALS ARE INTERPRETED +BY THEIR BONES</h2> + + +<p>There is only one safe path which the naturalist +may follow who would tell the story of the +meaning and nature of an extinct type of animal +life, and that is to compare it as fully as possible in +its several bones, and as a whole, with other animals, +especially with those which survive. It is easy to +fix the place in nature of living animals and determine +their mutual relations to each other, because all +the organs—vital as well as locomotive—are available +for comparison. On such evidence they are +grouped together into the large divisions of Beasts, +Birds, and Reptiles; as well as placed in smaller +divisions termed Orders, which are based upon less +important modifications of fundamental structures. +All these characteristic organs have usually disappeared +in the fossil. Hence a new method of +study of the hard parts of the skeleton, which alone +are preserved, is used in the endeavour to discover +how the Flying Reptile or other extinct animal is to +be classified, and how it acquired its characters or +came into existence.</p> +<p><span class='pagenum'><a name="Page_38" id="Page_38">[Pg 38]</a></span></p> + +<h4>VARIATIONS OF BONES AMONG MAMMALIA</h4> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_14" id="Fig_14"></a> +<span class="caption">FIG. 14. THE FORE LIMB IN FOUR TYPES OF MAMMALS</span> +<img src="images/i_055.jpg" width="640" height="454" alt="FIG. 14." title="FIG. 14." /> +<p class="center">Comparison of the fore limb in mammals, showing variation +of form of the bones with function</p> +</div> + + +<p>Resemblances and differences in the bones are +easily over-estimated in importance as evidence of +pedigree relationship. The Mammalia show, by +means of such skeletons as are exhibited in any +Natural History Museum, how small is the importance +to be attached to even the existence of any +group of bones in determining its grade of organisation. +The whole Whale tribe suckle their young and +conform to the distinctive characters in brain and +lungs which mark them as being mammals. But if +there is one part of the skeleton more than another +which distinguishes the Mammalia, it is the girdle of +bones at the hips which supports the hind limbs. It +is characterised by the bone named the ilium being<span class='pagenum'><a name="Page_39" id="Page_39">[Pg 39]</a></span> +uniformly directed forward. Yet in the Whale tribe +the hip-girdle and the hind limb which it usually +supports are so faintly indicated as to be practically +lost; while the fore limb becomes a paddle without +distinction of digits, and is therefore devoid of hoofs +or claws, which are usual terminations of the extremities +in mammals. Yet this swimming paddle, with +its ill-defined bones—sometimes astonishing in number, +as well as in fewness of the finger bones—is +represented by the burrowing fore limb of the Mole, +which lives underground; by the elongated hoofed +legs of the Giraffe, which lives on plains; and the +extended arm and finger bones of the Bat, which are +equally mammals with the Whale. From such comparison +it is seen that no proportion, or form, or +length, or use of the bones of the limbs, or even the +presence of limbs, is necessarily characteristic of a +mammal. No limitation can be placed upon the +possible diversity of form or development of bones +in unknown animals, when they are considered in the +light of such experience of varied structural conditions +in living members of a single class.</p> + +<p>What is true for the limbs and the bony arches +which support them is true for the backbone also, for +the ribs, and to some extent for the skull. The neck +in the Whale is shortened almost beyond recognition. +In the Giraffe the same seven vertebræ are elongated +into a marvellous neck; so that in the technical +definition of a mammal both are said to have seven +neck vertebræ. Yet exceptions show a capacity for +variation. One of the Sloths reduces the number to +six, while another has nine vertebræ in the neck; +proving that there is no necessary difference between +a mammal and a reptile when judged by a character<span class='pagenum'><a name="Page_40" id="Page_40">[Pg 40]</a></span> +which is typically so distinctive of mammals as the +number of the neck bones.</p> + +<p>The skull varies too, though to a less extent. The +Great Ant-eater of South America is a mammal absolutely +without teeth. The Porpoises have a simple +unvarying row of conical teeth with single roots extending +along the jaw. And the dental armature of +the jaws, and relative dimensions of the skull bones, +exhibit such diversity, in evidence of what may be +parted with or acquired, that recognition of the many +reptilian structures and bones in the skull of Ornithorhynchus, +the Australian Duckbill, demonstrates +that the difficulties in recognising an animal by its +bones are real, unless we can discover the Animal +Type to which the bones belong; and that there is +very little in osteology which may not be lost without +affecting an animal's grade of organisation.</p> + + +<h4>VARIATION IN SKIN COVERING OF MAMMALS</h4> + +<p>Even the covering of the body varies in the same +class, or even order of animals, so that the familiar +growth on the skin is never its only possible covering. +The Indian ant-eater, named Manis, which +looks like a gigantic fir-cone, the Armadillo, which +sheathes the body in rings of bone, bearing only a +scanty development of hair, are examples of mammalian +hair, as singular as the quills of a Porcupine, +the horn of a Rhinoceros, or the growth of hair of +varying length and stoutness on different parts of the +body in various animals, or the imperfect development +of hair in the marine Cetacea. Among living +animals it is enough for practical purposes to say +that a mammal is clothed with hair, but in a fossil<span class='pagenum'><a name="Page_41" id="Page_41">[Pg 41]</a></span> +state the hair must usually be lost beyond recognition +from its fineness and shortness of growth.</p> + + +<h4>VARIATION IN SKIN COVERING OF BIRDS</h4> + +<p>No Class of living animals is more homogeneous +than Birds; and well-preserved remains prove that, +at least as far back in time as the Upper Oolites, birds +were clothed with feathers of essentially the same +mode of growth and appearance as the feathers of +living birds. There may, therefore, be no ground for +assuming that the covering was ever different, though +some regions of the skin are free from feathers. Yet +the variations from fine under-down to the scale-like +feathers on the wings of a Penguin, or the great +feathers in the wings of birds of flight, or the double +quill of the Ostrich group, are calculated to yield +dissimilar impressions in a fossil state, even if the +fine down would be preserved in any stratum.</p> + + +<h4>VARIATION IN THE BONES OF BIRDS</h4> + +<p>Osteologically there is less variety in the skeleton +of birds than in other great groups of animals. The +existing representatives do not exhaust its capability +for modification. The few specimens of birds hitherto +found in the Secondary strata have rudely removed +many differences in the bones which separated living +birds from reptiles; so that if only the older fossil +birds were known, and the Tertiary and living birds +had not existed, a bird might have been defined as +an animal having its jaw armed with teeth, instead of +devoid of teeth; with vertebræ cupped at both ends, +instead of with a saddle-shaped articulation which in +front is concave from side from side, and convex from +above downwards; in which the bones of the hand<span class='pagenum'><a name="Page_42" id="Page_42">[Pg 42]</a></span> +are separate, so that three digits terminating in claws +can be applied to the ground, instead of the metacarpal +bones being united in a solid mass with clawless +digits; and in which the tail is elongated like +the tail of a lizard. Yet the limits to variation are +not to be formulated till Nature has exhausted all +her resources in efforts to preserve organic types by +adapting them to changed circumstances. Birds may +be regarded theoretically as equally capable with +mammals of parting with almost every distinctive +structure in the skeleton by which it is best known. +Even the living frigate bird blends the early joints of +the backbone into a compact mass like a sacrum. +The Penguin has a cup-and-ball articulation in the +early dorsal vertebræ, with the ball in front. And the +genus Cypselus has the upper arm bone almost as +broad as long, unlike the bird type. Such examples +prove that we are apt to accept the predominant +structures in an animal type as though they were +universal, and forget that inferences based, like those +of early investigators, on limited materials may be +re-examined with advantage.</p> + + +<h4>VARIATION IN THE BONES OF REPTILES</h4> + +<p>The true Reptilia, notwithstanding some strong resemblances +to Birds in technical characters of the +skeleton, display among their surviving representatives +an astonishing diversity in the bony framework +of the body, exceeding that of the mammalia. This +unlooked-for capacity for varying the plan of construction +of the skeleton is in harmony with the +diversity of structure in groups of extinct animals +to which the name reptiles has also been given. The +interval in form is so vast between Serpent and<span class='pagenum'><a name="Page_43" id="Page_43">[Pg 43]</a></span> +Tortoise, and so considerable in structure of the +skeleton between these and the several groups of +Lizards, Crocodiles, and Hatteria, that any other +diversity could not be more surprising. And the +inference is reasonable that just as mammals live +in the air, in the sea, on the earth, and burrow under +the earth, similar modes of existence might be +expected for birds and reptiles, though no bird is +yet known to have put on the aspect of a fish, and +no reptiles have been discovered which roamed in +herds like antelopes, or lived in the air like birds +or bats, unless these fossil flying animals prove on +examination to justify the name by which they are +known.</p> + +<p>Comparative study of structure in this way demolishes +the prejudice, born of experience of the +life which now remains on earth, that the ideas +of Reptile and of Flight are incongruous, and not +to be combined in one animal. The comparative +study of the parts of animals does not leave the +student in a chaos of possibilities, but teaches us +that organic structures, which mark the grades of +life, have only a limited scope of change; while +Nature flings away every part of the skeleton which +is not vital, or changes its form with altering circumstances +of existence, enforced by revolutions of the +Earth's surface in geological time, in her efforts to +save organisms from extinction and pass the grade +of life onward to a later age.</p> + +<p>The bones are only of value to the naturalist as +symbols, inherited or acquired, and vary in value as +evidence of the nature and association of those vital +organs which differentiate the great groups of the +vertebrata.<span class='pagenum'><a name="Page_44" id="Page_44">[Pg 44]</a></span></p> + +<p>These distinctive structures, which separate Mammals, +Birds, and Reptiles, are sometimes demonstrated +by the impress of their existence left on the bones; +or sometimes they may be inferred from the characters +of the skeleton as a whole.</p> + + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_45" id="Page_45">[Pg 45]</a></span></p> +<h2><a name="CHAPTER_VII" id="CHAPTER_VII"></a><small>CHAPTER VII</small><br /><br /> + +INTERPRETATION OF PTERODACTYLES +BY THEIR SOFT PARTS</h2> + +<h4>THE ORGANS WHICH FIX AN ANIMAL'S PLACE +IN NATURE</h4> + + +<p>We shall endeavour to ascertain what marks +of its grade of organisation the Pterodactyle +has to show. The organs which are capable of modifying +the bones are probably limited to the kidneys, +the brain, and the organs of respiration. It may be +sufficient to examine the latter two.</p> + + +<h4>PNEUMATIC FORAMINA IN PTERODACTYLES</h4> + + + +<p>Hermann von Meyer, the historian of the Ornithosaurs +of the Lithographic Slate, as early as 1837 +described some Pterodactyle bones from the Lias +of Franconia, which showed that air was admitted +into the interior of the bones by apertures near their +extremities, which, from this circumstance, are known +as pneumatic foramina. He drew the inference, +naturally enough, that such a structure is absolute +proof that the Pterodactyle was a flying animal. +It was not quite the right form in which the conclusion +should have been stated, because the Ostrich +and other birds which do not fly have the principal +<span class='pagenum'><a name="Page_46" id="Page_46">[Pg 46]</a></span> +bones pneumatic. Afterwards, in 1859, the larger +bones which Professor Sedgwick, of Cambridge, +transmitted to Sir Richard Owen +established this condition as characteristic +of the Flying Reptiles of +the Cambridge Greensand. It was +thus found as a distinctive structure +of the bones both at the beginning +and the close of the geological +history of these animals. Von +Meyer remarks that the supposition +readily follows that in the respiratory +process there was some similarity +between Pterodactyles and Birds. +This cautious statement may perhaps +be due to the circumstance +that in many animals air cavities +are developed in the skull without +being connected with organs of respiration. It +is well known that the bulk of the Elephant's head +is due to the brain cavity being protected with an +envelope formed of large air cells. Small air cells +are seen in the skulls of oxen, pigs, and many other +mammals, as well as in the human forehead. The +head of a bird like the Owl owes something of its +imposing appearance to the way in which its mass +is enlarged by the dense covering of air cells in the +bones above the brain, like that seen in some Cretaceous +Pterodactyles. Nor are the skulls of Crocodiles +or Tortoises exceptions to the general rule that an +animal's head bones may be pneumatic without +implying a pneumatic prolongation of air from the +lungs. The mere presence of air cells without specification +of the region of the skeleton in which they<span class='pagenum'><a name="Page_47" id="Page_47">[Pg 47]</a></span> +occur is not remarkable. The holes by which air +enters the bones are usually much larger in Pterodactyles +than in Birds, but the entrance to the air cell +prolonged into the bones is the same in form and +position in both groups. So far as can be judged +by this character, there is no difference between them. +The importance of the comparison can only be appreciated +by examining the bones side by side. In +the upper arm bone of a bird, on what is known +as the ulnar border, near to the shoulder joint, and +on the side nearest to it, is the entrance to the air +cell in the humerus. In the Pterodactyle the corresponding +foramen has the same position, form, and +size, and is not one large hole, but a reticulation +of small perforations, one beyond another, exactly +such as are seen in the entrance to the air cell in the +bone of a bird, in which the pneumatic character +is found. For it is not every bird of flight which has +this pneumatic condition of the bones; and Dr. Crisp +stated that quite a number of birds—the Swallow, +Martin, Snipe, Canary, Wood-wren and Willow-wren, +Whinchat, Glossy-starling, Spotted-fly-catcher, and +Black-headed Bunting—have no air in their bones. +And it is well known that in many birds, especially +water birds, it is only the upper bones of the limbs +which are pneumatic, while the smaller bones retain +the marrow.</p> + +<div class="figcenter" style="width: 366px;"> +<a name="Fig_15" id="Fig_15"></a> +<span class="caption">FIG. 15. HEAD OF THE HUMERUS OF THE PTERODACTYLE ORNITHOCHEIRUS</span> +<img src="images/i_063.jpg" width="366" height="640" alt="FIG. 15." title="FIG. 15." /> +<p class="center">Showing position of the pneumatic foramen on the +ulnar side of the bone as in a bird</p> +</div> + + +<p><span class='pagenum'><a name="Page_48" id="Page_48">[Pg 48]</a></span></p> +<h4>LUNGS AND AIR CELLS</h4> + + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_16" id="Fig_16"></a> +<span class="caption">FIG. 16. LUNGS OF THE BIRD APTERYX +PARTLY OPENED ON THE RIGHT-HAND SIDE</span> +<img src="images/i_065.jpg" width="640" height="420" alt="FIG. 16." title="FIG. 16." /> +<p class="center">The circles are openings of the bronchial tubes on the surface of the lung +The notches on the inner edges of the lungs are impressions of the ribs<br /> + +(After R. Owen)</p> +</div> + + + +<p>It may be well to remember that the lungs of a +bird are differently conditioned from those of any +other animal. Instead of hanging freely suspended +in the cone-shaped chamber of the thorax formed by +the ribs and sternum, they are firmly fixed on each +side, so that the ribs deeply indent them and hold +them in place. The lungs have the usual internal +structure, being made up of branching cells. The +chief peculiarity consists in the way in which the air +passes not only into them, but through them. The +air tube of the throat of a bird, unlike that of a +man, has the organ of voice, not at the upper end +in the form of a larynx, but at the lower end, forming +what is termed a syrinx. There is no evidence +of this in a fossil state, although in a few birds the +rings of the trachæa become ossified, and are preserved. +But below the syrinx the trachæa divides +into two bronchi, tubes which carry the ringed +character into the lungs for some distance, and +these give off branches termed bronchial tubes, the +finer subdivisions from which, in their clustered +minute branching sacs, make up the substance of +the lung. There is nothing exceptional in that. But +towards the outer or middle part of the ventral or +<span class='pagenum'><a name="Page_49" id="Page_49">[Pg 49]</a></span> +under surface of the lungs, four or five rounded +openings are seen on each side. Each of these +openings resembles the entrance of the air cell into +a bone, since it displays several smaller openings +which lead to it. Each opening from the lung +leads to an air cell. Those cells may be regarded as +the blowing out of the membrane which covers the +lungs into a film which holds air like a mass of soap +bubbles, until the whole cavity of the body of a +bird from neck to tail is occupied by sacculated air +cells, commonly ten in number, five on each side, +though two frequently blend at the base of the neck +in the region of the <b>V</b>-shaped bone named the +clavicle or furculum, popularly known as the merry-thought. +Most people have seen some at least of +<span class='pagenum'><a name="Page_50" id="Page_50">[Pg 50]</a></span> +these semi-transparent bladder-like air cells beneath +the skin in the abdominal region of a fowl. The cells +have names from their positions, and on each side +one is abdominal, two are thoracic, one clavicular, +and one cervical, which last is at the base of the +neck. The clavicular and abdominal air cells are +perhaps the most interesting. The air cell termed +clavicular sends a process outward towards the arm, +along with the blood vessels which supply the arm. +Thus this air cell, entering the region of the axilla +or arm-pit, enters the upper arm bone usually on its +under side, close to the articular head of the humerus, +and in the same way the air may pass from bone to +bone through every bone in the fore limb. The hind +limbs similarly receive air from the abdominal air +cell, which supplies the femur and other bones of +the leg, the sacrum, and the tail. But the joints of +the backbone in front of the sacrum receive their air +from the cervical air sac. The air cells are not +limited to the bones, but ramify through the body, +and in some cases extend among the muscles. A +bird may be said to breathe not only with its lungs, +but with its whole body. And it is even affirmed +that respiration has been carried on through a broken +arm bone when the throat was closed, and the bird +under water.</p> + +<div class="figcenter" style="width: 488px;"> +<a name="Fig_17" id="Fig_17"></a> +<span class="caption">FIG. 17. THE BODY OF AN OSTRICH LAID OPEN +TO SHOW THE AIR CELLS WHICH EXTEND +THROUGH ITS LENGTH</span> +<p class="center">(After Georges Roché)</p> +<img src="images/i_066.jpg" width="488" height="480" alt="FIG. 17." title="FIG. 17." /> +</div> + + +<p>Birds differ greatly in the extent to which the aircell +system prolonged from the lungs is developed, +some having the air absent from every bone, while +others, like the Swift, are reputed to have air in every +bone of the body.</p> + +<p>Comparison shows that in so far as the bones are +the same in Bird and Ornithosaur, the evidence of +the air cells entering them extends to resemblance,<span class='pagenum'><a name="Page_51" id="Page_51">[Pg 51]</a></span> +if not coincidence, in every detail. No living group +of animals except birds has pneumatic limb bones, +in relation to the lungs; so that it is reasonable to +conclude that the identical structures in the bones +were due to the same cause in both the living and +extinct groups of animals. It is impossible to say +that the lungs were identical in Birds and Pterodactyles, +but so far as evidence goes, there is no +ground for supposing them to have been different.</p> + + +<h4>THE LUNGS OF REPTILES</h4> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_18" id="Fig_18"></a> +<span class="caption">FIG. 18. THE SIDE OF THE BODY OF A CHAMELEON</span> +<p class="center">Ribs removed to show the sacculate branched form of the lung</p> +<img src="images/i_068.jpg" width="640" height="339" alt="FIG. 18." title="FIG. 18." /> +</div> + + +<p>There is nothing comparable to birds, either in the +lungs of living reptiles or in their relation to the +bones. The Chameleon is remarkable in that the +lung is not a simple bladder prolonged through +the whole length of the body cavity, as in a serpent, +but it develops a number of large lateral branches +visible when the body is laid open. Except near +the trachæa, where the tissue has the usual density +of a lizard lung, the air cell is scarcely more complicated +than the air bladder of a fish, and does not +enter into any bone of the skeleton. And although +<span class='pagenum'><a name="Page_52" id="Page_52">[Pg 52]</a></span> +many fishes like the Loach have the swim bladder +surrounded by bone connected with the head, it offers +no analogy to the pneumatic condition of the bones +in the Pterodactyle.</p> + + +<h4>THE FORM OF THE BRAIN CAVITY</h4> + +<p>But the identity of the pneumatic foramina in +Birds and Flying Reptiles is not a character which +stands by itself as evidence of organisation, for a +mould of the form of the brain case contributes +evidence of another structural condition which throws +some light on the nature of Ornithosaurs. Among +many of the lower animals, such as turtles, the brain +does not fill the chamber in the dry skull, in which +the same bones are found as are moulded upon the +brain in higher animals. For the brain case in such +reptiles is commonly an envelope of cartilage, as +among certain fishes; and except among serpents, +the Ophidia, the bones do not completely close the +reptilian brain case in front. The brain fills the brain +case completely among birds. A mould from its +interior is almost as definite in displaying the several +parts of which it is formed as the actual brain would +be. And the chief regions of the brain in a bird—cerebrum, +optic lobes, cerebellum—show singularly +little variation in proportion or position. The essential +fact in a bird's brain, which separates it absolutely +from all other animals, is that the pair of nerve +masses known as the optic lobes are thrust out at +the sides, so that the large cerebral hemispheres +extend partly over them as they extend between +them to abut against the cerebellum. This remarkable +condition has no parallel among other vertebrate +animals. In Fishes, Amphibians, Reptiles, and +<span class='pagenum'><a name="Page_53" id="Page_53">[Pg 53]</a></span> +Mammals the linear succession of the several parts +of the brain is never departed from; and any appearance +of variation from it among mammals is more +apparent than real, for the linear succession may be +seen in the young calf till the cerebral hemispheres +grow upward and lop backward, so as to hide the +relatively small brain masses which correspond to +the optic lobes of reptiles, extending over these +corpora-quadrigemina, as they are named, so as to +cover more or less of the mass of the cerebellum. +From these conditions of the brain and skull, it +would not be possible to mistake a mould from +<span class='pagenum'><a name="Page_54" id="Page_54">[Pg 54]</a></span> +the brain case of a bird for that of a reptile, though +in some conditions of preservation it is conceivable +that the mould of the brain of a bird might be distinguished +with difficulty from that of the brain in the +lowest mammals. Taken by itself, the avian form of +brain in an animal would be as good evidence that +its grade of organisation was that of a bird as could +be offered.</p> + +<div class="figcenter" style="width: 480px;"> +<a name="Fig_19" id="Fig_19"></a> +<span class="caption">FIG. 19. THE FORM OF THE BRAIN</span> +<img src="images/i_070.jpg" width="480" height="543" alt="FIG. 19." title="FIG. 19." /> +</div> + + +<h4>THE BRAIN IN SOLENHOFEN PTERODACTYLES</h4> + +<p>It happens that moulds of the brain of Pterodactyles, +more or less complete, are met with of +all geological ages—Liassic, Oolitic, and Cretaceous. +The Solenhofen Slate is the only deposit in Europe +in which Pterodactyle skulls can be said to be fairly +numerous. They commonly have the bones so thin +as to show the form of the upper surface of the +mould of the brain, or the bones have scaled off +the mould, or remain in the counterpart slab of stone, +so as to lay bare the shape of the brain mass.</p> + +<p>In the Museum at Heidelberg a skull of this kind +is seen in the long-tailed genus of Pterodactyles +named Rhamphorhynchus. It shows the large +rounded cerebral hemispheres, which extend in +front of cerebral masses of smaller size a little +below them in position, which perhaps are as like +the brain of a monotreme mammal as a bird.</p> + +<p>The short-tailed Pterodactylus described by Cuvier +has the cerebral hemispheres very similar to those +of a bird, but the relations of the hinder parts of +the brain to each other are less clear.</p> + +<p>The first specimen to show the back of the brain +was found by Mr. John Francis Walker, <small>M.A.</small>, in the +Cambridge Greensand. I was able to remove the +<span class='pagenum'><a name="Page_55" id="Page_55">[Pg 55]</a></span> +thick covering of cellular bone which originally +extended above it, and thus expose evidence that +in the mutual relations of the fore and hind parts +of the brain bird and ornithosaur were practically +identical. Another Cambridge Greensand skull +showed that in the genus Ornithocheirus the optic +lobes of the brain are developed laterally, as in birds. +That skull was isolated and imperfect. But about the +same time the late Rev. W. Fox, of Brixton, in the +Isle of Wight, obtained from Wealden beds another +skull, with jaws, teeth, and the principal bones of +the skeleton, which showed that the Wealden Pterodactyle +Ornithodesmus had a similar and bird-like +brain. In 1888 Mr. E. T. Newton, <small>F.R.S.</small>, obtained a +skull from the Upper Lias, uncrushed and free from +distortion. This made known the natural mould of +the brain, which shows the cerebral hemispheres, optic +lobes, and cerebellum more distinctly than in the specimens +previously known. In some respects it recalls +the Heidelberg brain of Rhamphorhynchus in the +apparently transverse subdivision of the optic lobes, +but it is unmistakably bird-like, and quite unlike any +reptile.</p> + + +<h4>IMPORTANCE OF THE BRAIN AND BREATHING +ORGANS</h4> + +<p>So far as the evidence goes, it appears that these +fossil flying animals show no substantial differences +from birds, either in the mould of the brain or the +impress of the breathing organs upon the bones. +These approximations to birds of the nervous and +respiratory systems, which are beyond question two +of the most important of the vital organs of an +animal, and distinctive beyond all others of birds,<span class='pagenum'><a name="Page_56" id="Page_56">[Pg 56]</a></span> +place the naturalist in a singular dilemma. He must +elect whether he will trust his interpretation to the +soft organs, which among existing animals never vary +their type in the great classes of vertebrate animals, +and on which the animal is defined as something +distinct from its envelope the skeleton and its appendages +the limbs, or whether he will ignore them. +The answer must choose substantially between belief +that the existing order of Nature gives warrant for +believing that these vital characteristics which have +been discussed might equally coexist with the skeleton +of a mammal or a reptile, as with that of a bird, +for which there is no particle of evidence in existing +life. Or, as an alternative, the fact must be accepted +that birds only have such vital organs as are here +found, and therefore the skeleton, that may be associated +with them, cannot affect the reference of the +type to the same division of the animal kingdom as +birds. The decision need not be made without further +consideration. But brain and breathing organs of the +avian type are structures of a different order of +stability in most animals from the bones, which vary +to a remarkable extent in almost every ordinal group +of animals.</p> + + +<h4>TEMPERATURE OF THE BLOOD</h4> + +<p>The organs of circulation and digestion are necessarily +unknown. There are reasons why the blood +may have been hot, such as the evidences from the +wings of exceptional activity; though the temperature +depends more upon the amount of blood in the +body than upon the apparatus by which it is distributed. +We speak of a Crocodile as cold-blooded, +yet it is an animal with a four-chambered heart not<span class='pagenum'><a name="Page_57" id="Page_57">[Pg 57]</a></span> +incomparable with that of a bird. On the other hand, +the Tunny, a sort of giant Mackerel, is a fish with a +three-chambered heart, only breathing the air dissolved +in water, which has blood as warm as a +mammal, its temperature being compared to that of +a pig. Several fishes have blood as warm as that of +Manis, the scaly ant-eater; and many birds have +hotter blood than mammals. The term "hot-blooded," +as distinct from "cold-blooded," applied to animals, is +relative to the arbitrary human standard of experience, +and expresses no more than the circumstance +that mammals and birds are warmer animals than +reptiles and fishes.</p> + +<p>The exceptional temperature of the Flying Fish +has led to a vague impression that physical activity +and its effect upon the amount of blood which vigour +of movement circulates, are more important in raising +an animal's temperature than possession of the circulatory +organs commonly associated with hot blood, +which drive the blood in distinct courses through the +body and breathing organs. Yet the kind of heart +which is always associated with vital structures such +as Pterodactyles are inferred to have possessed from +the brain mould and the pneumatic foramina in the +bones, is the four-chambered heart of the bird and +the mammal. Considering these organs alone—of +which the fossil bones yield evidence—we might +anticipate, by the law of known association of structures, +that nothing distinctly reptilian existed in the +other soft part of the vital organisation, because there +is no evidence in favour of or against such a possibility.</p> + + + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_58" id="Page_58">[Pg 58]</a></span></p> +<h2><a name="CHAPTER_VIII" id="CHAPTER_VIII"></a><small>CHAPTER VIII</small><br /><br /> + +THE PLAN OF THE SKELETON</h2> + + +<p>While these animals are incontestably nearer +to birds than to any other animals in their +plan of organisation, thus far no proof has been +found that they are birds, or can be included in +the same division of vertebrate life with feathered +animals. It is one of the oldest and soundest teachings +of Linnæus that a bird is known by its feathers; +and the record is a blank as to any covering to the +skin in Pterodactyles. There is the strongest probability +against feathers having existed such as are +known in the Archæopteryx, because every Solenhofen +Ornithosaur appears to have the body devoid +of visible or preservable covering, while the two birds +known from the Solenhofen Slate deposit are well +clothed with feathers in perfect preservation. We +turn from the skin to the skeleton.</p> + +<p>The plan on which the skeleton is constructed +remains as evidence of the animal's place in nature, +which is capable of affording demonstration on which +absolute reliance would have been placed, if the brain +and pneumatic foramina had remained undiscovered. +With the entire skeleton before us, it is inconceivable +that anatomical science should fail to discover the<span class='pagenum'><a name="Page_59" id="Page_59">[Pg 59]</a></span> +true nature of the animal to which it belonged, by +the method of comparing one animal with another. +There is no lack of this kind of evidence of Pterodactyles +in the three or four scores of skeletons, and +thousands of isolated or associated bones, preserved +in the public museums of Europe and America.</p> + +<p>I may recall the circumstance that the discovery of +skeletons of fossil animals has occasionally followed +upon the interpretation of a single fragment, from +which the animal has been well defined, and sometimes +accurately drawn, before it was ever seen. So +I propose, before drawing any conclusions from the +skeletons in the entirety of their construction, to +examine them bone by bone, and region by region, +for evidence that will manifest the nature of this +brood of Dragons. Their living kindred, and perhaps +their extinct allies, assembled as a jury, may be able +to determine whether resemblances exist between +them, and whether such similarity between the bones +as exists is a common inheritance, or is a common +acquisition due to similar ways of life, and no evidence +of the grade of the organism among vertebrate +animals.</p> + +<p>The bones of these Ornithosaurs, when found +isolated, first have to be separated from the organisms +with which they are associated and mixed in the +geological strata. This discrimination is accomplished +in the first instance by means of the texture of the +surface. The density and polish of the bones is +even more marked than in the bones of birds, and is +usually associated with a peculiar thinness of substance +of the bone, which is comparable to the condition +in a bird, though usually a little stouter, so +that the bones resist crushing better. Pterodactyle<span class='pagenum'><a name="Page_60" id="Page_60">[Pg 60]</a></span> +bones in many instances are recognised by their +straightness and comparatively uniform dimensions, +due to the exceptional number of long bones which +enter into the structure of the wing as compared +with birds. When the bones are unerringly determined +as Ornithosaurian, they are placed side by +side with all the bones which are most like them, till, +judged by the standard of the structures of living +animals, the fossil is found to show a composite construction +as though it were not one animal but many, +while its individual bones often show equally composite +characters, as though parts of the corresponding +bone in several animals had been cunningly fitted +together and moulded into shape.</p> + + +<h4>THE PLAN OF THE HEAD IN ORNITHOSAURS</h4> + +<p>The head is always the most instructive part of an +animal. It is less than an inch long in the small +Solenhofen skeleton named <i>Pterodactylus brevirostris</i>, +and is said to be three feet nine inches long in the +toothless Pterodactyle Ornithostoma from the Chalk +of Kansas. Most of these animals have a long, +slender, conical form of head, tapering to the point +like the beak of a Heron, forming a long triangle +when seen from above or from the side. Sometimes +the head is depressed in front, with the beak flattened +or rounded as in a Duck or Goose, and occasionally in +some Wealden and Greensand species the jaws are +truncated in front in a massive snout quite unlike +any bird. The back of the head is sometimes +rounded as among birds, showing a smooth pear-shaped +posterior convexity in the region of the brain. +Sometimes the back of the head is square and vertical +or oblique. Occasionally a great crest of cellular<span class='pagenum'><a name="Page_61" id="Page_61">[Pg 61]</a></span> +tissue is extended backward from above the brain +case over the spines of the neck bones.</p> + +<p>There are always from two to four lateral openings +in the skull. First, the nostril is nearest to the extremity +of the beak. Secondly, the orbits of the +eyes are placed far backward. These two openings +are always present. The nostril may incline upward. +The orbits of the eyes are usually lateral, though +their upper borders sometimes closely approximate, +as in the woodpecker-like types from the Solenhofen +Slate named <i>Pterodactylus Kochi</i>, now separated +as another genus. In most genera there is an opening +in the side of the head, between the eye hole and the +nostril, known as the antorbital vacuity; and another +opening, which is variable in size and known as the +temporal vacuity, is placed behind the eye. The +former is common in the skulls of birds, the latter is +absent from all birds and found in many reptiles.</p> + +<p>The palate is usually imperfectly seen, but English +and American specimens have shown that it has +much in common with the palate in birds, though it +varies greatly in form of the bones in representatives +from the Lias, Oolites, and Cretaceous rocks.</p> + +<p>From the scientific aspect the relative size of the +head, its form, and the positions and dimensions of +its apertures and processes, are of little importance +in comparison with its plan of construction, as evidenced +by the positions and relations to each other +of the bones of which it is formed. There usually is +some difficulty in stating the limits of the bones of +the skull, because in Pterodactyles, as among birds, +they usually blend together, so that in the adult +animal the sutures between the bones are commonly +obliterated.<span class='pagenum'><a name="Page_62" id="Page_62">[Pg 62]</a></span></p> + +<p>Bones have relations to each other and places in +the head which can only change as the organs with +which they are associated change their positions. No +matter what the position of a nostril may be—at the +extremity of a long snout, as in an ant-eater, or far +back at the top of the head in a porpoise, or at the +side of the head in a bird—it is always bordered by +substantially the same bones, which vary in length +and size with the changing place of the nostril and +the form of the head. Every region of the head is +defined by this method of construction; so that eye +holes and nose holes, brain case and jaw bones, +palate and teeth, beak, and back of the skull are all +instructive to those who seek out the life-history of +these animals. We may briefly examine the head +of an Ornithosaurian.</p> + + +<h4>BONES ABOUT THE NOSTRIL</h4> + +<p>No matter what its form may be, the head of an +Ornithosaur always terminates in front in a single +bone called the intermaxillary. It sends a bar of +bone backward above the visible nostrils, between +them; and a bar on each side forms the margin of +the jaw in which teeth are implanted. The bone +varies in depth, length, sharpness, bluntness, slenderness, +and massiveness. As the bone becomes long +the jaw is compressed from side to side, and the +openings of the nostrils are removed backward to +an increasing distance from the extremity of the +beak.</p> + +<p>The outer and hinder border of the nostril is made +by another bone named the maxillary bone, which is +usually much shorter than the premaxillary. It +contains the hindermost teeth, which rarely differ +<span class='pagenum'><a name="Page_63" id="Page_63">[Pg 63]</a></span> +from those in front, except in sometimes being +smaller.</p> + +<p>The nasal bones, which always make the upper +and hinder border of the nostrils, meet each other +above them, in the middle line of the beak.</p> + +<div class="figcenter" style="width: 594px;"> +<a name="Fig_20" id="Fig_20"></a> +<span class="caption">FIG. 20</span> +<img src="images/i_080.jpg" width="594" height="480" alt="FIG. 20" title="FIG. 20" /> +<p class="center">Showing that the extremity of the jaws in Rhamphorhynchus was +sheathed in horn as in the giant Kingfisher, since the jaws +similarly gape in front.<br /> +<br /> +The hyoid bones are below the lower jaw in the Pterodactyle.</p> +</div> + + +<p>The nostrils are unusually large in the Lias genus +named Dimorphodon, and small in species of the +genus Rhamphorhynchus from Solenhofen. Such +differences result from the relative dimensions and +proportions of these three bones which margin the +nasal vacuity, and by varying growth of their front +margins or of their hinder margins govern the form +of the snout.</p> + +<p>The jaws are most massive in the genera known from +the Wealden beds to the Chalk. The palatal surface is +<span class='pagenum'><a name="Page_64" id="Page_64">[Pg 64]</a></span> +commonly flat or convex, and often marked by an +elevated median ridge which corresponds to a groove +in the lower jaw, though the median ridge sometimes +divides the palate into two parallel concave channels. +The jaw is margined with teeth which are rarely +fewer than ten or more than twenty on each side. +They are sharp, compressed from side to side, curved +inward, and never have a saw-like edge on the back +and front margins. No teeth occur upon the bones +of the palate.</p> + +<p>In most birds there is a large vacuity in the side +of the head between the nostril and the orbit of the +eye, partly separated from it by the bone which +carries the duct for tears named the lachrymal bone. +The same preorbital vacuity is present in all long-tailed +Pterodactyles, though it is either less completely +defined or absent in the group with short +tails. It affords excellent distinctive characters for +defining the genera. In the long-tailed genus +Scaphognathus from Solenhofen this preorbital opening +is much larger than the nostril, while in Dimorphodon +these vacuities are of about equal size. +Rhamphorhynchus is distinguished by the small size +of the antorbital vacuity, which is placed lower than +the nostril on the side of the face. The aperture is +always imperfectly defined in Pterodactylus, and is +a relatively small vacuity compared with the long +nostril. In Ptenodracon the antorbital vacuity +appears to have no existence separate from the nostril +which adjoins the eye hole. And so far as is known at +present there is no lateral opening in advance of the +eye in the skull in any Ornithosaur from Cretaceous +rocks, though the toothless Ornithostoma is the only +genus with the skull complete. When a separate +<span class='pagenum'><a name="Page_65" id="Page_65">[Pg 65]</a></span> +antorbital vacuity exists, it is bordered by the maxillary +bone in front, and by the malar bone behind. +The prefrontal bone is at its upper angle. That bone +is known in a separate state in reptiles and, I think, in +monotreme mammals. Its identity is soon lost in +the mammal, and its function in the skull is different +from the corresponding bone in Pterodactyles.</p> + + +<h4>BONES ABOUT THE EYES</h4> + +<div class="figcenter" style="width: 492px;"> +<a name="Fig_21" id="Fig_21"></a> +<span class="caption">FIG. 21. UPPER SURFACE OF SKULL OF THE HERON</span> +<p class="center">Compared with the same aspect of the skull of Rhamphorhynchus</p> +<img src="images/i_082.jpg" width="492" height="480" alt="FIG. 21." title="FIG. 21." /> +</div> + + +<p>The third opening in the side of the head, counting +from before backward, is the orbit of the eye. In this +vacuity is often seen the sclerotic circle of overlapping +bones formed in the external membrane of the eye, +like those in nocturnal birds and some reptiles. The +<span class='pagenum'><a name="Page_66" id="Page_66">[Pg 66]</a></span> +eye hole varies in form from an inverted pear-shape +to an oblique or transverse oval, or a nearly circular +outline. It is margined by the frontal bone above; +the tear bone or lachrymal, and the malar or cheek +bone in front; while the bones behind appear to be +the quadrato-jugal and post-frontal bones, though the +bones about the eye are somewhat differently arranged +in different genera.</p> + +<p>The eyes were frequently, if not always, in contact +with the anterior walls of the brain case, as in many +birds, and are always far back in the side of the head. +In Dimorphodon they are in front of the articulation +of the lower jaw; in Rhamphorhynchus, above that +articulation; while in Ornithostoma they are behind +the articulation for the jaw. This change is governed +by the position of the quadrate bone, which is vertical +in the Lias genus, inclined obliquely forward in the +fossils from the Oolites, and so much inclined in the +Chalk fossil that the small orbit is thrown relatively +further back.</p> + +<p>Thus far the chief difference in the Pterodactyle +skull from that of a bird is in the way in which the +malar arch is prolonged backward on each side. It is +a slender bar of bone in birds, without contributing +ascending processes to border vacuities in the side +of the face, while in these fossil animals the lateral +openings are partly separated by the ascending processes +of these bones. This divergence from birds, +in the malar bone entering the orbit of the eye +is approximated to among reptiles and mammals, +though the conditions, and perhaps the presence of a +bone like the post-orbital bone, are paralleled only +among Reptiles. The Pterodactyles differ among +themselves enough for the head to make a near +<span class='pagenum'><a name="Page_67" id="Page_67">[Pg 67]</a></span> +approach to Reptiles in Dimorphodon, and to Birds +in Pterodactylus. In the Ground Hornbill and the +Shoebill the lachrymal bones in front of the orbits +of the eyes grow down to meet the malar bars without +uniting with them. The post-frontal region also +is prolonged downward almost as far as the malar +bar, as though to show that a bird might have its +orbital circle formed in the same way and by the +same bones as in Pterodactylus. Cretaceous Ornithosaurs +sometimes differ from birds apparently in admitting +the quadrato-jugal bone into the orbit. It +then becomes an expanded plate, instead of a slender +bar as in all birds.</p> + + +<h4>THE TEMPORAL FOSSA</h4> + +<p>A fourth vacuity is known as the temporal fossa. +When the skull of such a mammal as a Rabbit, or +Sheep, is seen from above, there is a vacuity behind +the orbits for the eyes, which in life is occupied by +the muscles which work the lower jaw. It is made +by the malar bone extending from the back of the +orbit and the process of bone, called the zygomatic +process, extending forward from the articulation of +the jaw, which arches out to meet the malar bone.</p> + +<p>In birds there is no conspicuous temporal fossa, +because the malar bar is a slender rod of bone in a +line with the lower end of the quadrate bone.</p> + +<p>Reptile skulls have sometimes one temporal vacuity +on each side, as among tortoises, formed by a single +lateral bar. These vacuities, which correspond to +those of mammals in position, are seen from the top +of the head, as lateral vacuities behind the orbits +of the eyes, and are termed superior temporal vacuities. +In addition to these there is often in other +<span class='pagenum'><a name="Page_68" id="Page_68">[Pg 68]</a></span> +reptiles a lateral opening behind the eye, termed +the inferior temporal vacuity, seen in Crocodiles, in +Hatteria, and in Lizards; and in such skulls there are +two temporal bars seen in side view, distinguished as +superior and inferior. The superior arch always includes +the squamosal bone, which is at the back +of the single bar in mammals. The lower arch +includes the malar bone, which is in front in the single +arch of mammals. The circumstance that both these +arches are connected with the quadrate bone makes +the double temporal arch eminently reptilian.</p> + +<p>In Ornithosaurs the lateral temporal vacuity varies +from a typically reptilian condition to one which, +without becoming avian, approaches the bird type. In +skulls from the Lias, Dimorphodon and Campylognathus, +there is a close parallel to the living New +Zealand reptile Hatteria, in the vertical position +of the quadrate bone and in the large size of the +vacuity behind and below the eye, which extends +nearly the height of the skull. In the species of the +genus Pterodactylus, the forward inclination of the +quadrate bone recalls the Curlew, Snipe, and other +birds. The back of the head is rounded, and the +squamosal bone, which appears to enter into the +wall of the brain case as in birds and mammals, +is produced more outward than in birds, but less +than in mammals, so as to contribute a little to +the arch which is in the position of the post-frontal +bone of reptiles. It is triangular, and stretches from +the outer angle of the frontal bone at the back of the +orbit to the squamosal behind, where it also meets +the quadrate bone. Its third lower branch meets the +quadratojugal, which rests upon the front of the quadrate +bone, as in Iguanodon, and is unlike Dimorphodon +<span class='pagenum'><a name="Page_69" id="Page_69">[Pg 69]</a></span> +in its connexions. In that genus the supra-temporal +bone, or post-orbital bone, appears to rest upon the +post-frontal and connect it with the quadrato-jugal. +In Dimorphodon the malar bone is entirely removed +from the quadrate, but in Pterodactylus it meets its +articular end. Between the post-frontal bone above +and the quadrato-jugal bone below is a small lunate +opening, which represents the lateral temporal +vacuity; and so far, this is a reptilian character. +But if the thin post-frontal bone were absorbed, +Pterodactylus would resemble birds. There is no +evidence that the quadrate bone is free in any +Ornithosaurs, as it is in all birds, while in Dimorphodon +it unites by suture with the squamosal bone. +In Ornithostoma the lateral temporal vacuity is little +more than a slit between the quadrate bone below, +the quadrato-jugal in front, and what may be the +post-frontal bone behind (see <a href="#Fig_2">Fig. 2, p. 12</a>).</p> + + +<h4>BONES ABOUT THE BRAIN</h4> + +<p>The bones containing the brain appear to be the +same as form the brain case in birds. The form of +the back of the skull varies in two ways. First it +may be flat above and flat at the back, when the +back of the head appears to be square. This condition +is seen in all the long-tailed genera, such as +Campylognathus from the Lias and Rhamphorhynchus, +and is associated with a high position for the +upper temporal bar. Secondly, the back of the head +may be rounded convexly, both above and behind. +That condition is seen in the short-tailed genera, +such as Pterodactylus. But in the large Cretaceous +types, such as Ornithocheirus and Ornithostoma, +the superior longitudinal ridge which runs back in +<span class='pagenum'><a name="Page_70" id="Page_70">[Pg 70]</a></span> +the middle line of the face becomes elevated and +compressed from side to side at the back of the head +as a narrow deep crest, prolonged backward over the +neck vertebræ for some inches of length. All these +three types are paralleled more or less in birds which +have the back of the head square like the Heron, or +rounded like the Woodpecker; or crested, though the +crest of the Cormorant is not quite identical with +Ornithocheirus, being a distinct bone at the back of +the head in the bird which never blends with the +skull. In so far as the crest is reptilian it suggests +the remarkable crest of the Chameleon. In the +structure of the back of the skull the bones are a +modification of the reptilian type of Hatteria in +the Lias genus Campylognathus, but the reptilian +characters appear to be lost in the less perfectly +preserved skulls of Cretaceous genera.</p> + +<p>The palate is well known in the chief groups of +Ornithosaurs, such as Campylognathus, Scaphognathus, +and Cycnorhamphus.</p> + +<p>Mr. E. T. Newton, <small>F.R.S.</small>, has shown that in the +English skull from the Lias of Whitby, the forms of +the bones are similar to the palate in birds and unlike +the conditions in reptiles. There is one feature, however, +which may indicate a resemblance to Dicynodon +and other fossil reptiles from South Africa. A +slender bone extends from the base of the brain case, +named the basi-sphenoid bone, outward and forward +to the inner margin of the quadrate bone (<a href="#Fig_22">Fig. 22</a>). +A bone is found thus placed in those South African +Reptiles, which show many resemblances to the Monotreme +and Marsupial Mammals. It is not an ordinary +element of the skeleton and is unknown in living +animals of any kind in that position. It has been +<span class='pagenum'><a name="Page_71" id="Page_71">[Pg 71]</a></span> +thought possible that it may represent one of the +bones which among mammals are diminutive and +are included in the internal ear. The resemblance +may have some interest hereafter, as helping to show +that certain affinities of the Ornithosaurs may lie +outside the groups of existing reptiles. Instead of +being directed transversely outward, as in the palatal +region of <i>Dicynodon lacerticeps</i>, they diverge outward +and forward to the inner border of the articulation +for the lower jaw which is upon the quadrate +bone.</p> + + +<div class="figcenter" style="width: 635px;"> +<a name="Fig_22" id="Fig_22"></a> +<span class="caption">FIG. 22</span> +<img src="images/i_088.jpg" width="635" height="480" alt="FIG. 22" title="FIG. 22" /> +</div> + + + +<h4>BONES OF THE PALATE</h4> + +<p>There is a pair of bones which extend forward +from these inner articular borders of the quadrate +bones, and converge in a long <b>V</b>-shape till they +merge in the hard palate formed by the bones of the +front of the beak, named intermaxillary and maxillary +bones. The limits of the bones of the palate are<span class='pagenum'><a name="Page_72" id="Page_72">[Pg 72]</a></span> +not distinct, but there can be no doubt that the front +of the <b>V</b> is the bone named vomer, that the palatine +bones are at its sides, and that its hinder parts are +the pterygoid bones as in birds. There is a long, +wide, four-sided, open space in the middle of the +palate, between the vomer and the basi-sphenoid +bone, unlike anything in birds or other animals.</p> + +<p>Professor Marsh, in a figure of the palate in the +great skull of the toothless Pterodactyle named Ornithostoma +(Pteranodon), from the Chalk of Kansas, +found a large oval vacuity in this region of the palate. +In that genus the pterygoid bones meet each other +between the quadrate bones as in Dicynodon (<a href="#Fig_73">Fig. 73, +p. 182</a>). Hence the great palatal vacuity here seen in +the Ornithosaur is paralleled by the small vacuity in +the South African reptile, which is sometimes distinct +and sometimes partly separated from the anterior +part of the vacuity which forms the openings of the +nostrils on the palate.</p> + +<p>The Solenhofen skulls which give any evidence of +the palate are exposed in side view only, and the +bones, imperfectly seen through the lateral vacuities, +are displaced by crushing. They include long strips +like the vomerine bones in the Lias fossil, and they +diverge in the same way as they extend back to the +quadrate bones. The oblique division into vomer in +front and pterygoid bone behind is shown by Goldfuss +in his original figure of Scaphognathus. Thus +there is some reason for believing that all Ornithosaurs +have the palate formed upon the same general plan, +which is on the whole peculiar to the group, especially +in not having the palatal openings of the nares +divided in the middle line. It would appear probable +that the short-tailed animals have the pterygoid bones<span class='pagenum'><a name="Page_73" id="Page_73">[Pg 73]</a></span> +meeting in the middle line and triangular; and that +they are slender rods entirely separate from each +other in the long-tailed genera.</p> + + +<h4>THE TEETH</h4> + +<p>The teeth are all of pointed, elongated shape, without +distinction into the kinds seen in most mammals +and named incisors, canines, and grinders. They are +organs for grasping, like the teeth of the fish-eating +Crocodile of India, and are not unlike the simple teeth +of some Porpoises. They are often implanted in +oblique oval sockets with raised borders, usually at +some distance apart from each other, and have the +crown pointed, flattened more on the outer side than +on the inner side, usually directed forward and curved +inward. As in many extinct animals allied to existing +reptiles, the teeth are reproduced by germs, which +originate on the inner side of the root and grow till +they gradually absorb the substance of the old tooth, +forming a new one in its place. Frequently in Solenhofen +genera, like Scaphognathus and Pterodactylus, +the successional tooth is seen in the jaw on the hinder +border of the tooth in use. There is some variation +in the character of bluntness or sharpness of the +crowns in the different genera, and in their size.</p> + +<p>The name Dimorphodon, given to the animal from +the Lias of Lyme Regis, expresses the fact that the +teeth are of two kinds. In the front of the jaw three or +four large long teeth are found in the intermaxillary +bone on each side, as in some Plesiosaurs, while the +teeth found further back in the maxillary bone are +smaller, and directed more vertically downward. This +difference is more marked in the lower jaw than in the +upper jaw. In Rhamphorhynchus the teeth are all<span class='pagenum'><a name="Page_74" id="Page_74">[Pg 74]</a></span> +relatively long and large, and directed obliquely +forward, but absent from the extremities of the beak, +as in the German genus from the Lias named Dorygnathus, +in which the bone of the lower jaw (which +alone is known) terminates in a compressed spear. +In Scaphognathus the teeth are few, more vertical, +and do not extend backward so far as in Rhamphorhynchus, +but are carried forward to the extremity of +the blunt, deep jaw.</p> + +<p>In the short-tailed Pterodactyles the teeth are +smaller, shorter, wider at the base of the crown, +closer together, and do not extend so far backward +in the jaw. In Ornithocheirus two teeth always +project forward from the front of the jaw. Ornithostoma +is toothless.</p> + + +<h4>SUPPOSED HORNY BEAK</h4> + +<p>Sometimes a horny covering has been suggested +for the beak, like that seen in birds or turtles, but no +such structure has been preserved, even in the Solenhofen +Slate, in which such a structure would seem as +likely to be preserved as a wing membrane, though +there is one doubtful exception. There are marks of +fine blood vessels on some of the jaws, indicating a +tough covering to the bone. In Rhamphorhynchus +the jaws appear to gape towards their extremities as +though the interspace had originally been occupied +by organic substance like a horny beak.</p> + + +<h4>LOWER JAW</h4> + +<p>The lower jaw varies in relative length with the +vertical or horizontal position of the quadrate bone in +the skull. In Dimorphodon the jaw is as long as the +skull; but in the genera from the Oolitic rocks the<span class='pagenum'><a name="Page_75" id="Page_75">[Pg 75]</a></span> +mandible is somewhat shorter, and in Ornithostoma +the discrepancy reaches its maximum. The hinder +part of the jaw is never prolonged backward much +beyond the articulation, differing in this respect from +Crocodiles and Plesiosaurs.</p> + +<p>The depth of the jaw varies. It is slender in +Pterodactylus, and is probably stronger relatively to +the skull in Scaphognathus than in any other form. +It fits between the teeth and bones of the alveolar +border in the skull, in all the genera. In Dimorphodon +its hinder border is partly covered by the +descending edge of the malar process which these +animals develop in common with some Dinosaurs, +and some Anomodont reptiles, and many of the lower +mammals. In this hinder region the lower jaw is +sometimes perforated, in the same way as in Crocodiles. +That condition is observed in Dimorphodon, +but is not found in Pterodactylus. The lower jaw is +always composite, being formed by several bones, as +among reptiles and birds. The teeth are in the +dentary bone or bones, and these bones are almost +always blended as in most birds and Turtles, and not +separate from each other as among Crocodiles, Lizards, +and Serpents.</p> + +<p>An interesting contour for the lower border of the +jaw is seen in Ornithostoma, as made known in +figures of American examples by Professors Marsh +and Williston. It deepens as it extends backwards +for two-thirds its length, stops at an angle, and then +the depth diminishes to the articulation with the +skull. This angle of the lower jaw is a characteristic +feature of the jaws of Mammals. It is seen in the +monotreme Echidna, and is characteristic of some +Theriodont Reptiles from South Africa, which in +<span class='pagenum'><a name="Page_76" id="Page_76">[Pg 76]</a></span> +many ways resemble Mammals. The character is +not seen in the jaws of specimens from the Oolitic +rocks, but is developed in the toothed Ornithocheirus +from the Cambridge Greensand, and is absent from +the jaws of existing reptiles and birds.</p> + + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_23" id="Fig_23"></a> +<span class="caption">FIG. 23. COMPARISON OF THE LOWER JAW IN +ECHIDNA AND ORNITHOSTOMA</span> +<img src="images/i_093.jpg" width="640" height="417" alt="FIG. 23." title="FIG. 23." /> +</div> + + +<h4>SUMMARY OF CHARACTERS OF THE HEAD</h4> + +<p>Taken as a whole, the head differs from other types +of animals in a blending of characters which at the +present day are found among Birds and Reptiles, with +some structures which occur in extinct groups of +animals with similar affinities, and perhaps a slight +indication of features common to the lowest mammals. +It is chiefly upon the head that the diverse views of +earlier writers have been based. Cuvier was impressed +with the reptilian aspect of the teeth; but in +later times discoveries were made of Birds with teeth—Archæopteryx, +Ichthyornis, Hesperornis. The teeth +are quite reptilian, being not unlike miniature teeth +<span class='pagenum'><a name="Page_77" id="Page_77">[Pg 77]</a></span> +of Mosasaurus. If those birds had been found prior +to the discovery of Pterodactyles, the teeth might +have been regarded as a link with the more ancient +birds, rather than a crucial difference between birds +and reptiles.</p> + +<p>All the specimens show a lateral temporal hole in +the bones behind the eye, and this is found in no +bird or mammal, and is typical of such reptiles as +Hatteria. The quadrate bone may not be so decisive +as Cuvier thought it to be, for its form is not unlike +the quadrate of a bird, and different, so far as I have +seen, from that of living reptiles. This region of the +head is reptilian, and if it occurred in a bird the character +would be as astonishing as was the discovery of +teeth in extinct birds. These characters of the head +are also found in fossil animals named Dinosaurs, in +association with many resemblances to birds in their +bones.</p> + +<p>The palate might conceivably be derived from +that of Hatteria by enlarging the small opening in +the middle line in that reptile till it extended forward +between the vomera; but it is more easily compared +with a bird, which the animal resembles in its beak, +and in the position of the nares. Excepting certain +Lizards, all true existing Reptiles have the nostrils +far forward and bordered by two premaxillary bones +instead of one intermaxillary, as in Birds and Ornithosaurs. +If nothing were known of the animal but +its head bones, it would be placed between Reptiles +and Birds.</p> + + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_78" id="Page_78">[Pg 78]</a></span></p> +<h2><a name="CHAPTER_IX" id="CHAPTER_IX"></a><small>CHAPTER IX</small><br /><br /> + +THE BACKBONE, OR VERTEBRAL +COLUMN</h2> + + +<p>The backbone is a more deep-seated part of the +skeleton than the head. It is more protected +by its position, and has less varied functions to perform. +Therefore it varies less in distinctive character +within the limits of each of the classes of vertebrate +animals than either the head or limbs. It is divided +into neck bones, the cervical vertebræ; back bones, +the dorsal vertebræ; loin bones, the lumbar vertebræ; +the sacrum, or sacral vertebræ, which support the +hind limbs; and the tail. Of these parts the tail is +the least important, though it reaches a length in +existing reptiles which sometimes exceeds the whole +of the remainder of the body, and includes hundreds +of vertebræ. It attains its maximum among serpents +and lizards. In frogs it is practically absent. In +some of the higher mammals it is a rudiment, which +does not extend beyond the soft parts of the body.</p> + + +<h4>THE NECK</h4> + +<p>The neck is more liable to vary than the back, with +the habit of life of the animal. And although +mammals almost always preserve the same number +<span class='pagenum'><a name="Page_79" id="Page_79">[Pg 79]</a></span> +of seven bones in the neck, the bones vary in length +between the short condition of the porpoise, in +which the neck is almost lost, and the long bones +which form the neck of the Llama, though even these +may be exceeded by some fossil reptiles like Tanystrophœus. +In many mammals the neck bones do +not differ in length or size from those of the back. +In others, like the Horse and Ox, they are much +broader and larger.</p> + +<p>There is the same sort of variation in the bones of +the neck among birds, some being slender like the +Heron, others broad like the Swan. But there is also +a singular variation in number of vertebral bones +in a bird's neck. At fewest there are nine, which +equals the exceptionally large number found among +mammals in the neck of one of the Sloths. Usually +birds have ten to fifteen cervical vertebræ, and in the +Swan there are twenty-three. Most of the neck bones +of birds are relatively long, and the length of the neck +is often greater than the remainder of the vertebral +column.</p> + +<p>Reptiles usually have short necks. The common +Turtle has eight bones in the neck, ten in the back. +The two regions are sharply defined by the dorsal +shield. Their articular ends are sometimes cupped in +front, in the neck, sometimes cupped behind, or convex +at both ends, or even flattened, or the articulation +may be made exceptionally by the neural arch alone. +Nine is the largest number of neck bones in existing +Lizards, and there are usually nine in Crocodiles; so +that reptiles closely approach mammals in number of +the neck bones. It is remarkable that the maximum +number in a mammal and in living reptiles should +coincide with the minimum number in birds. Therefore +<span class='pagenum'><a name="Page_80" id="Page_80">[Pg 80]</a></span> +the number of cervical vertebræ as an attribute +of Mammal, Bird, or Reptile, can only be important +from its constancy.</p> + +<p>German naturalists affirm on clear evidence that +the Solenhofen Pterodactyles have seven cervical vertebræ. +In many specimens there can be no doubt +about the number, because the neck bones are easily +distinguished from those of the back by their size; +but the number is not always easy to count.</p> + +<p>As in Birds, the first vertebra, or atlas, in Pterodactyles +is extremely short, and is generally—if not +always—blended with the much longer second vertebra, +named the axis. The front of the atlas forms +a small rounded cup to articulate with the rounded +ball of the basioccipital bone at the back of the skull. +The third and fourth vertebræ are longer, but the +length visibly shortens in the sixth and seventh.</p> + +<p>Sometimes the vertebræ are slender and devoid of +strong spinous processes. This is the condition in +the little <i>Pterodactylus longirostris</i> and in the comparatively +large <i>Cycnorhamphus Fraasii</i>, in which +there is a slight median ridge along the upper surface +of the arch of the vertebra. This condition is paralleled +in birds with long necks, especially wading +birds such as the Heron. Other Ornithosaurs, such +as Ornithocheirus from the Cretaceous rocks, have the +neck much more massive. The vertebræ are flattened +on the under side. The arch above the nervous +matter of the spinal cord has a more or less considerable +transverse expansion, and may even be as +wide as long. These vertebræ have proportions and +form such as may be seen in Vultures or in the +Swan. In either case the form of the neck bones +is more or less bird-like, and the neural spine may +<span class='pagenum'><a name="Page_81" id="Page_81">[Pg 81]</a></span> +be elevated, especially in Pterodactyles with long +tails.</p> + +<p>One of the most distinctive features of the neck +bones of a bird is the way in which the cervical ribs +are blended with the vertebræ. They are small, and +each is often prolonged in a needle-like rod at the +side of the neck bone.</p> + +<p>In Ornithocheirus the cervical rib similarly blends +with the vertebra by two articulations, as in mammals, +so that it might escape notice but for the +channel of a blood vessel which is thus inclosed. +In several of the older Pterodactyles from Solenhofen +the ribs of the neck vertebræ remain separated, +as in a Crocodile, though still bird-like in their +form, anterior position, and mode of attachment. In +Terrapins and Tortoises the long neck vertebræ have +no cervical ribs.</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_24" id="Fig_24"></a> +<span class="caption">FIG. 24 UNITED ATLAS AND AXIS OF ORNITHOCHEIRUS</span> +<p class="center">(Cambridge Greensand)</p> +<img src="images/i_098.jpg" width="640" height="337" alt="FIG. 24" title="FIG. 24" /> +</div> + +<p>The articular surfaces between the bodies of the +vertebræ, in the neck, are transversely oval. The middle +part of this articular joint is made by the body of +the vertebra; its outer parts are in the neural arch. +In front this surface is a hollow channel, often more +depressed than in any other animals. The corresponding +surface behind is convex, with a process on +<span class='pagenum'><a name="Page_82" id="Page_82">[Pg 82]</a></span> +each side at its lower outer angles (<a href="#Fig_25">Fig. 25</a>). It is a +modification of the cup-and-ball form of vertebral +articulation, which at the present day is eminently +reptilian. Serpents and Crocodiles have the articulations +similarly vertical, but in both the form of the +articulation is a circle. In Lizards the articular cup is +usually rather wider than deep, when the cup and +ball are developed in the vertebræ; it differs from +the vertical condition in pterodactyles in being oblique +and much narrower from side to side. Only among +Crocodiles and Hatteria is there a double articulation +for the cervical rib, though in neither order have rib +or vertebra in the neck the bird-like proportions +which are usual in these animals. Pterodactyles show +no resemblance to birds in this vertebral articulation. +A Bird has the corresponding surface concave from +side to side in front, but it is also convex from above +downward, producing what is known as the saddle-shaped +form which is peculiarly avian, being found +in existing birds except in part of the back in Penguins. +It is faintly approximated to in one or two +neck vertebræ in man. Professor Williston remarks +that in the toothless Pterodactyles of Kansas the +hinder ball of the vertebral articulation is continued +downward and outward as a concave articulation +upon the processes at its outer corners. There are +no mammals with a cup-and-ball articulation between +the vertebræ, so that for what it is worth the character +now described in Ornithosaurs is reptilian, when +judged by comparison with existing animals.</p> + +<p>Low down on each side of the vertebra, at the +junction of its body with the neural arch, is a large +ovate foramen, transversely elongated, and often a +little impressed at the border, which is the entrance +<span class='pagenum'><a name="Page_83" id="Page_83">[Pg 83]</a></span> +of the air cell into the bone. These foramina are +often one-third of the length of the neck vertebræ +in specimens from the Cambridge Greensand, where +the neck bones vary from three-quarters of an inch +to about two and a half inches in length, and in +extreme forms are as wide as long. The width of +the interspace between the foramina is one-half the +width of the vertebræ, though this character varies +with different genera and species. Several species +from the Solenhofen Slate have the neck long and +slender, on the type of the Flamingo. In others the +neck is thick and short—in the <i>Scaphognathus crassirostris</i> +and <i>Pterodactylus spectabilis</i>. Some genera +with slender necks have the bones preserved with a +curved contour, such as might suggest a neck carried +like that of a Llama or a Camel. The neck is occasionally +preserved in a curve like a capital <b>S</b>, as +though about to be darted forward like that of a +bird in the act of striking its prey. The genera of +Pterodactyles with short necks may have had as great +mobility of neck as is found among birds named +Ducks and Divers; but those Pterodactyles with +stout necks, such as Dimorphodon and Ornithocheirus, +in which the vertebræ are large, appear to +<span class='pagenum'><a name="Page_84" id="Page_84">[Pg 84]</a></span> +have been built more for strength than activity, and +the neck bones have been chiefly concerned in the +muscular effort to use the fighting power of the jaws +in the best way.</p> + + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_25" id="Fig_25"></a> +<span class="caption">FIG. 25. CERVICAL VERTEBRA OF ORNITHOCHEIRUS</span> +<p class="center">From the Cambridge Greensand</p> +<img src="images/i_100.jpg" width="640" height="276" alt="FIG. 25." title="FIG. 25." /> +</div> + + +<h4>THE BACK</h4> + +<p>The region of the back in a Pterodactyle is short +as compared with the neck, and relatively is never +longer than the corresponding region in a bird. The +shortness results partly from the short length of the +vertebræ, each of which is about as long as wide. +There is also a moderate number of bones in the +back. In most skeletons from Solenhofen these +vertebræ between the neck and girdle of hip bones +number from twelve to sixteen. They have a general +resemblance in form to the dorsal vertebræ in birds. +The greatest number of such vertebræ in birds is +eleven. The number is small because some of the +later vertebræ in birds are overlapped by the bones +of the hip girdle, which extend forward and cover +them at the sides, so that they become blended with +the sacrum. This region of the skeleton in the +Dimorphodon from the Lias is remarkable for the +length of the median process, named the neural +spine, which is prolonged upward like the spines of +the early dorsal vertebræ of Horses, Deer, and other +mammals. In this character they differ from living +reptiles, and parallel some Dinosaurs from the Weald. +The bones of the back in Ornithocheirus from the +Cambridge Greensand show the under side to be well +rounded, so that the articular surfaces between the +vertebræ, though still rather wider than deep, are +much less depressed than in the region of the neck. +The neural canal for the spinal cord has become +<span class='pagenum'><a name="Page_85" id="Page_85">[Pg 85]</a></span> +larger and higher, and the sides of the bone are +somewhat compressed. Strong transverse processes +for the support of the ribs are elevated above the +level of the neural canal, at the sides of vertebræ +compressed on the under sides, and directed outward. +Between these lateral horizontal platforms +is the compressed median neural spine, which varies +in vertical height. The articulation of the ribs is not +seen clearly. Isolated ribs from the Stonesfield Slate +have double-headed dorsal ribs, like those of birds. +In some specimens from the Solenhofen Slate like +the Scaphognathus, in the University Museum at +Bonn, dorsal ribs appear to be attached by a notch +in the transverse process of the dorsal vertebra, which +resembles the condition in Crocodiles. Variations in +the mode of attachment of ribs among mammals +may show that character to be of subordinate importance. +Von Meyer has described the first pair +of ribs as frequently larger than the others, and +there appear in Rhamphorhynchus to be examples +preserved of the sternal ribs, which connect the +dorsal ribs with the sternum. Six pairs have been +counted. A more interesting feature in the ribs +consists in the presence behind the sternum, which +is shorter than the corresponding bone in most birds, +of median sternal ribs. They are slender <b>V</b>-shaped +bones in the middle line of the abdomen, which +overlapped the ends of the dorsal ribs like the +similar sternal bones of reptiles. Such structures +are unknown among Birds and Mammals. There is +no trace in the dorsal ribs of the claw-like process, +which extends laterally from rib to rib as a marked +feature in many birds. Its presence or absence may +not be important, because it is represented by fibro-cartilage +<span class='pagenum'><a name="Page_86" id="Page_86">[Pg 86]</a></span> +in the ribs of crocodiles, and may be a small +cartilage near the head of the rib in serpents, and is +only ossified in some ribs of the New Zealand reptile +Hatteria. So that it might have been present in a +fossil animal without being ossified and preserved. +Although the structure is associated with birds, it +is possibly also represented by the great bony plates +which cover the ribs in Chelonians, and combine to +form the shield which covers the turtle's back. The +structure is as characteristic of reptiles as of birds, +but is not necessarily associated with either.</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_26" id="Fig_26"></a> +<span class="caption">FIG. 26</span> +<img src="images/i_103.jpg" width="640" height="441" alt="FIG. 26" title="FIG. 26" /> +<p class="center">The upper figures show the side and back of a dorsal vertebra of +Ornithocheirus compared with corresponding views of the +side and back of a dorsal vertebra of a Crocodile</p> +</div> + +<p>There are two remarkable modifications of the +early dorsal vertebræ in some of the Cretaceous +Pterodactyles. First, in the genus Ornithodesmus +from the Weald the early dorsal vertebræ are blended +together into a continuous mass, like that which is +found in the corresponding region of the living +Frigate-bird, only more consolidated, and similar to +<span class='pagenum'><a name="Page_87" id="Page_87">[Pg 87]</a></span> +that consolidated structure found behind the dorsal +vertebræ, known as the sacrum, made by the blending +of the vertebræ into a solid mass which supports +the hip bones. Secondly, in some of the Cretaceous +genera of Pterodactyles of Europe and America the +vertebræ in the front part of the back are similarly +blended, but their union is less complete; and in +genera Ornithocheirus and Ornithostoma—the former +chiefly English, the latter chiefly American—the +sides of the neural spines are flattened to form an +oval articular surface on each side, which gives +attachment to the flattened ends of their shoulder-blade +bones named the scapulæ. This condition is +found in no other animals. Three vertebræ appear +to have their neural arches thus united together. +The structure so formed may be named the notarium +to distinguish it from the sacrum.</p> + + +<h4>SACRUM</h4> + +<p>For some mysterious reason the part of the backbone +which lies between the bones of the hips and +supports them is termed the sacrum. Among living +reptiles the number of vertebræ in this region is +usually two, as in lizards and crocodiles. There are +other groups of fossil reptiles in which the number +of sacral vertebræ is in some cases less and in other +cases more. There is, perhaps, no group in which the +sacrum makes a nearer approach to that of birds +than is found among these Pterodactyles, although +there are more sacral vertebræ in some Dinosaurs. +In birds the sacral vertebræ number from five to +twenty-two. In bats the number is from five to six. +In some Solenhofen species, such as <i>Pterodactylus +dubius</i> and <i>P. Kochi</i> and <i>P. grandipelvis</i>, the number +<span class='pagenum'><a name="Page_88" id="Page_88">[Pg 88]</a></span> +is usually five or six. The vertebræ are completely +blended. The pneumatic foramina in the sacrum, so +far as they have been observed, are on the under +sides of the transverse processes; +while in the corresponding notarial +structure in the shoulder +girdle the foramina are in front +of the transverse processes. Almost +any placental mammal in +which the vertebræ of the sacral +region are anchylosed together +has a similar sacrum, which +differs from that of birds in the +more complete individuality of +the constituent bones remaining +evident. The transverse processes +in front of the sacrum are +wider than in its hinder part; so +that the pelvic bones which are +attached to it converge as they +extend backward, as among +mammals. The bodies of the vertebræ forming the +sacrum are similar in length to those of the back. +Each transverse process is given off opposite the +body of its own vertebra, but from a lower lateral +position than in the region of the back, in which the +vertebræ are free.</p> + +<div class="figcenter" style="width: 406px;"> +<a name="Fig_27" id="Fig_27"></a> +<span class="caption">FIG. 27. SACRUM OF RHAMPHORHYNCHUS</span> +<img src="images/i_105.jpg" width="406" height="640" alt="FIG. 27." title="FIG. 27." /> +<p class="center">Showing the complete blending +of the vertebræ and ribs as +in a bird, with the well-defined +Iliac bones, produced chiefly +in front of the acetabulum for +the head of the femur.</p> +</div> + + +<p>The hip bones are closely united with the sacrum +by bony union, and rarely appear to come away from +the sacral vertebræ, as among mammals and reptiles, +though this happens with the Lias Pterodactyles. In +the Stonesfield Slate and Solenhofen Slate the slender +transverse processes from the vertebræ blend with the +ilium of the hip girdle, and form a series of transverse +<span class='pagenum'><a name="Page_89" id="Page_89">[Pg 89]</a></span> +foramina on each side of the bodies of the vertebræ. +In the Cambridge Greensand genera the part +of the ilium above the acetabulum for the articular +head of the femur appears to be always broken away, +so that the relation of the sacrum to the pelvis has +not been observed. This character is no mark of +affinity, but only shows that ossification obliterated +sutures among these animals in the same way as +among birds.</p> + +<p>The great difference between the sacrum of a +Pterodactyle and that of a bird has been rendered +intelligible by the excellent discussion of the sacral +region in birds made by Professor Huxley. He +showed that it is only the middle part of the sacrum +of a chicken which corresponds to the true sacrum of +a reptile, and comprises the five shortest of the vertebræ; +while the four in front correspond to those of +the lower part of the back, which either bear no ribs +or very short ribs, and are known as the lumbar +region in mammals, so that the lower part of the +back becomes blended with the sacrum, and thus +reduces the number of dorsal vertebræ. Similarly +the five vertebræ which follow the true sacral vertebræ +are originally part of the tail, and have been +blended with the other vertebræ in front, in consequence +of the extension along them of the bird's +hip bones. This interpretation helps to account for +the great length of the sacrum in many birds, and +also explains in part the singular shortness of the +tail in existing birds. The Ornithosaur sacrum has +neither the lumbar nor the caudal portions of the +sacrum of a bird.</p> +<p><span class='pagenum'><a name="Page_90" id="Page_90">[Pg 90]</a></span></p> + +<h4>THE TAIL</h4> + +<p>The tail is perhaps the least important part of the +skeleton, since it varies in character and length in +different genera. The short tails seen in typical +pterodactyles include as few as ten vertebræ in +<i>Pterodactylus grandipelvis</i> and <i>P. Kochi</i>, and as many +as fifteen vertebræ in <i>Pterodactylus longirostris</i>. The +tails are more like those of mammals than existing +birds, in which there are usually from six to ten +vertebræ terminating in the ploughshare bone. But +just as some fossil birds, like the Archæopteryx, have +about twenty long and slender vertebræ in the tail, +so in the pterodactyle Rhamphorhynchus this region +becomes greatly extended, and includes from thirty-eight +to forty vertebræ. In Dimorphodon the tail +vertebræ are slightly fewer. The earliest are very +short, and then they become elongated to two or +three times the length of the early tail vertebræ, and +finally shorten again towards the extremity of the +tail, where the bones are very slender. In all long-tailed +Ornithosaurians the vertebræ are supported +and bordered by slender ossified ligaments, which +extend like threads down the tail, just as they do +in Rats and many other mammals and in some +lizards.</p> + +<p>Professor Marsh was able to show that the extremity +of the tail in Rhamphorhynchus sometimes +expands into a strong terminal caudal membrane of +four-sided somewhat rhomboidal shape. He regards +this membrane as having been placed vertically. It +is supported by delicate processes which represent +the neural spines of the vertebræ prolonged upward. +They are about fifteen in number. A corresponding +<span class='pagenum'><a name="Page_91" id="Page_91">[Pg 91]</a></span> +series of spines on the lower border, termed chevron +bones, equally long, were given off from the junctions +of the vertebræ on their under sides, and produced +downward. This vertical appendage is of some +interest because its expansion is like the tail of a +fish. It suggests the possibility of having been used +in a similar way to the caudal fin as an organ for +locomotion in water, though it is possible that it may +have also formed an organ used in flight for steering +in the air.</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_28" id="Fig_28"></a> +<span class="caption">FIG. 28. EXTREMITY OF THE TAIL OF +<i>RHAMPHORHYNCHUS PHYLLURUS</i> (<span class="smcap">Marsh</span>)</span> +<img src="images/i_108.jpg" width="640" height="249" alt="FIG. 28." title="FIG. 28." /> +<p class="center">Showing the processes on the upper and under sides of the vertebræ +which make the terminal leaf-like expansion</p> +</div> + + +<p>The tail vertebræ from the Cambridge Greensand +are mostly found isolated or with not more than four +joints in association. They are very like the slender +type of neck vertebræ seen in long-necked pterodactyles, +but are depressed, and though somewhat +wider are not unlike the tail vertebræ of the Rhamphorhynchus. +The pneumatic foramen in them is a +mere puncture. They have no transverse processes +or neural spines, nor indications of ribs, or chevron +bones.</p> + +<p>The hindermost specimens of tail vertebræ observed +have the neural arch preserved to the end, as among +reptiles; whereas in mammals this arch becomes +lost towards the end of the tail. The processes +by which the vertebræ are yoked together are +<span class='pagenum'><a name="Page_92" id="Page_92">[Pg 92]</a></span> +small. There is nothing to suggest that the tail was +long, except the circumstance that the slender caudal +vertebræ are almost as long as the stout cervical +vertebræ in the same animal. No small caudal +vertebræ have ever been found in the Cambridge +Greensand. The tail is very short, according to +Professor Williston, in the toothless Ornithostoma +in the Chalk of Kansas.</p> + + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_93" id="Page_93">[Pg 93]</a></span></p> +<h2><a name="CHAPTER_X" id="CHAPTER_X"></a><small>CHAPTER X</small><br /><br /> + +THE HIP-GIRDLE AND HIND +LIMB</h2> + + +<p>The bones of the hip-girdle form a basin which +incloses and protects the abdominal vital organs. +It consists on each side of a composite bone, the +unnamed bones—<i>ossa innominata</i> of the older anatomists—which +are each attached to the sacrum on +their inner side, and on the outer side give attachment +to the hind limbs. As a rule three bones enter +into the borders of this cup, termed the acetabulum, in +which the head of the thigh bone, named the Femur, +moves with a more or less rotary motion.</p> + +<p>There are a few exceptions in this division of the +cup between three bones, chiefly among Salamanders +and certain Frogs. In Crocodiles the bone below the +acetabular cup is not divided into two parts. And +in certain Plesiosaurs from the Oxford Clay—Murænosaurus—the +actual articulation appears to be made +by two bones—the ilium and ischium. The three +bones which form each side of the pelvis are known +as the ilium, or hip bone, sometimes termed the aitch +bone; secondly, the ischium, or sitz bone, being the +bone by which the body is supported in a sitting +position; and thirdly the pubis, which is the bone in +<span class='pagenum'><a name="Page_94" id="Page_94">[Pg 94]</a></span> +front of the acetabulum. The pubic bones meet in +the middle line of the body on the under side of the +pelvis in man, and on each side are partly separated +from the ischia by a foramen, spoken of as the +obturator foramen, which in Pterodactyles is minute +and almost invisible, when it exists.</p> + +<p>There is often a fourth bony element in the pelvis. +In some Salamanders a single cartilage is directed +forward, and forked in front. According to Professor +Huxley something of this kind is seen in the Dog. +The pair of bones which extend forward in front +of the pelvis in Crocodiles may be of the same kind, +in which case they should be called prepubic bones. +But among the lower mammals named marsupials +a pouch is developed for the protection of the young +and supported by two slender bones attached to the +pubes, and these bones have long been known as +marsupial bones. In a still lower group of mammalia +named monotremata, which lay eggs, and in many +ways approximate to reptiles and birds, stronger +bones are developed on the front edge of the pubes, +and termed prepubic bones. They do not support a +marsupium.</p> + +<p>Naturalists have been uncertain as to the number +of bones in the pelvis of Pterodactyles, because the +bones blend together early in life, as in birds. Some +follow the Amphibian nomenclature, and unite the +ischium and pubis into one bone, which is then +termed ischium, when the prepubis is termed the +pubis, and regarded as removed from the acetabulum. +There is no ground for this interpretation, for the +sutures are clear between the three pelvic bones in +the acetabulum in some specimens, like <i>Cycnorhamphus +Fraasii</i>, from Solenhofen, and some examples +<span class='pagenum'><a name="Page_95" id="Page_95">[Pg 95]</a></span> +of Ornithocheirus from the Cambridge Greensand. +Pterodactyles all have prepubic bones, which are +only known in Ornithorhynchus and Echidna among +mammals, and are absent from the higher mammals +and birds. They are unknown in any other existing +animals, unless present in Crocodiles, in which ischium +and pubis are always undivided. Therefore it is +interesting to examine the characters of the Ornithosaurian +pelvis.</p> + +<p>The acetabulum for the head of the femur is imperforate, +being a simple oval basin, as in Chelonian +reptiles and the higher Mammals. It never shows +the mark of the ligamentous attachment to the head +of the femur, which is seen in Mammals. In Birds +the acetabulum is perforated, as in many of the fossils +named Dinosaurs, and in Monotremata.</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_29" id="Fig_29"></a> +<span class="caption">FIG. 29. COMPARISON OF THE LEFT SIDE OF THE +PELVIS IN A BIRD AND A PTERODACTYLE</span> +<img src="images/i_112.jpg" width="640" height="259" alt="FIG. 29." title="FIG. 29." /> +</div> + + +<p>Secondly, the ilium is elongated, and extends quite +as much in front of the acetabulum as behind it. +The bone is not very deep in this front process. +Among existing animals this relation of the bone is +nearer to birds than to any other type, since birds +alone have the ilium extended from the acetabulum +in both directions. The form of the Pterodactyle +ilium is usually that of the embryo bird, and its +slender processes compare in relative length better +with those of the unhatched fowl and Apteryx of +<span class='pagenum'><a name="Page_96" id="Page_96">[Pg 96]</a></span> +New Zealand than with the plate-like form in adult +birds.</p> + +<p>In mammals the ilium is directed forward, and +even in the Cape ant-eater Orycteropus there is only +an inappreciable production of the bone backward +behind the acetabulum. Among reptiles the general +position of the acetabulum is at the forward termination +of the ilium, though the Crocodile has some +extension of the bone in both directions, without +forming distinct anterior and posterior processes. +This anterior and posterior extension of the ilium +is seen in the Theriodont reptiles of Russia and of +South Africa, as well as in Dinosaurs.</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_30" id="Fig_30"></a> +<span class="caption">FIG. 30. LEFT PELVIC BONES WITH PREPUBIC BONE IN<br /> +<i>PTERODACTYLUS LONGIROSTRIS</i></span> +<img src="images/i_113.jpg" width="640" height="195" alt="FIG. 30." title="FIG. 30." /> +</div> + +<p>Thirdly, in all pterodactyles the ischium and pubis +are more or less completely blended into a sheet of +bone, unbroken by perforation, though there is usually +a minute vascular foramen; or the lower border may +be notched between the ischium and the pubis, as +in some of the Solenhofen species, and the pubis +does not reach the median line of the body. But +in Dimorphodon the pelvic sheet of bone is unbroken +by any notch or perforation. The notch between +the ischium and pubis is well marked in <i>Pterodactylus +longirostris</i>, and better marked in <i>Pterodactylus dubius</i>, +<i>Cycnorhamphus Fraasii</i>, and Rhamphorhynchus. The +fossil animals which appear to come nearest to the +Pterodactyles in the structure of the pelvis are +<span class='pagenum'><a name="Page_97" id="Page_97">[Pg 97]</a></span> +Theriodonts from the Permian rocks of Russia. The +type known as Rhopalodon has the ilium less prolonged +front and back, and is much deeper than in any +Pterodactyle; but the acetabulum is imperforate, and +the ischium and pubis are not always completely +separated from each other by suture. In the pelvis +referred to the Theriodont Deuterosaurus there is +some approximation to the pelvis of Rhamphorhynchus +and of <i>Pterodactylus dubius</i> in the depth +of the division between the pubis and ischium.</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_31" id="Fig_31"></a> +<span class="caption">FIG. 31 PELVIS AND PREPUBIC BONES OF RHAMPHORHYNCHUS</span> +<img src="images/i_114.jpg" width="640" height="240" alt="FIG. 31" title="FIG. 31" /> +<p class="center">On the left-hand side the two prepubic bones are separate. On +the right-hand they are united into a transverse bar which +overlaps the front of pelvis seen from the under side</p> +</div> + + +<p>There are three modifications of the Ornithosaurian +pelvis. First, the type of Rhamphorhynchus, +in which the pubis and ischium are inclined somewhat +backward, and in which the two prepubic bones +are triangular, and are often united together to form +a transverse bow in front of the pubic region.</p> + +<p>Secondly, there is the ordinary form of pelvis in +which the pubis and ischium usually unite with each +other down their length, as in Dimorphodon, but +sometimes, as in <i>Pterodactylus dubius</i>, divide immediately +below the acetabulum. All these types +possess the paddle-shaped prepubic bones, which are +never united in the median line.</p> + +<p>Thirdly, there is the cretaceous form indicated by +<span class='pagenum'><a name="Page_98" id="Page_98">[Pg 98]</a></span> +Ornithocheirus and Ornithostoma, in which the +posterior half of the ilium is modified in a singular +way, since it is more elevated towards the sacrum +than the anterior half, suggesting the contour of the +upper border of the ilium in a lizard. Without being +reptilian—the anterior prolongation of the bone +makes that impossible—it suggests the lizards. This +type also possesses prepubic bones. They appear, +according to Professor Williston, to be more like +the paddle-shaped bones of Pterodactylus than like +the angular bones in Rhamphorhynchus. The prepubic +bones are united in the median line as in +Rhamphorhynchus. But their median union in that +genus favours the conclusion that the bones were +united in the median line in all species, though they +are only co-ossified in these two families.</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_32" id="Fig_32"></a> +<span class="caption">FIG. 32. THE PELVIC BONES OF AN ALLIGATOR SEEN FROM BELOW</span> +<img src="images/i_115.jpg" width="640" height="403" alt="FIG. 32." title="FIG. 32." /> +<p class="center">The bones in front are here regarded as prepubic, but are commonly +named pubic</p> +</div> + + +<p>This median union of the prepubic bones is a +difference from those mammals like the Ornithorhynchus +and Echidna, which approach nearest to +the Reptilia. In them the prepubic bones have a long +<span class='pagenum'><a name="Page_99" id="Page_99">[Pg 99]</a></span> +attachment to the front margin of the pubis, and +extend their points forward without any tendency +for the anterior extremities to approximate or unite. +The marsupial mammals have the same character, +keeping the marsupial bones completely distinct +from each other at their free extremities. The +only existing animals in which an approximation +is found to the prepubic bones in Pterodactyles +are Crocodiles, in bones which most writers term the +pubic bones. This resemblance, without showing +any strong affinity with the Crocodilia, indicates +that Crocodiles have more in common with the +fossil flying animals than any other group of existing +reptiles; for other reptiles all want prepubic bones, +or bones in front of the pubic region.</p> + + +<h4>THE HIND LIMB</h4> + +<p>The hind limb is exceptionally long in proportion +to the back. This is conspicuous in the skeletons of +the short-tailed Pterodactyles, and is also seen in +Dimorphodon. In Rhamphorhynchus the hind limb +is relatively much shorter, so that the animal, when +on all fours, may have had an appearance not unlike +a Bat in similar position. The limb is exceptionally +short in the little <i>Ptenodracon brevirostris</i>. The +bones of the hind limb are exceptionally interesting. +One remarkable feature common to all the specimens +is the great elongation of the shin bones relatively to +the thigh bones. The femur is sometimes little more +than half the length of the tibia, and always shorter +than that bone. The proportions are those of +mammals and birds. Some mammals have the leg +shorter than the thigh, but mammals and birds +alone, among existing animals, have the proportions +<span class='pagenum'><a name="Page_100" id="Page_100">[Pg 100]</a></span> +which characterise Pterodactyles. The foot appears +to have been applied to the ground not always as in +a bird, but more often in the manner of reptiles, or +mammals in which the digits terminate in claws.</p> + + +<h4>THE FEMUR</h4> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_33" id="Fig_33"></a> +<span class="caption">FIG. 33. THE FEMUR</span> +<img src="images/i_117.jpg" width="640" height="345" alt="FIG. 33." title="FIG. 33." /> +<p class="center">On the right is a front view of femur of a bear. In the middle are front and +side views of the femur of Ornithocheirus. On the left is the femur +of Echidna. These comparisons illustrate the mammalian +characters of the Pterodactyle thigh bone</p> +</div> + + +<p>The thigh bone, on account of the small size of +many of the specimens, is not always quite clear +evidence as an indication of technical resemblance to +other animals. The bone is always a little curved, +has always a rounded, articular head, and rounded +distal condyles. Its most remarkable features are +shown in the large, well-preserved specimens from +the Cambridge Greensand. The rounded, articular +head is associated with a constricted neck to the +bone, followed by a comparatively straight shaft with +distal condyles, less thickened than in mammals. No +bird is known, much less any reptile, with a femur +like Ornithocheirus. Only among Mammals is a +similar bone known with a distinct neck; and only +a few mammals have the exceptional characters of +<span class='pagenum'><a name="Page_101" id="Page_101">[Pg 101]</a></span> +the rounded head and constricted neck at all +similar to the Cretaceous Pterodactyles. A few +types, such as the higher apes, the Hyrax, and +animals especially active in the hind limb, have a +femur at all resembling the Pterodactyle in the pit +for the obturator externus muscle, behind the trochanter +major, such as is seen in a small femur from +Ashwell. The femur varies in different genera, so as +to suggest a number of mammalia rather than any +particular animal for comparison. These approximations +may be consequences of the ways in which +the bones are used. When functional modifications +of the skeleton are developed, so as to produce +similar forms of bones, the muscles to which they +give attachment, which act upon the bones, and +determine their growth, are substantially the same. +In the <i>Pterodactylus longirostris</i> the femur corresponds +in length to about eleven dorsal vertebræ. +The end next the shin bone is less expanded than +is usual among Mammals, and rather suggests an +approach to the condition in Crocodiles, in the moderate +thickness and breadth of the articular end, and +the slight development of the terminal pulley-joint. +One striking feature of the femur is the circumstance +that the articular head, as compared with the distal +end, is directed forward and very slightly inward and +upward. So that allowing for the outward divergence +of the pelvic bones, as they extend forward, there +must have been a tendency to a knock-kneed approximation +of the lower ends of the thigh bones, +as in Mammals and Birds, rather than the outward +divergence seen in Reptiles.</p> + +<p>Apparently the swing of the leg and foot, as it +hung on the distal end of the femur, must have +<span class='pagenum'><a name="Page_102" id="Page_102">[Pg 102]</a></span> +tended rather to an inward than to an outward +direction, so that the feet might be put down upon +the same straight line; this arrangement suggests +rapid movement.</p> + + +<h4>TIBIA AND FIBULA</h4> + +<div class="figcenter" style="width: 512px;"> +<a name="Fig_34" id="Fig_34"></a> +<span class="caption">FIG. 34. COMPARISON OF THE TIBIA AND FIBULA +IN ORNITHOSAUR AND VULTURE</span> +<img src="images/i_119.jpg" width="512" height="480" alt="FIG. 34." title="FIG. 34." /> +</div> + + +<p>In <i>Pterodactylus longirostris</i> the tibia is slender, +more than a fifth longer than the femur. A crest is +never developed at the proximal end, like that seen +in the Guillemot and Diver and other water birds. +The bone is of comparatively uniform thickness down +the shaft in most of the Solenhofen specimens, as in +most birds. At the distal end the shin bone commonly +has a rounded, articular termination, like that +seen in birds. This is conspicuous in the <i>Pterodactylus +grandis</i>. In other specimens the tarsal bones, +which form this pulley, remain distinct from the tibia; +and the upper row of these bones appears to consist +<span class='pagenum'><a name="Page_103" id="Page_103">[Pg 103]</a></span> +of two bones, like those which in many Dinosaurs +combine to form the pulley-like end of the tibia +which represents the bird's drum-stick bone. They +correspond with the ankle bones in man named +astragalus and os calcis.</p> + +<p>Complete English specimens of tibia and fibula are +found in the genus Dimorphodon from the Lias, in +which the terminal pulley of the distal end has some +expansion, and is placed forward towards the front of +the tibia, as in some birds. The rounded surface of +the pulley is rather better marked than in birds. +The proximal end of the shaft is relatively stout, and +is modified by the well-developed fibula, which is a +short external splint bone limited to the upper half +of the tibia, as in birds; but contributing with it to +form the articular surface for the support of the +lower end of the femur, taking a larger share in that +work than in birds. Frequently there is no trace of +the fibula visible in Solenhofen specimens as preserved; +or it is extremely slender and bird-like, as in +<i>Pterodactylus longirostris</i>. In Rhamphorhynchus it +appears to extend the entire length of the tibia, as in +Dinosaurs. In the specimens from the Cambridge +Greensand there is indication of a small proximal crest +to the tibia with a slight ridge, but no evidence that +this is due to a separate ossification. The patella, or +knee-cap, is not recognised in any fossil of the group. +There is no indication of a fibula in the specimens +thus far known from the Chalk rocks either of Kansas +in America, or in England.</p> + +<p>The region of the tarsus varies from the circumstance +that in many specimens the tibia terminates +downward in a rounded pulley, like the drum-stick of +a bird; while in other specimens this union of the +<span class='pagenum'><a name="Page_104" id="Page_104">[Pg 104]</a></span> +proximal row of the tarsal bones with the tibia does +not take place, and then there are two rows of +separate tarsal bones, usually with two bones in each +row. When the upper row is united with the tibia +the lower row remains distinct from the metatarsus, +though no one has examined these separate tarsal +bones so as to define them.</p> + + +<h4>THE FOOT</h4> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_35" id="Fig_35"></a> +<span class="caption">FIG. 35. METATARSUS AND DIGITS IN THREE TYPES +OF ORNITHOSAURS</span> +<img src="images/i_121.jpg" width="640" height="393" alt="FIG. 35." title="FIG. 35." /> +</div> + +<p>The foot sometimes has four toes, and sometimes +five. There are four somewhat elongated, slender +metatarsal bones, which are separate from each other +and never blended together, as in birds. There has +been a suspicion that the metatarsal bones were +separate in the young Archæopteryx. In the young +of many birds the row of tarsal bones at the proximal +end of the metatarsus comes away, and there is a +partial division between the metatarsal bones, though +they remain united in the middle. And among Penguins, +in which the foot bones are applied to the +ground instead of being carried in the erect position +of ordinary birds, there is always a partial separation +<span class='pagenum'><a name="Page_105" id="Page_105">[Pg 105]</a></span> +between the metatarsal bones, though they become +blended together. The Pterodactyle is therefore +different from birds in preserving the bones distinct +through life, and this character is more like Reptiles +than Mammals. The individual bones are not +like those of Dinosaurs, and diverge in Rhamphorhynchus +as though the animals were web-footed. +There is commonly a rudimentary fifth metatarsal. +It is sometimes only a claw-shaped appendage, like +that seen in the Crocodile. It is sometimes a short +bone, completely formed, and carrying two phalanges +in Solenhofen specimens: though no trace of these +phalanges is seen in the large toothless Pterodactyles +from the Cretaceous rocks of North America. In the +<i>Pterodactylus longirostris</i> the number of foot bones +on the ordinary digits is two, three, four, five, as in +lizards; but the short fifth metatarsal has only two +toe bones. In Dimorphodon the fifth digit was bent +upward, and supported a membrane for flight. There +are slight variations in the number of foot bones. +In the species <i>Pterodactylus scolopaciceps</i> the number +of bones in the toes follows the formula two, three, +three, four. In <i>Pterodactylus micronyx</i> the number is +two, three, three, three. The terminal claws are much +less developed than is usual with Birds; and there is +a difference from Bats in the unequal length of the +digits. Taken as a whole, the foot is perhaps more +reptilian than avian, and in some genera is crocodilian.</p> + +<p>The foot is the light foot of an active animal. Von +Meyer thought that the hind legs were too slender +to enable the animal to walk on land; and Professor +Williston, of the University of Kansas, remarks that +the rudimentary claws and weak toes indicate that +<span class='pagenum'><a name="Page_106" id="Page_106">[Pg 106]</a></span> +the animal could not have used the feet effectively +for grasping, while the exceedingly free movement +of the femur indicates great freedom of movement of +the hind legs; and he concludes that the function +of the legs was chiefly for guidance in flight through +their control over the movements, and expresses his +belief that the animal could not have stood upon the +ground with its feet. There may be evidence to +sustain other views. If the limb bones are reconstructed, +they form limbs not wanting in elegance +or length. If it is true, as Professor Williston suggests, +that the weight of his largest animals with the +head three feet long, and a stretch of wing of eighteen +or nineteen feet, did not exceed twenty pounds, there +can be no objection to regarding these animals as +quadrupeds, or even as bipeds, on the ground of the +limbs lacking the strength necessary to support the +body. The slender toes of many birds, and even the +two toes of the ostrich, may be thought to give less +adequate support for those animals than the metatarsals +and digits of Pterodactyles.</p> + + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_107" id="Page_107">[Pg 107]</a></span></p> +<h2><a name="CHAPTER_XI" id="CHAPTER_XI"></a><small>CHAPTER XI</small><br /><br /> + +SHOULDER-GIRDLE AND +FORE LIMB</h2> + + +<h4>STERNUM</h4> + +<p>The sternum is always a distinguishing part of +the bony structure of the breast. In Crocodiles +it is a cartilage to which the sternal ribs unite; and +upon its front portion a flat knife-like bone called +the interclavicle is placed. In lizards like the Chameleon, +it is a lozenge-shaped structure of thin bony +texture, also bearing a long interclavicle, which supports +the clavicular bones, named collar bones in +man, which extend outward to the shoulder blades. +Among mammals the sternum is usually narrow and +flat, and often consists of many successive pieces in +the middle line, on the under side of the body. +Among Bats the anterior part is somewhat widened +from side to side, to give attachment to the collar +bones, but the sternum still remains a narrow bone, +much narrower than in Dolphins, and not differing +in character from many other Mammals, notwithstanding +the Bat's power of flight. The bone develops +a median keel for the attachment of the +muscles of the breast, but something similar is seen +in burrowing Insectivorous mammals like the Moles. +<span class='pagenum'><a name="Page_108" id="Page_108">[Pg 108]</a></span> +So that, as Von Meyer remarked, the presence of a +keel on the sternum is not in itself sufficient evidence +to prove flight.</p> + +<p>Among birds the sternum is greatly developed. +Broad and short in the Ostrich tribe, it is devoid of +a keel; and therefore the keel, if present in a bird, +is suggestive of flight. The keel is differently developed +according to the mode of attachment of the +several pectoral muscles which cover a bird's breast. +In several water birds the keel is strongly developed +in front, and dies away towards the hinder part of +the sternum, as in the Cormorant and its allies. The +sternum in German Pterodactyles is most nearly +comparable to these birds.</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_36" id="Fig_36"></a> +<span class="caption">FIG. 36. COMPARISON OF THE STERNUM</span> +<img src="images/i_125.jpg" width="640" height="376" alt="FIG. 36." title="FIG. 36." /> +</div> + +<p>In the Solenhofen Slate the sternum is fairly well +preserved in many Ornithosaurs. It is relatively +shorter than in birds, and is broader than long; but +not very like the sternum of reptile or mammal in +form. The keel is limited to the anterior part of the +shield of the sternum, as in Merganser and the Cormorant, +and is prolonged forward for some distance in +advance of it. Von Meyer noticed the resemblance of +this anterior process to the interclavicle of the Croco<span class='pagenum'><a name="Page_109" id="Page_109">[Pg 109]</a></span>dile +in position; but it is more like the keel of a bird's +sternum, and is not a separate bone as in Reptiles. +In Pterodactyles from the Cretaceous rocks, the side +bones, called coracoids, are articulated to saddle-shaped +surfaces at the hinder part of the base of +this keel, which are parallel in Ornithocheirus, as in +most birds, but overlap in Ornithodesmus, as in +Herons and wading birds.</p> + +<div class="figcenter" style="width: 484px;"> +<a name="Fig_37" id="Fig_37"></a> +<span class="caption">FIG. 37. STERNUM IN ORNITHOCHEIRUS FROM THE +CAMBRIDGE GREENSAND</span> +<img src="images/i_126.jpg" width="484" height="480" alt="FIG. 37." title="FIG. 37." /> +<p class="center">Showing the strong keel and the facets for the coracoid bones on its hinder +border above the lateral constrictions</p> +</div> + +<p>The keel was pneumatic, and when broken is seen +to be hollow, and appears to have been exceptionally +high in Rhamphorhynchus, a genus in which the +wing bones are greatly elongated. Von Meyer found +<span class='pagenum'><a name="Page_110" id="Page_110">[Pg 110]</a></span> +in Rhamphorhynchus on each side of the sternum a +separate lateral plate with six pairs of sternal ribs, +which unite the sternum with the dorsal ribs, as in +the young of some birds. The hinder surface of the +sternum is imperfectly preserved in the toothless +Pterodactyles of Kansas. Professor Williston states +that the bone is extremely thin and pentagonal in +outline, projecting in front of the coracoids, in a +stout, blunt, keel-like process, similar to that seen in +the Pterodactyles of the Cambridge Greensand. +American specimens have not the same notch behind +the articulation for the coracoid to separate it +from the transverse lateral expansion of the sternal +shield. The lateral margin in the Cambridge Greensand +specimens figured by Professor Owen and myself +is broken; but Professor Williston had the good +fortune to find on the margin of the sternum the +articular surfaces which gave attachment to the sternal +ribs. The margin of the sternal bone thickens at these +facets, four of which are preserved. The sternum in +Ornithostoma was about four and a half inches long +by less than five and a half inches wide. The median +keel extends forward for rather less than two inches, +while in the smaller Cambridge species of Ornithocheirus +it extends forward for less than an inch and +a half.</p> + +<p>A sternum of this kind is unlike that of any other +animal, but has most in common with a bird; and +may be regarded as indicating considerable power +of flight. The bone cannot be entirely attributed to +the effect of flight, since there is no such expanded +sternal shield in Bats. The small number of sternal +ribs is even more characteristic of birds than mammals +or reptiles.</p> +<p><span class='pagenum'><a name="Page_111" id="Page_111">[Pg 111]</a></span></p> + +<h4>THE SHOULDER-GIRDLE</h4> + +<p>The bones which support the fore limb are one +of the distinctive regions of the skeleton defining +the animal's place in nature. Among most of the +lower vertebrata, such as Amphibians and Reptiles, +the girdle is a double arch—the arch of the +collar bone or clavicles in front, and the arch of the +shoulder-blade or scapula behind. The clavicular +arch, when it exists, is formed of three or five parts—a +medium bar named the interclavicle, external to +which is a pair of bones called clavicles, reaching to +the front of the scapulæ when they are present; and +occasionally there is a second pair of bones called +supraclavicles, extending from the clavicles up the +front margins of the scapulæ. Thus the clavicular +arch is placed in front of the scapular arch. The +supraclavicles are absent from all living Reptiles, and +the clavicles are absent from Crocodiles. The interclavicle +is absent from all mammals except Echidna +and Ornithorhynchus. Clavicles also may be absent +in some orders of mammals. Hence the clavicular +arch may be lost, though the collar bones are retained +in man.</p> + +<p>The scapular arch also is more complicated and +more important in the lower than in the higher +vertebrata. It may include three bones on each side +named coracoid, precoracoid, and scapula. But in +most vertebrates the coracoid and precoracoid appear +never to have been segmented so as to be separated +from each other; and it is only among extinct types +of reptiles, which appear to approximate to the Monotreme +mammals, that separate precoracoid bones are +found; though among most mammals, probably, +<span class='pagenum'><a name="Page_112" id="Page_112">[Pg 112]</a></span> +there are stages of early development in which precoracoids +are represented by small cartilages, though +few mammals except Edentata like the Sloths and +Ant-eaters, retain even the coracoids as distinct bones. +Therefore, excepting the Edentata and the Monotremes, +the distinctive feature of the mammalian +shoulder-girdle appears to be that the limbs are supported +by the shoulder-blades, termed the scapulæ.</p> + +<p>Among reptiles there are several distinct types +of shoulder-girdle. Chelonians possess a pair of +bones termed coracoids which have no connexion +with a sternum; and their scapulæ are formed of two +widely divergent bars, divided by a deeper notch than +is found in any fossil reptiles. Among Lizards both +scapula and coracoid are widely expanded, and the +coracoid is always attached to the sternum. Chameleons +have the blade of the scapula long and slender, +but the coracoid is always as broad as it is long. +Crocodiles have the bone more elongated, so that it +has somewhat the aspect of a very strong first sternal +rib when seen on the ventral face of the animal. The +bone is perforated by a foramen, which would probably +lie in the line of separation from the precoracoid +if any such separation had ever taken place. The +scapula, or shoulder-blade, of Crocodiles is a similar +flat bone, very much shorter than the scapula of a +Chameleon, and more like that of the New Zealand +Hatteria. Thus there is very little in common between +the several reptilian types of shoulder-girdle.</p> + +<p>In birds the apparatus for the support of the wings +has a far-off resemblance to the Crocodilian type. +The coracoid bones, instead of being directed laterally +outward and upward from the sternum, as among +Crocodiles, are directed forward, so as to prolong the +<span class='pagenum'><a name="Page_113" id="Page_113">[Pg 113]</a></span> +line of the breast bone, named the sternum. The +bird's coracoid is sometimes flattened towards the +breast bone among Swans and other birds; yet as a +rule the coracoid is a slender bar, which combines +with the still more slender and delicate blade of the +scapula, which rests on the ribs, to make the articulation +for the upper arm bone. Among reptiles the +scapula and coracoid are more or less in the same +straight line, as in the Ostrich, but in birds of flight +they meet at an angle which is less than a right angle, +and where they come in contact the external surface +is thickened and excavated to make the articulation +for the head of the humerus. There is nothing like +this shoulder-girdle outside the class of birds, until it +is compared with the corresponding structure in these +<span class='pagenum'><a name="Page_114" id="Page_114">[Pg 114]</a></span> +extinct animals called Pterodactyles. The resemblance +between the two is surprising. It is not +merely the identity of form in the coracoid bone and +the scapula, but the similar angle at which they meet +and the similar position of the articulation for the +humerus. Everything in the Pterodactyle's shoulder-girdle +is bird-like, except the absence of the representative +of the clavicles, that forked <b>V</b>-shaped bone +of the bird which in scientific language is known as +the furculum, and is popularly termed the "merry-thought." +This kind of shoulder-girdle is found in +the genera from the Lias and the Oolitic rocks, both +of this country and Germany.</p> + + +<div class="figcenter" style="width: 553px;"> +<a name="Fig_38" id="Fig_38"></a> +<span class="caption">FIG. 38. COMPARISON OF SCAPULA AND CORACOID IN +THREE PTERODACTYLES AND A BIRD</span> +<img src="images/i_130.jpg" width="553" height="480" alt="FIG. 38." title="FIG. 38." /> +</div> + + + + +<p>In the Cretaceous rocks the scapula presents, in +most cases, a different appearance. The coracoid is +an elongated, somewhat triangular bone, compressed +on the outer margin as in birds, but differing alike +from birds and other Pterodactyles in not being +prolonged forward beyond the articulation for the +humerus. In these Cretaceous genera, toothed and +toothless alike, the articulation for the upper arm +bone truncates the extremity of the coracoid, so that +the bone is less like that of a bird in this feature. +Perhaps it shows a modification towards the crocodilian +direction. The scapula, which unites with the +coracoid at about a right angle, is similarly truncated +by the articular surface for the humerus; but the +bone is somewhat expanded immediately beyond the +articulation, and compressed; and instead of being +directed backward, it is directed inward over the ribs +to articulate with the neural arches of the early +dorsal vertebræ in the genera found in strata associated +with the Chalk. As the bone approaches +this articulation, it thickens and widens a little, +<span class='pagenum'><a name="Page_115" id="Page_115">[Pg 115]</a></span> +becoming suddenly truncated by an ovate facet, +which exactly corresponds to the transversely ovate +impression, concave from front to back, which is seen +in the neural arches of the dorsal vertebræ on which +it fits. This condition is not present in all Cretaceous +Pterodactyles. It does not occur in the Kansas fossil, +named by Professor Marsh, Nyctodactylus. And it +<span class='pagenum'><a name="Page_116" id="Page_116">[Pg 116]</a></span> +appears to be absent from the Pterodactyles of the +English Weald, named Ornithodesmus.</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_39" id="Fig_39"></a> +<span class="caption">FIG. 39. THE NOTARIUM</span> +<img src="images/i_132a.jpg" width="640" height="252" alt="FIG. 39. THE NOTARIUM" title="FIG. 39." /> +<p class="center">An ossification which gives attachment to the scapulæ seen in +the early dorsal vertebra of Ornithocheirus<br /> +(From the Cambridge Greensand)</p> +</div> + + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_40" id="Fig_40"></a> +<span class="caption">FIG. 40. RESTORATION OF THE SHOULDER-GIRDLE IN THE +CRETACEOUS ORNITHOCHEIRUS</span> +<img src="images/i_132b.jpg" width="640" height="466" alt="FIG. 40." title="FIG. 40." /> +<p class="center">Showing how the scapulæ articulate with a vertebra and the articulation +of the coracoids with the sternum. The humeral articulation with +the coracoid is unlike the condition shown in other Ornithosaurs</p> +</div> + + +<p>There is no approach to this transverse position of +the scapulæ among birds. And while the form of +the bones in the older genera of Ornithosaurs is +singularly bird-like, the angular arrangement in this +Cretaceous genus is obtained by closely approximating +the articulations on the sternum, so that the +coracoids extend outward as in reptiles, instead of +forward as in birds; and the extremities of the +scapulæ similarly approximate towards each other. +This rather recalls the relative positions of scapula +and coracoid among crocodiles. If crocodile and +bird had been primitive types of animals instead of +surviving types, it might almost seem as though +there had been a cunning and harmonious blending +of one with the other in evolving this form of +shoulder-girdle.</p> + + +<h4>THE FORE LIMB</h4> + +<p>The bones of the fore limb, generally, correspond +in length with the similar parts of the hind limb. +The upper arm bone corresponds with the upper leg +bone, and the fore-arm bone is as long as the fore-leg +bone; then differences begin. The bones which +correspond to the back of the hand in man, termed +the metacarpus, are variable in length in Pterodactyles—sometimes +very long and sometimes short. The +wing metacarpal bone is always stout, and the others +are slender. The extremity of the metacarpus was +applied to the ground. Three small digits of the +hand are developed from the three small metacarpal +bones, and terminate in large claws.</p> + +<p>The great wing finger was bent backward, and only +<span class='pagenum'><a name="Page_117" id="Page_117">[Pg 117]</a></span> +touched the ground where it fitted upon the wing +metacarpal bone. It appears sometimes to have +been as long as the entire vertebral column.</p> + +<p>Owing to the circumstance that the joint in the +arm in Pterodactyles was not at the wrist as among +birds, but between the metacarpus and the phalanges, +it follows that the fore limb was longer than the hind +limb when the metacarpus was long; but the difference +would not interfere with the movements of the +animal, either upon four feet or on two feet, for in bats +and birds the disproportion in length is greater.</p> + + +<h4>HUMERUS OR UPPER ARM BONE</h4> + +<p>The first bone in the fore-arm, the humerus, is +remarkable chiefly for the compressed crescent form +of its upper articular end, which is never rounded +like the head of the upper arm bone in man, and +secondly for the great development of the external +process of bone near that end, termed the radial +crest. Sir Richard Owen compared the bone to the +humerus of both birds and crocodiles, but in its upper +articular end the crocodile bone may be said to be +more like a bird than it is like the Pterodactyle. In +flying reptiles the articular surface next to the shoulder-girdle +is somewhat saddle-shaped, being concave from +side to side above and convex vertically, while most +animals with which it can be compared have the +articular head of the bone convex in both directions. +A remarkable exception to this general rule is found +in some fossil animals from South Africa, which, from +resemblance to mammals in their teeth, have been +termed Theriodonts. They sometimes have the head +of the bone concave from side to side and convex in +the vertical direction. To this condition Ornithorhynchus +<span class='pagenum'><a name="Page_118" id="Page_118">[Pg 118]</a></span> +makes a slight approximation. The singular +expansion of the structure called the radial crest +finds no close parallel in reptiles, though Crocodiles +have a moderate crest on the humerus in the same +position; and in Theriodonts the radial crest extends +much further down the shaft of the humerus. No +bird has a radial crest of a similar kind, though it +is prolonged some way down the shaft in Archæopteryx. +In Pterodactyles it sometimes terminates +outward in a smooth, rounded surface, which might +have been articular if any structure could have articulated +with it. There is also a moderate expansion of +the bone on the ulnar side in some Pterodactyles, so +that the proximal end often incloses nearly three-fourths +of an ovate outline. The termination of the +radial crest is at the opposite end of this oval to the +wider articular part of the head of the bone, in +some specimens from the Cambridge Greensand. The +radial crest is more extended in Rhamphorhynchus. +All specimens of the humerus show a twist in the +length of the bone, so that the end towards the fore-arm, +which is wider than the shaft, makes a right +angle with the radial crest on the proximal end, +which is not seen in birds. The shaft of the humerus +is always stouter than that of the femur, though +different genera differ in this respect.</p> + +<p>The humerus in genera from rocks associated with +the Chalk presents two modifications, chiefly seen in +the characters of the distal end of the bone. One of +these is a stout bone with a curiously truncated end +where it joins the two bones of the fore-arm; and +the other is more or less remarkable for the rounded +form of the distal condyles. Both types show distinct +articular surfaces. The inner one is somewhat oblique +<span class='pagenum'><a name="Page_119" id="Page_119">[Pg 119]</a></span> +and concave, the outer one rounded; the two being +separated by a concave channel, so that the ulna +makes an oblique articulation with the bone as in +birds, and the radius articulates by a more or less +truncated or concave surface.</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_41" id="Fig_41"></a> +<span class="caption">FIG. 41. COMPARISON OF THE HUMERUS IN +PTERODACTYLE AND BIRD</span> +<img src="images/i_136.jpg" width="640" height="434" alt="FIG. 41." title="FIG. 41." /> +</div> + + + + +<h4>ULNA AND RADIUS</h4> + +<p>The bones of the fore-arm are similar to each other +in size, and if there be any difference between them +the ulna is slightly the larger. There is some evidence +that in Rhamphorhynchus the upper end of the ulna +was placed behind the radius, probably in consequence +of the mode of attachment of those bones to the +humerus. The ulna abutted towards the inner and +lower border, while the radius was towards the upper +border, consequent upon the twist in the humerus. +This condition corresponds substantially with the +arrangement in birds, but differs from birds in the +relatively more important part taken by the radius +in making the articulation. The bones are compared +in Dimorphodon with the Golden Eagle drawn of the +same size (<a href="#Fig_42">Fig. 42</a>). In birds the ulna supports the great +feathers of the wing, and this may account for the +<span class='pagenum'><a name="Page_120" id="Page_120">[Pg 120]</a></span> +size of the bone. The ulna is best seen at its proximal +end in the specimens from the Cambridge +Greensand, where there is a terminal olecranon ossification +forming an oblique articulation, which frequently +comes away and is lost. It is sometimes +well preserved, and indicated by a suture. The +examples of ulna from the Lias show a slight expansion +of the bone at both ends, and at the distal +end toward the wrist the articulation is well defined, +where the bone joins the carpus. The larger specimens +of the bone are broken. The distal articular +surface is only connected with the proximal end of +the bone in small specimens: it always shows on +the one margin a concavity, followed by a prominent +boss, and an oblique articulation beyond the boss. +On the side towards the radius, on the lower end of +the shaft there is an angular ridge, which marks the +<span class='pagenum'><a name="Page_121" id="Page_121">[Pg 121]</a></span> +line along which the ulna overlaps the radius. The +lower end of the radius has a simple, slightly convex +articulation, somewhat bean-shaped. No rotation +of these bones on each other was possible as in +man. There is a third bone in the fore-arm. This +bone, named the pteroid, is commonly seen in skeletons +from Solenhofen. It was regarded by Von +Meyer as having supported the wing membrane in +flight. Some writers have interpreted it as an essential +part of the Pterodactyle skeleton, and Von +Meyer thought that it might possibly indicate a fifth +digit in the hand. The only existing structure at all +like it is seen in the South African insectivorous +mammal named <i>Chrysochloris capensis</i>, the golden +mole, which also has three bones in the fore-arm, +the third bone extending half-way up towards the +humerus. In that animal the third bone appears to +be behind the others and adjacent to the ulna. In +the German fossils the pteroid articulated with a +separate carpal or metacarpal bone, placed on the side +of the arm adjacent to the radius, and the radius +is always more inward than the ulna. If the view +suggested by Von Meyer is adopted, this bone would +be a first digit extending outward and backward +towards the humerus. That view was adopted by +Professor Marsh. It involves the interpretation of +what has been termed the lateral carpal as the first +metacarpal bone, which would be as short as that +of a bird, but turned in the opposite direction backward. +The first digit would then only carry one +phalange, and would not terminate in a claw, but lie +in the line of the tendon which supports the anterior +wing membrane of a bird.</p> + +<div class="figcenter" style="width: 552px;"> +<a name="Fig_42" id="Fig_42"></a> +<span class="caption">FIG. 42. COMPARISON OF THE BONES OF THE FORE-ARM +IN BIRD AND ORNITHOSAUR</span> +<img src="images/i_137.jpg" width="552" height="480" alt="FIG. 42." title="FIG. 42." /> +</div> + + +<p>The third bone in the fore-arm of Chrysochloris +<span class='pagenum'><a name="Page_122" id="Page_122">[Pg 122]</a></span> +does not appear to correspond to a digit. The bone +is on the opposite side of the arm to the similar +bone of a Pterodactyle, and therefore cannot be the +same structure in the Golden Mole. The interpretation +which makes the pteroid bone the first digit +has the merit of accounting for the fifth digit of the +hand. All the structures of the hand are consistent +with this view. The circumstance that the bone is +rarely found in contact with the radius, but diverging +from it, shows that it plays the same part in stretching +the membrane in advance of the arm, that the fifth +digit holds in supporting the larger wing membrane +behind the arm.</p> + +<p>According to Professor Williston, the American +toothless Pterodactyle Ornithostoma has but a single +phalange on the corresponding first toe of the hind +foot, and that bone he describes as long, cylindrical, +gently curved, and bluntly pointed. There is some +support for this interpretation; but I have not seen +any English or German Pterodactyles with only one +phalange in the first toe.</p> + +<p>The wing in Pterodactyles would thus be stretched +between two fingers which are bent backward, the +three intermediate digits terminating in claws.</p> + + +<h4>THE CARPUS</h4> + +<p>The wrist bones in the reptilia usually consist of +two rows. In Crocodiles, in the upper row there is +a large inner and a small outer bone, behind which +is a lunate bone, the remainder of the carpus being +cartilaginous. Only one carpal is converted into +bone in the lower row. It is placed immediately +under the smaller upper carpal. In Chelonians, the +turtle and tortoise group, the characters of the carpus +<span class='pagenum'><a name="Page_123" id="Page_123">[Pg 123]</a></span> +vary with the family. In the upper row there are +usually two short carpals, which may be blended, +under the ulna; while the two under the radius are +commonly united. The lower row is made up of +several small bones. Lizards, too, usually have three +bones in the proximal row and five smaller bones +in the distal row.</p> + +<p>The correspondence of the distal carpals with +the several metacarpal bones of the middle hand +is a well-known feature of the structure of the +wrist.</p> + +<p>Von Meyer remarks that the carpus is made up of +two rows of small bones in the Solenhofen Pterodactyles; +while in birds there is one row consisting +of two bones. The structure of the carpus is not +distinct in all German specimens; but in the short-tailed +Solenhofen genera the bones in the two rows +retain their individuality.</p> + +<p>In all the Cretaceous genera the carpal bones of +each row are blended into a single bone, so that two +bones are superimposed, which may be termed the +proximal and distal carpals. One specimen shows +by an indication of sutures the original division of +the distal carpal into three bones; and the separated +constituent bones are very rarely met with. Two +bones of the three confluent elements contribute to the +support of the wing metacarpal, and the third gives +an articular attachment to the bone which extends +laterally at the inner side of the carpus, which I +now think may be the first metacarpal bone turned +backward towards the humerus. The three component +bones meet in the circular pneumatic foramen +in the middle of the under side of the distal +carpal. There is no indication of division of the +<span class='pagenum'><a name="Page_124" id="Page_124">[Pg 124]</a></span> +proximal carpal in these genera into constituent +bones.</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_43" id="Fig_43"></a> +<span class="caption">FIG. 43. CARPUS FROM ORNITHOCHEIRUS</span> +<p class="center">(Cambridge Greensand)</p> +<img src="images/i_141.jpg" width="640" height="254" alt="FIG. 43." title="FIG. 43." /> +</div> + + + +<p>This condition is somewhat different from birds. +In 1873 Dr. Rosenberg, of Dorpat, showed that +there is in the bird a proximal carpal formed of two +elements, and a distal carpal also formed of two +elements. Therefore the two constituents of the +distal carpal in the bird which blends in the mature +animal with the metacarpus, forming the rounded +pulley joint, may correspond with two of the three +bones in the Cretaceous Pterodactyle <i>Ornithocheirus.</i></p> + +<p>The width of a proximal carpal rarely exceeds two +inches, and that of a distal carpal is about an inch +and three-quarters. Two such bones when in contact +would not measure more than one inch in depth. +The lower surface shows that the wing had some +rotary movement upon the carpus outward and +backward.</p> + + +<h4>METACARPUS</h4> + +<p>The metacarpus consists of bones which correspond +to the back of the hand. The first digit of +the hand in clawed animals has the metacarpal bone +short, or shorter than the others. Among mammals +metacarpal bones are sometimes greatly elongated; +and a similar condition is found in Pterodactyles, in +which the metacarpal bone may be much longer +<span class='pagenum'><a name="Page_125" id="Page_125">[Pg 125]</a></span> +than the phalange which is attached to it. Two +metacarpal bones appear to be singularly stouter +than the others. The first bone of the first digit, if +rightly determined, is much shorter than the others, +and is, in fact, no longer than the carpus (<a href="#Fig_43">Fig. 43</a>). It +is a flat oblong bone, attached to the inner side of +the lower carpal, and instead of being prolonged +distally in the same direction as the other metacarpal +bones, is turned round and directed upward, +so that its upper edge is flush with the base of the +radius, and gives attachment to a bone which resembles +a terminal phalange of the wing finger. +According to this interpretation it is the first and +only phalange in the first digit. The bone is often +about half as long as the fore-arm, terminates upward +in a point, is sometimes curved, and frequently +diverges outward from the bones of the fore-arm, +as preserved in the associated skeleton, being +stretched towards the radial crest of the humerus. +This mode of attachment of the supposed first metacarpal, +which is true for all Cretaceous pterodactyles, +has not been shown to be the same for all those +from the Solenhofen Slate. There is no greater +anomaly in this metacarpal and phalange on the +<span class='pagenum'><a name="Page_126" id="Page_126">[Pg 126]</a></span> +inner side being bent backward, than there is in the +wing finger being bent backward on the outer side. +The three slender intervening digits extend forward +between them, as though they were applied to the +ground for walking.</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_44" id="Fig_44"></a> +<span class="caption">FIG. 44. METACARPUS IN TWO ORNITHOSAURS</span> +<img src="images/i_142.jpg" width="640" height="336" alt="FIG. 44." title="FIG. 44." /> +</div> + + +<p>The bone which is usually known as the wing +metacarpal is frequently stouter at the proximal end +towards the carpus than towards the phalange. At +the carpal end it is oblong and truncated, with a short +middle process, which may have extended into the +pit in the base of the carpal bone; while the distal +terminal end is rounded exactly like a pulley. There +is great difference in the length of the metacarpus. +In the American genus Ornithostoma it is much +longer than the fore-arm. In Rhamphorhynchus it +is remarkably short, though perhaps scarcely so +short as in Dimorphodon or in Scaphognathus. The +largest Cretaceous examples are about two inches +wide where they join the carpus. The bone is sometimes +a little curved.</p> + +<p>Between the first and fifth or wing metacarpal are +the three slender metacarpal bones which give attachment +to the clawed digits. They bear much the +same relation to the wing metacarpal that the large +metatarsal of a Kangaroo has to the slender bones +of the instep which are parallel to it.</p> + +<p>The facet for the wing metacarpal on the carpus is +clearly recognised, but as a rule there is no surface +with which the small metacarpals can be separately +articulated. One or two exceptional specimens from +the Cambridge Greensand appear to have not only +surfaces for the wing metacarpal, but two much +smaller articular surfaces, giving attachment to +smaller metacarpals; while in one case there appears +<span class='pagenum'><a name="Page_127" id="Page_127">[Pg 127]</a></span> +to be only one of these additional impressions. It +is certain that all the animals from the Lias and +Oolites have three clawed digits, but at present I +have seen no evidence that there were three in the +Cretaceous genera, though Professor Williston's statements +and restoration appear to show that the +toothless Pterodactyles have three. Another difference +from the Oolitic types, according to Professor +Williston, is in the length of the slender metacarpals +of the clawed phalanges being about one-third that +of the wing metacarpal, but this is probably due to +imperfect ossification at the proximal end; for at the +distal end the bones all terminated on the same level, +showing that the four outer digits were applied to +the ground to support the weight of the body. The +corresponding bone in the Horse and Oxen is carried +erect, so as to be in a vertical line with the bones of +the fore-arm; and the same position prevails usually, +though not invariably, with the corresponding bone +in the hind limb, while in many clawed mammals the +metacarpus and metatarsus are both applied upon the +ground. In Pterodactyles the metatarsal bones are +preserved in the rock in the same straight line with +the smaller bones of the foot, or make an angle with +the shin bone, leading to the conviction that the bones +of the foot were applied to the ground as in Man, +and sometimes as in the Dog, and were thus modified +for leaping. Just as the human metacarpus is extended +in the same line with the bones of the fore-arm, +and the movement of jointing occurs where the +fingers join the metacarpus, so Pterodactyles also +had these bones differently modified in the fore and +hind limbs for the functions of life. The result is to +lengthen the fore limb as compared with the hind +<span class='pagenum'><a name="Page_128" id="Page_128">[Pg 128]</a></span> +limb by introducing into it an elevation above the +ground which corresponds to the length of the metacarpus, +always supposing that the animal commonly +assumed the position of a quadruped when upon the +earth's surface.</p> + +<p>This position of the metacarpus is a remarkable +difference from Birds, because when the bird's wing +is at rest it is folded into three portions. The upper +arm bone extends backward, the bones of the fore-arm +are bent upon it so as to extend forward, and +then at the wrist the third portion, which includes +the metacarpus and finger bones, is bent backward. +So that the metacarpus in the Pterodactyle differs from +birds in being in the same line as the bones of the +fore-arm, whereas in birds it is in the same line with +the digit bones of the hand. It is worthy of remark +that in Bats, which are so suggestive of Pterodactyles +in some features of the hand, the metacarpals and +phalanges are in the same straight line; so that in +this respect the bat is more like the bird. But Pterodactyles +in the relation of these bones to flight are +quite unlike any other animal, and have nothing in +common with the existing animals named Reptiles.</p> + + +<h4>THE HAND</h4> + +<p>From what has just been said it follows that the +construction of the hand is unique. It may be contrasted +with the foot of a bird. The bone which +is called, in the language of anatomists, the tarso-metatarsus, +and is usually free from feathers and +covered with skin, is commonly carried erect in birds, +so that the whole body is supported upon it; and +from it the toes diverge outward. It is formed in +birds of three separate bones blended together. In +<span class='pagenum'><a name="Page_129" id="Page_129">[Pg 129]</a></span> +the fore limb of the Pterodactyle the metacarpus +has the same relation to the bones of the fore-arm +that the metatarsus has to the corresponding bones +of the leg in a bird. But the three metacarpal bones +in the Pterodactyle remain distinct from each other, +perhaps because the main work of that region of the +skeleton has devolved upon the digit called the wing +finger, which is not recognised in the bird. In the +Pterodactyles from the Solenhofen Slate there is +a progressive number of phalanges in the three small +digits of the hand, which were applied to the ground. +This number in the great majority of species follows +the formula of two bones in the first, three bones in +second, and four in the third; so that in the innermost +of the clawed digits only one bone intervenes between +the metacarpal and the claw. The fingers slightly +increase in length with increase in number of bones +which form them.</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_45" id="Fig_45"></a> +<span class="caption">FIG. 45. CLAW PHALANGE FROM THE HAND IN ORNITHOCHEIRUS.<br /> +(Half natural size)</span> +<img src="images/i_146.jpg" width="640" height="208" alt="FIG. 45." title="FIG. 45." /> +</div> + + +<p>The terminal claw bones are unlike the claws of +Birds or Reptiles. They are compressed from side to +side, and extremely deep and strong, with evidence of +powerful attachment for ligaments, so that they rather +resemble in their form and large size the claws of +some of the carnivorous fossil reptiles, often grouped +as Dinosauria, such as have been termed Aristosuchus +and Megalosaurus. In the hand of the +Ostrich the first and second digits terminate in +claws, while the third is without a claw. But these +<span class='pagenum'><a name="Page_130" id="Page_130">[Pg 130]</a></span> +claws of the Ostrich and other birds are slender, +curved, and rather feeble organs. In the Archæopteryx, +a fossil bird which agrees with the Pterodactyles +in retaining the separate condition of the metacarpal +bones and in having the same number of phalanges +in two of the fingers of the fore limb, the terminal +claws are rather more compressed from side to side, +and stronger than in the Ostrich, but not nearly so +strong as in the Pterodactyle. The Archæopteryx +differs from the Pterodactyle in having no trace of a +wing finger. The first metacarpal bone is short, +as in all birds; and the first phalange scarcely +lengthens that segment of the first digit of the Bird's +hand to the same length as the other metacarpal +bones. It therefore was not bent backward like the +first digit in Pterodactyles. The wing finger, from +which the genius of Cuvier selected the scientific +name—Pterodactyle—for these fossils, yields their +<span class='pagenum'><a name="Page_131" id="Page_131">[Pg 131]</a></span> +most distinctive character. It is a feature which could +only be partly paralleled in the Bat, by making +changes of structure which would remove every +support to the wing but the outermost digit of that +animal's hand. In the Bat's hand the membrane for +flight is extended chiefly by four diverging metacarpal +bones. There are only two or three phalanges in each +digit in its four wing fingers. In Pterodactyles the +metacarpal bones are, as we have seen, arranged in +close contact, and take no part in stretching the wing.</p> + + +<div class="figcenter" style="width: 599px;"> +<a name="Fig_46" id="Fig_46"></a> +<span class="caption">FIG. 46. METACARPUS AND DIGITS OF THE HAND +IN BIRDS WITH CLAWS</span> +<img src="images/i_147.jpg" width="599" height="480" alt="FIG. 46." title="FIG. 46." /> +</div> + + + +<h4>THE WING FINGER</h4> + +<p>In Birds there is nothing whatever to represent the +wing finger of the Pterodactyle, for it is an organ +external to the finger bones of the bird, and contains +four phalanges. The first phalange is quite different +from the others. Its length is astonishing when compared +with the small phalanges of the clawed fingers. +The articular surface, which joins on to the wing +metacarpal bone, is a concave articulation, which fits +the pulley in which that bone ends. The pulley +articulation admits of an extension movement in +one direction only. Many specimens show the wing +finger to be folded up so as to extend backward. +The whole finger is preserved in other specimens +straightened out so as to be in line with the metacarpus. +This condition is well seen in Professor +Marsh's specimen of Rhamphorhynchus, which has +the wing membrane preserved, in which all bones +of the fore-arm metacarpus and wing finger are +extended in a continuous curve. The outer surface +of the end of the first bone of the wing finger +overlaps the wing metacarpal, so that a maximum +of strength and resistance is provided in the bony +<span class='pagenum'><a name="Page_132" id="Page_132">[Pg 132]</a></span> +structures by which the wing is supported. There +is, therefore, in flight only one angular bend in the +limb, and that is between the upper arm bone and +the fore-arm.</p> + +<p>An immense pneumatic foramen is situate in a +groove on the under side of the upper end of the +first phalange in Ornithocheirus, but is absent in +specimens from the Kimeridge clay. This bone is +long and stout. It terminates at the lower end in +an obliquely truncated articular surface. Specimens +occur in the Cambridge Greensand which are 2 inches +broad at the upper end and nearly 1½ inch wide at +the lower end. An imperfect bone from the Chalk +is 14½ inches long. The bones are all flattened. +Specimens from the Chalk of Kansas at Munich are +28 inches long. The second phalange is concave at +the upper articular end and convex in the longer +direction at the lower end. The articular points of +union between the several phalanges form prominences +on the under side of the finger in consequence +of the adjacent bones being a little widened at their +junction. It should be mentioned that there is a +proximal epiphysis or separate bone to the first +phalange, adjacent to the pulley joint of the metacarpal +bone, which is like the separate olecranon process +of the ulna of the fore-arm. It sometimes comes +away in specimens from the Chalk and Cambridge +Greensand, leaving a large circular pit with a depressed +narrow border. On the outer side of this +process is a rounded boss, which may possibly have +supported the bone, if it were applied to the ground +with the wing folded up, like the wing of a Bat directed +upward and backward at the animal's side.</p> + +<p>The four bones of the wing finger usually decrease +<span class='pagenum'><a name="Page_133" id="Page_133">[Pg 133]</a></span> +progressively in length, so that in Rhamphorhynchus, +in which the length of the animal's head only slightly +exceeds 3½ inches, the first phalange is nearly as long, +the second phalange is about 3¼ inches, the third 2¾ +inches, and the fourth a little over 2 inches. Thus +the entire length of the four phalanges slightly exceeds +11 inches, or rather more than three times the +length of the head. But the fore-arm and metacarpus +in this type only measure 3 inches. Therefore the +entire spread of wings could not have been more +than 2 feet 9 inches.</p> + +<p>The largest Ornithosaur in which accurate measurements +have been made is the toothless Pterodactyle +Ornithostoma, also named Pteranodon, from North +America. In that type the head appears to have +been about 3 or 4 feet long, and the wing finger +exceeded 5 feet; while the length of the fore-arm +and metacarpus exceeded 3 feet. The width of the +body would not have been more than 1 foot. The +length of the short humerus, which was about +11 inches, did not add greatly to the stretch of the +wing; so that the spread of the wings as stretched +in flight may be given as probably not exceeding +17 or 18 feet. A fine example of the wing bones of +this animal quite as large has been obtained by the +(British Museum Natural History). Many years ago, +on very fragmentary materials, I estimated the wings +in the English Cretaceous Ornithocheirus as probably +having a stretch of 20 feet in the largest specimens, +basing the calculation partly upon the extent of the +longest wings in existing birds relatively to their +bones, and partly upon the size of the largest associated +bones which were then known.</p> + + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_134" id="Page_134">[Pg 134]</a></span></p> +<h2><a name="CHAPTER_XII" id="CHAPTER_XII"></a><small>CHAPTER XII</small><br /><br /> + +EVIDENCES OF THE ANIMAL'S +HABITS FROM ITS REMAINS</h2> + + +<p>Such are the more remarkable characters of the +bones in a type of animal life which was more +anomalous than any other which peopled the earth +in the Secondary Epoch of geological time. Its +skeleton in different parts resembles Reptiles, Birds, +and Mammals; with modifications and combinations +so singular that they might have been deemed impossible +if Nature's power of varying the skeleton +could be limited. Since Ornithosaurs were provided +with wings, we may believe the animals to some extent +to have resembled birds in habit. Their modes of +progression were more varied, for the structures indicate +an equal capacity for movement on land as a +biped, or as a quadruped, with movement in the air. +There is little evidence to support the idea that they +were usually aquatic animals. The majority of birds +which frequent the water have their bodies stored +with fat and the bones of their extremities filled with +marrow. And a bird's marrow bones are stouter and +stronger than those which are filled with air. There +are few, if any, bones of Pterodactyles so thick as to +suggest the conclusion that they contained marrow,<span class='pagenum'><a name="Page_135" id="Page_135">[Pg 135]</a></span> +and the bones of the extremities appear to have been +constructed on the lightest type found among terrestrial +birds. Their thinness, except in a few specimens +from the Wealden rocks, is marvellous; and all the +later Pterodactyles show the arrangement, as in birds, +by which air from the lungs is conveyed to the +principal bones. No Pterodactyle has shown any +trace of the web-footed condition seen in birds which +swim on the water, unless the diverging bones of the +hind foot in Rhamphorhynchus supports that inference. +The bones of the hind foot are relatively +small, and if it were not that a bird stands easily +upon one foot, might be considered scarcely adequate +to support the animal in the position which terrestrial +birds usually occupy. Yet, as compared with the +length and breadth of the foot in an Ostrich, the toes +of an Ornithosaur are seen to be ample for support. +These facts appear to discourage the idea that the +animals were equally at home on land and water, and +in air.</p> + +<p>Some light may be thrown upon the animal's habits +by the geological circumstances under which the +remains are found. The Pterodactyle named Dimorphodon, +from the Lias of the south of England, is +associated with evidences of terrestrial land animals, +the best known of which is Scelidosaurus, an armoured +Dinosaur adapted by its limbs for progression +on land. And the Pterodactyle Campylognathus, +from the Lias of Whitby, is associated with trunks +of coniferous trees and remains of Insects. So that +the occurrence of Pterodactyles in a marine stratum +is not inconsistent with their having been transported +by streams from off the old land surface of the Lias, +on which coniferous trees grew and Dinosaurs lived.<span class='pagenum'><a name="Page_136" id="Page_136">[Pg 136]</a></span></p> + +<p>Similar considerations apply to the occurrence of +the Rhamphocephalus in the Stonesfield Slate of +England. The deposit is not only formed in shallow +water, but contains terrestrial Insects, a variety of +land plants, and many Reptiles and other animals +which lived upon land. The specimens from the +Purbeck beds, again, are in strata which yield a +multitude of the spoils of a nearly adjacent land +surface; while the numerous remains found in the +marine Solenhofen Slate in Germany are similarly +associated with abundant evidences of varied types +of terrestrial life. The evidence grows in force from +its cumulative character. The Wealden beds, which +yield many terrestrial reptiles and so much evidence +of terrestrial vegetation, and shallow-water conditions +of disposition, have afforded important Pterodactyle +remains from the Isle of Wight and Sussex.</p> + +<p>The chief English deposit in which these fossils +are found, the Upper Greensand, has afforded +thousands of bones, battered and broken on a +shore, where they have lain in little associated +groups of remains, often becoming overgrown with +small marine shells. Side by side with them are +found bones of true terrestrial Lizards and Crocodiles +of the type of the Gavial of the Indian rivers, many +terrestrial Dinosaurs, and other evidences of land +life, including fossil resins, such as are met with in +the form of amber or copal at the present day.</p> + +<p>The great bones of Pterodactyles found in the +Chalk of Kent, near Rochester, became entombed, +beyond question, far from a land surface. There is +nothing to show whether the animals died on land +and were drifted out to sea like the timber which is +found water-logged and sunken after being drilled by<span class='pagenum'><a name="Page_137" id="Page_137">[Pg 137]</a></span> +the ship-worm (Teredo) of that epoch. Seeing the +power of flight which the animal possessed, storms +may have struck down travellers from time to time, +when far from land.</p> + +<p>Evidence of habit of another kind may be found +in their teeth. They are brightly enamelled, sharp, +formidable; and are frequently long, overlapping the +sides of the jaws. They are organs which are often +better adapted for grasping than for tearing, as may +be seen in the inclined teeth of Rhamphocephalus of +the Stonesfield Slate; and better adapted for killing +than tearing, from their piercing forms and cutting +edges, in genera like Ornithocheirus of the Greensand. +The manner in which the teeth were implanted and +carried is better paralleled by the fish-eating crocodile +of Indian rivers than by the flesh-eating crocodiles, or +Muggers, which live indifferently in rivers and the +sea. As the Kingfisher finds its food (see <a href="#Fig_20">Fig. 20</a>) +from the surface of the water without being in the +common sense of the term a water bird, so some +Pterodactyles may have fed on fish, for which their +teeth are well adapted, both in the stream and by the +shore.</p> + +<p>A Pterodactyle's teeth vary a good deal in appearance. +The few large teeth in the front of the jaw +in Dimorphodon, associated with the many small +vertical teeth placed further backward, suggest that +the taking of food may have been a process requiring +leisure, since the hinder teeth adapted to +mincing the animal's meat are extremely small. The +way in which the teeth are shaped and arranged +differs with the genera. In Pterodactylus they are +short and broad and few, placed for the most part +towards the front of the jaws. Their lancet-shaped<span class='pagenum'><a name="Page_138" id="Page_138">[Pg 138]</a></span> +form indicates a shear-like action adapted to dividing +flesh. In the associated genus Rhamphorhynchus +the teeth are absent from the extremity of the jaw, +are slender, pointed, spaced far apart, and extend far +backward. When the jaws of the Rhamphorhynchus +are brought together there is always a gap between +them in front, which has led to belief that the teeth +were replaced by some kind of horny armature which +has perished. In the long-nosed English type of +Ornithocheirus the jaws are compressed together, so +that the teeth of the opposite sides are parallel to +each other, with the margins well filled with teeth, +which are never in close contact, though occasionally +closer and larger in front, in some of the forms with +thick truncated snouts.</p> + +<p>It is not the least interesting circumstance of the +dentition of Pterodactyles that, associated in the +same deposits with these most recent genera with +teeth powerfully developed, there is a genus named +Ornithostoma from the resemblance of its mouth to +that of a bird in being entirely devoid of teeth. It +is scarcely possible to distinguish the remains of the +toothed and toothless skeletons except in the dentary +character of the jaws. There is no evidence that +the toothless types ever possessed a tooth of any +sort. They were first found in fragments in England +in the Cambridge Greensand, but were afterwards +met with in great abundance in the Chalk of Kansas, +where the same animals were named Pteranodon. +A jaw so entirely bird-like suggests that the digestive +organs of Pterodactyles may in such toothless forms +at least have been characterised by a gizzard, which +is so distinctive of Birds. The absence of teeth in +the Great Ant-eater and some other allied Mammals<span class='pagenum'><a name="Page_139" id="Page_139">[Pg 139]</a></span> +has transferred the function which teeth usually perform +to the stomach, one part of which becomes +greatly thickened and muscular, adapting itself to +the work which it has to perform. It is probable +that the gizzard may be developed in relation to the +necessities which food creates, since even Trout, feeding +on the shell-fish in some Irish lochs, acquire such +a thickened muscular stomach, and a like modification +is recorded in other fishes as produced by +food.</p> + +<p>Closely connected with an animal's habits is the +protection to the body which is afforded by the skin. +In Pterodactyles the evidence of the condition of +the skin is scanty, and mostly negative. Sometimes +the dense, smooth texture of the jaw bones indicates +a covering like the skin of a Lizard or the hinder part +of the jaw of a Bird. Some jaws from the Cambridge +Greensand have the bone channeled over its +surface by minute blood vessels which have impressed +themselves into the bone more easily than +into its covering. Thus in the species of Ornithocheirus +distinguished as <i>microdon</i> the palate is +absolutely smooth, while in the species named +<i>machærorhynchus</i> it is marked by parallel impressed +vascular grooves which diverge from the median +line. This condition clearly indicates a difference in +the covering of the bone, and that in the latter +species the covering had fewer blood vessels and +more horny protection than in the other. The tissue +may not have been of firmer consistence than in the +palate of Mammals. The extremity of the beak is +often as full of blood vessels as the jaw of a Turtle +or Crocodile.</p> +<p><span class='pagenum'><a name="Page_140" id="Page_140">[Pg 140]</a></span></p> + +<h4>COVERING OF THE BODY</h4> + +<p>There is no trace even in specimens from the +Solenhofen or Stonesfield Slate of any covering to +the body. There are no specimens preserved like +mummies, and although the substance of the wings +is found there is no trace of fur or feathers, bones, +or scales on the skin. The only example in which +there is even an appearance suggesting feathers is in +the beautiful Scaphognathus at Bonn, and upon portions +of the wing membrane of that specimen are +preserved a very few small short and apparently +tubular bodies, which have a suggestive resemblance +to the quills of small undeveloped feathers. Such +evidences have been diligently sought for. Professor +Marsh, after examining the wing membranes of his +specimen of Rhamphorhynchus from Solenhofen, +stated that the wings were partially folded and +naturally contracted into folds, and that the surface +of the tissue is marked by delicate striæ, which +might easily be taken at first sight for a thin coating +of hair. Closer investigation proved the markings +to be minute wrinkles on the under surface of the +wing membrane. This negative evidence has considerable +value, because the Solenhofen Slate has +preserved in the two known examples of the bird +Archæopteryx beautiful details of the structure of +the larger feathers concerned in flight. It has preserved +many structures far more delicate. There is, +therefore, reason for believing that if the skin had +possessed any covering like one of those found in +existing vertebrate animals, it could scarcely have +escaped detection in the numerous undisturbed skeletons +of Pterodactyles which have been examined.<span class='pagenum'><a name="Page_141" id="Page_141">[Pg 141]</a></span></p> + +<p>The absence of a recognisable covering to the skin +in a fossil state cannot be accepted as conclusive +evidence of the temperature, habits, or affinities of +the animal. Although Mammalia are almost entirely +clothed with dense hair, which has never been +found in a recognisable condition in a fossil state in +any specimen of Tertiary age, one entire order, the +Cetacea, show in the smooth hairless skins of Whales +and Porpoises that the class may part with the +typical characteristic covering without loss of temperature +and without intelligible cause. That the absence +of hair is not due to the aquatic conditions of rivers +or sea is proved by other marine Mammals, like Seals, +having the skin clothed with a dense growth of hair, +which is not surpassed in any other order. The fineness +of the growth of hair in Man gives a superficial +appearance of the skin being imperfectly clothed, +and a similar skin in a fossil state might give the +impression that it was devoid of hair. There are +many Mammals in which the skin is scantily clothed +with hair as the animal grows old. Neither the +Elephant nor the Armadillo in a fossil state would be +likely to have the hair preserved, for the growth is +thin on the bony shields of the living Armadilloes. +Yet the difficulty need be no more inherent in the +nature of hair than in that of feathers, since the hair +of the Mammoth and Rhinoceros has been completely +preserved upon their skins in the tundras of +Siberia, densely clothing the body. This may go +to show that the Pterodactyle possessed a thin +covering of hair, or, more probably, that hair +was absent. Since Reptiles are equally variable in +the clothing of the skin with bony or horny plates, +and in sometimes having no such protection, it may<span class='pagenum'><a name="Page_142" id="Page_142">[Pg 142]</a></span> +not appear singular that the skin in Ornithosaurs has +hitherto given no evidence of a covering. From +analogy a covering might have been expected; +feathers of Birds and hair of Mammals are non-conducting +coverings suited to arrest the loss of heat.</p> + +<p>With the evidence, such as it is, of resemblance of +Ornithosaurs to Birds in some features of respiration +and flight, a covering to the skin might have been +expected. Yet the covering may not be necessary +to a high temperature of the blood. Since Dr. John +Davy made his observations it has been known that +the temperature of the Tunny, above 90° Fahrenheit, +is as warm as the African scaly ant-eater named the +Pangolin, which has the body more amply protected +by its covering. This illustration also shows that +hot blood may be produced without a four-celled +heart, with which it is usually associated, and that +even if the skin in Pterodactyles was absolutely +naked an active life and an abundant supply of blood +could have given the animal a high temperature.</p> + +<p>The circumstance that in several individuals the +substance of the wing membrane is preserved would +appear to indicate either that it was exceptionally +stout when there would have been small chance of +resisting decomposition, or that its preservation is +due to a covering which once existed of fur or down +or other clothing substance, which has proved more +durable than the skin itself.</p> + +<div class="figcenter" style="width: 556px;"> +<a name="Fig_48" id="Fig_48"></a> +<span class="caption">FIG. 48. REMAINS OF DIMORPHODON FROM THE LIAS OF LYME REGIS<br /><br /> +SHOWING THE SKULL, NECK, BACK AND SOME OF THE LONGER BONES OF THE SKELETON</span> +<img src="images/i_161.jpg" width="556" height="1024" alt="FIG. 48." title="" /> +<p class="center"><i>From a slab in the British Museum (Natural History)</i></p> +</div> + + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_143" id="Page_143">[Pg 143]</a></span></p> +<h2><a name="CHAPTER_XIII" id="CHAPTER_XIII"></a><small>CHAPTER XIII</small><br /><br /> + +ANCIENT ORNITHOSAURS FROM THE LIAS</h2> + + +<p>Cuvier's discourse on the revolutions of the +Earth made the Pterodactyle known to English +readers early in the nineteenth century. Dr. Buckland, +the distinguished professor of Geology at +Oxford, discovered in 1829 a far larger specimen in +the Lias of Lyme Regis, and it became known by a +figure published by the Geological Society, and by the +description in his famous Bridgewater Treatise, p. 164. +This animal was tantalising in imperfect preservation. +The bones were scattered in the clay, so as to give no +idea of the animal's aspect. Knowledge of its limbs +and body has been gradually acquired; and now, for +some years, the tail and most parts of the skeleton +have been well known in this oldest and most +interesting British Pterodactyle.</p> + +<p>Sir Richard Owen after some time separated the +fossil as a distinct genus, named Dimorphodon; for +it was in many ways unlike the Pterodactyles described +from Bavaria. The name Dimorphodon indicated +the two distinct kinds of teeth in the jaws, +a character which is still unparalleled among Pterodactyles +of newer age. There are a few large pointed, +<span class='pagenum'><a name="Page_144" id="Page_144">[Pg 144]</a></span> +piercing and tearing teeth in the front of the jaws, +with smaller teeth further back, placed among the +tearing teeth in the upper jaw; while in the lower +jaw the small teeth are continuous, close-set, and +form a fine cutting edge like a saw.</p> + +<div class="figcenter" style="width: 1024px;"> +<a name="Fig_49" id="Fig_49"></a> +<span class="caption">FIG. 49. LEFT SIDE OF DIMORPHODON (RESTORED) AT REST</span> +<img src="images/i_163.jpg" width="1024" height="470" alt="FIG. 49." title="FIG. 49." /> +</div> + +<p>The Dimorphodon has a short beak, a deep head, +and deep lower jaw, which is overlapped by the cheek +bones. The side of the head is occupied by four +vacuities, separated by narrow bars of bone. First, in +front, is the immense opening for the nostril, triangular +in form, with the long upper side following the rounded +curve of the face. A large triangular opening intervenes +between the nose hole and the eye hole, scarcely +smaller than the former, but much larger than the +orbit of the eye. The eye hole is shaped like a kite +or inverted pear. Further back still is a narrower vertical +opening known as the lateral or inferior temporal +vacuity. The back of the head is badly preserved. +The two principal skulls differ in depth, probably +from the strains under which they were pressed flat +in the clay. A singular detail of structure is found +in the extremity of the lower jaw, which is turned +slightly downward, and terminates in a short toothless +<span class='pagenum'><a name="Page_145" id="Page_145">[Pg 145]</a></span> +point. The head of Dimorphodon is about eight inches +long.</p> + +<div class="figcenter" style="width: 1024px;"> +<a name="Fig_50" id="Fig_50"></a> +<span class="caption">FIG. 50. DIMORPHODON MACRONYX<br /><br /> +RESTORED FORM OF THE ANIMAL</span> +<img src="images/i_165.jpg" width="1024" height="523" alt="FIG. 50." title="FIG. 50." /> +</div> + +<p>The neck bones are of suitable stoutness and width +to support the head. The bones are yoked together +by strong processes. The neck was about 6 inches +long, did not include more than seven bones, and +appeared short owing only to the depth and size of +the head. The length of the backbone which supported +the ribs was also about 6 inches. Its joints +are remarkably short when compared with those of +the neck. The tail is about 20 inches long.</p> + +<p>The extreme length of the animal from the tip +of the nose to the end of the tail may have been +3 feet 4 inches, supposing it to have walked on all +fours in the manner of a Reptile or Mammal. This +may have been a common position, but Dimorphodon +may probably also have been a biped. Before +1875, when the first restoration appeared in the +<i>Illustrated London News</i>, the legs had been regarded +as too short to have supported the animal, standing +upon its hind limbs. They are here seen to be well +adapted for such a purpose. The upper leg bone is +3¼ inches long, the lower leg bone is 4½ inches +long, and the singularly strong instep bones are +firmly packed together side by side as in a leaping +or jumping Mammal, and measure 1½ inches +in length. Dimorphodon differs from several other +Pterodactyles in having the hind limb provided +with a fifth outermost short instep bone, to which +two toe bones are attached. These bones are elongated +in a way that may be compared, on a small +scale, with the elongation of the wing finger in the +fore limb. The digit was manifestly used in the same +way as the wing finger, in partial support of a flying +<span class='pagenum'><a name="Page_146" id="Page_146">[Pg 146]</a></span> +membrane, though its direction may have been upward +and outward, rather than inward. There is no +evidence of a pulley joint between the metatarsal +and the adjacent phalange.</p> + +<p>The height of the Dimorphodon, standing on its +hind legs in the position of a Bird, with the wings +folded upon the body in the manner of a Bird, was +about 20 inches. An ungainly, ill-balanced animal +in aspect, but not more so than many big-headed +birds, and probably capable of resting upon the instep +bones as many birds do. The chief point of variation +from the Pterodactyle wing is in the relative +length of the metacarpus in Dimorphodon. It is +shorter than the other bones in the wing, never +exceeding 1½ inches. The total length of all +the arm bones down to the point where the metacarpus +might have touched the ground, or where +the wing finger is bent upon it, is about 9 inches, +which gives a length of less than 6 inches below +the upper arm bone. The four bones of the wing +finger measure, from the point where the first bone +bends upon the metacarpus, less than 18 inches. +So that the wings could only have been carried +in the manner of the wings of a Bat, folded at the +side and directed obliquely over the back when the +animal moved on all fours. Its body would appear +to have been raised high above the ground, in a +manner almost unparalleled in Reptiles, and comparable +to Birds and Mammals. Dimorphodon is to +be imagined in full flight, with the body extended +like that of a Bird, when the wings would have +had a spread from side to side of about 4 feet 4 +inches. As in other animals of this group, the three +claws on the front feet are larger than the similar +four claws on the hind feet; as though the fingers +might have functions in grasping prey, which were +not shared by the toes.</p> + +<div class="figcenter" style="width: 1024px;"> +<a name="Fig_51" id="Fig_51"></a> +<span class="caption">FIG. 51. DIMORPHODON MACRONYX WALKING AS A QUADRUPED<br /><br /> +RESTORATION OF THE SKELETON</span> +<img src="images/i_168.jpg" width="1024" height="538" alt="FIG. 51." title="FIG. 51." /> +</div> + +<p><span class='pagenum'><a name="Page_147" id="Page_147">[Pg 147]</a></span>The +restorations give faithful pictures of the skeleton, +and the form of the body is built upon the indications +of muscular structure seen in the bones.</p> + + +<div class="figcenter" style="width: 1024px;"> +<a name="Fig_52" id="Fig_52"></a> +<span class="caption">FIG. 52. DIMORPHODON MACRONYX WALKING AS A BIPED</span> +<p class="center"><i>Based chiefly on remains in the British Museum</i></p> +<img src="images/i_172.jpg" width="1024" height="650" alt="FIG. 52." title="FIG. 52." /> +</div> + +<p>A second English Pterodactyle is found in the +Upper Lias of Whitby. It is only known from +an imperfect skull, published in 1888. It has the +great advantage of preserving the bones in their +natural relations to each other, and with a length of +head probably similar to Dimorphodon shows that +the depth at the back of the eye was much less; and +the skull wants the arched contour of face seen in +Dimorphodon. The head has the same four lateral +vacuities, but the nostril is relatively small and elongated, +extending partly above the oval antorbital +opening, which was larger. There is thus a difference +of proportion, but it is precisely such as might result +from the species having the skull flatter. The head +is easily distinguished by the small nostril, which +is smaller than the orbit of the eye. The animal +is referred to another genus. The quadrate bones +which give attachment to the lower jaw send a process +inward to meet the bones of the palate, which differ +somewhat from the usual condition. Two bony rods +extend from the quadrate bones backward and upward +to the sphenoid, and two more slender bones +extend from the quadrate bones forward, and converge +in a <b>V</b>-shape, to define the division between +the openings of the nostrils on the palate. The +<b>V</b>-shaped bone in front is called the vomer, while +the hinder part is called pterygoid. The bones that +extend backward to the sphenoid are not easily identified. +<span class='pagenum'><a name="Page_148" id="Page_148">[Pg 148]</a></span> +This animal is one of the most interesting +of Pterodactyles from the very reptilian character +exhibited in the back of the head, which appears to +be different from other specimens, which are more +like a bird in that region. Yet underneath this +reptilian aspect, with the bony bar at the side of the +temporal region of the head formed by the squamosal +and quadrate bones, defining the two temporal vacuities +as in Reptiles, a mould is preserved of the +cavity once occupied by the brain, showing the chief +details of structure of that organ, and proving that +in so far as it departs from the brain of a Bird it +appears to resemble the brain of a Mammal, and is +unlike the brain of a Reptile.</p> + +<p>The Pterodactyles from the Lias of Germany are +similar to the English types, in so far as they can be +compared. In 1878 I had the opportunity of studying +those which were preserved in the Castle at Banz, +which Professor Andreas Wagner, in 1860, referred to +the new genus Dorygnathus. The skull is unknown, +but the lower jaw, 6½ inches long, is less than 2½ inches +wide at the articulation with the quadrate bone in the +skull. The depth of the lower jaw does not exceed +¼ inch, so that it is in marked contrast to Buckland's +Dimorphodon. The symphysis, which completely +blends the rami of the jaw, is short. As far as it +extends it contains large tearing teeth, followed by +smaller teeth behind, like those of Dimorphodon. +But this German fossil appears to differ from the +English type in having the front of the lower jaw, for +about ¾ inch, compressed from side to side into a +sharp blade or spear, more marked than in any +other Pterodactyle, and directed <i>upward</i> instead of +downward as in Dimorphodon. Nearly all the +<span class='pagenum'><a name="Page_149" id="Page_149">[Pg 149]</a></span> +measurements in the skeleton are practically identical +with those of the English Dimorphodon, and +extend to the jaw, humerus, ulna and radius, wing +metacarpal, first phalange of the wing finger. The +principal bones of the hind limb appear to be a little +shorter; but the scapula and coracoid are slightly +larger. All these bones are so similar in form to +Dimorphodon that they could not be separated from +the Lyme Regis species, if they were found in the +same locality.</p> + + +<div class="figcenter" style="width: 558px;"> +<a name="Fig_53" id="Fig_53"></a> +<span class="caption">FIG. 53. LOWER JAW OF DORYGNATHUS SEEN FROM BELOW</span> +<p class="center">From the Lower Lias of Germany, showing the spear in front +of the tooth sockets</p> +<img src="images/i_174.jpg" width="558" height="480" alt="FIG. 53." title="FIG. 53." /> +</div> + +<p>Just as the Upper Lias in England has yielded a +second Pterodactyle, so the Upper Lias in Germany +has yielded a skeleton, to which Felix Plieninger, in +1894, gave the name Campylognathus. It is an instructive +skeleton, with the head much smaller than +in Dimorphodon, being less than 6 inches long, but, +unfortunately, broken and disturbed. A lower jaw +<span class='pagenum'><a name="Page_150" id="Page_150">[Pg 150]</a></span> +gives the length 4½ inches. Like the other Pterodactyles +from the Lias, it has the extremity of the +beak toothless, with larger teeth in the region of the +symphysis in front and smaller teeth behind. The jaw +is deeper than in the Banz specimen from the Lower +Lias, but not so deep as in Dimorphodon. The teeth +of the upper jaw vary in size, and there appears to +be an exceptionally large tooth in the position of +the Mammalian canine at the junction of the bones +named maxillary and intermaxillary.</p> + +<p>The nasal opening is small and elongated, as in the +English specimen from Whitby. As in that type +there is little or no indication of the convex contour +of the face seen in Dimorphodon.</p> + +<p>The neck does not appear to be preserved. In the +back the vertebræ are about <sup>3</sup>/<sub>10</sub> inch long, so that +twelve, which is the usual number, would only occupy +a length of a little more than 3½ inches. The tail is +elongated like that of Dimorphodon, and bordered +in the same way by ossified ligaments. There are +thirty-five tail vertebræ. Those which immediately +follow the pelvis are short, like the vertebræ of the +back. But they soon elongate, and reach a maximum +length of nearly 1½ inches at the eighth, and +then gradually diminish till the last scarcely exceeds +<sup>1</sup>/<sub>8</sub> inch in length. The length of the tail is +about 22 inches; this appears to be an inch or +two longer than in Dimorphodon. The longest rib +measures 2½ inches, and the shortest 2 inches. These +ribs probably were connected with the sternum, which +is imperfectly preserved.</p> + + +<div class="figcenter" style="width: 1024px;"> +<a name="Fig_54" id="Fig_54"></a> +<span class="caption">FIG. 54. DIMORPHODON MACRONYX<br /><br /> +SHOWING THE MAXIMUM SPREAD OF THE WING MEMBRANES</span> +<img src="images/i_176.jpg" width="1024" height="588" alt="FIG. 54." title="FIG. 54." /> +</div> + +<p>The bones of the limbs have about the same length +as those of Dimorphodon, so far as they can be compared, +except that the ulna and radius are shorter. +<span class='pagenum'><a name="Page_151" id="Page_151">[Pg 151]</a></span> +The wing metacarpal is of about the same length, +but the first phalange of the wing finger measures +6¼ inches, the second is about 8¼ inches, the third +6½ inches, and the fourth 4¾ inches; so that the total +length of the wing finger was about half an inch short +of 2 feet. One character especially deserves attention +in the apparent successive elongation of the first three +phalanges in the wing finger in Dimorphodon. The +third phalange is the longest in the only specimen in +which the finger bones are all preserved. Usually the +first phalange is much longer than the second, so that +it is a further point of interest to find that this German +type shares with Dimorphodon a character of +the wing finger which distinguishes both from some +members of the group by its short first phalange.</p> + + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_55" id="Fig_55"></a> +<span class="caption">FIG. 55. THE LEFT SIDE OF THE PELVIS OF DIMORPHODON +SHOWING THE TWO PREPUBIC BONES</span> +<img src="images/i_178.jpg" width="640" height="398" alt="FIG. 55." title="FIG. 55." /> +</div> + +<p>The pelvis is exceptionally strong in Campylognathus, +and although it is crushed the bones manifestly +met at the base of the ischium, while the pubic +bones were separated from each other in front. The +bones of the hind limb are altogether shorter in the +German fossil than in Dimorphodon, especially in +<span class='pagenum'><a name="Page_152" id="Page_152">[Pg 152]</a></span> +the tibia; but the structure of the metatarsus is just +the same, even to the short fifth metatarsal with its +two digits, only those bones are extremely short, instead +of being elongated as in Dimorphodon. It is +therefore convenient, from the different proportions +of the body, that Campylognathus may be separated +from Dimorphodon; but so much as is preserved of +the English specimen from the Upper Lias of Whitby +rather favours the belief that our species should also +be referred to Campylognathus, which had not been +figured when the Whitby skull was referred to Scaphognathus +by Mr. Newton. It may be doubtful whether +there is sufficient evidence to establish the distinctness +of the other German genus Dorygnathus, though +it may be retained pending further knowledge.</p> + +<p>In these characters are grounds for placing the +Lias Pterodactyles in a distinct family, the Dimorphodontidæ, +as was suggested in 1870. This evidence +is found in the five metatarsal bones, of which four +are in close contact, the middle two being slightly +the longest, so as to present the general aspect of the +corresponding bones in a Mammal rather than a Bird. +Secondly, the very slender fibula, prolonged down the +length of the shin bone, which ends in a rounded +pulley like the corresponding bone of a Bird. Thirdly, +the great elongation of the third wing phalange. +Fourthly, the prolongation of the coracoid bone beyond +the articulation for the humerus, as in a Bird. +And the toothless, spear-shaped beak, and jaw with +large teeth in front and small teeth behind, are also +distinctive characters.</p> + + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_153" id="Page_153">[Pg 153]</a></span></p> +<h2><a name="CHAPTER_XIV" id="CHAPTER_XIV"></a><small>CHAPTER XIV</small><br /><br /> + +ORNITHOSAURS FROM THE +MIDDLE SECONDARY ROCKS</h2> + + +<h4>RHAMPHOCEPHALUS</h4> + +<p>The Stonesfield Slate in England, which corresponds +in age with the lower part of the Great +or Bath Oolite, yields many evidences of terrestrial +life—land plants, insects, and mammals—preserved in +a marine deposit. A number of isolated bones have +been found of Pterodactyles, some of them indicating +animals of considerable size and strength. The +nature of the limestone was unfavourable to the preservation +of soft wing membranes, or even to the bones +remaining in natural association. Very little is known +of the head of Rhamphocephalus. One imperfect specimen +shows a long temporal region which is wide, and +a very narrow interspace between the orbits; with a +long face, indicated by the extension of narrow +nasal bones. The lower jaw has an edentulous beak +or spear in front, which is compressed from side to +side in the manner of the Liassic forms, but turned +upward slightly, as in Dorygnathus or Campylognathus. +Behind this extremity are sharp, tall teeth, +few in number, which somewhat diminish in size as +they extend backward, and do not suddenly change +<span class='pagenum'><a name="Page_154" id="Page_154">[Pg 154]</a></span> +to smaller series, as in the Lias genera. A few small +vertebræ have been found, indicating the neck and +back. The sacrum consists of five vertebræ. One +small example has a length of only an inch. It is +a little narrower behind than in front, and would be +consistent with the animal having had a long tail, +which I believe to have been present, although I +have not seen any caudal vertebræ. The early ribs +are like the early ribs of a Crocodile or Bird in the +well-marked double articulation. The later ribs +appear to have but one head. <b>V</b>-shaped abdominal +ribs are preserved. Much of the animal is unknown. +The coracoid seems to have been directed forward, +and, as in a bird, it is 2½ inches long. The humerus +is 3½ inches long, and the fore-arm measured 6 +inches, so that it was relatively longer than in Dimorphodon. +The metacarpus is 1¾ inches long. The +wing finger was exceptionally long and strong. Professor +Huxley gave its length at 29 inches. My own +studies lead to the conclusion that the first finger +bone of the wing was the shorter, and that although +they did not differ greatly in length, the second was +probably the longest, as in Campylognathus.</p> + +<p>Professor Huxley makes the second and third +phalanges 7¾ inches long, and the first only about +<sup>3</sup>/<sub>8</sub> inch shorter, while the fourth phalange is 6½ inches. +These measurements are based upon some specimens +in the Oxford University Museum. There is only +one first phalange which has a length of 7¾ inches. +The others are between 5 and 6 inches, or but little +exceed 4 inches; so that as all the fourth phalanges +which are known have a length of 6½ inches, it is +possible that the normal length of the first phalange +in the larger species was 5½ inches. The largest +<span class='pagenum'><a name="Page_155" id="Page_155">[Pg 155]</a></span> +of the phalanges which may be classed as second or +third is 8½ inches, and that, I suppose, may have been +associated with the 7¾ inches first phalange. But +the other bones which could have had this position +all measure 5½ and 7¾ inches. The three species +indicated by finger bones may have had the measurements:—</p> + +<div class='center'> +<table border="0" cellpadding="2" cellspacing="0" summary=""> +<tr><th align='center' colspan='4'>Phalanges of the wing finger</th></tr> +<tr><td align='center'>I.</td><td align='center'>II.</td><td align='center'>III.</td><td align='center'>IV.</td></tr> +<tr><td align='center'>7¾</td><td align='center'>8½</td><td align='center'>[7?]</td><td align='center'>6½</td><td align='center' rowspan='3'><span class="ft30">}</span> length of each bone in inches.</td></tr> +<tr><td align='center'>5½</td><td align='center'>7¾</td><td align='center'>5½</td><td align='center'>[4½?]</td></tr> +<tr><td align='center'>4½</td><td align='center'>——</td><td align='center'>——</td><td align='center'>——</td></tr> +</table></div> + +<p>The femur is represented by many examples—one +3¾ inches long, and others less than 3 inches long +(2<sup>9</sup>/<sub>10</sub>). In Campylognathus, which has so much in +common with the jaw and the wing bones in size, +the upper leg bone is 2<sup>8</sup>/<sub>10</sub> inches. Therefore if we +assign the larger femur to the larger wing, the femur +will be relatively longer in all species of Rhamphocephalus +than in Campylognathus. Only one example +of a tibia is preserved. It is 3½ inches long, +or only <sup>1</sup>/<sub>10</sub> inch shorter than the bone in Campylognathus, +which has the femur 2<sup>8</sup>/<sub>10</sub> inches, so that I +refer the tibia of Rhamphocephalus to the species +which has the intermediate length of wing. These +coincidences with Campylognathus establish a close +affinity, and may raise the question whether the +Upper Lias species may not be included in the +Stonesfield Slate genus Rhamphocephalus.</p> + +<p>The late Professor Phillips, in his <i>Geology of Oxford</i>, +attempted a restoration of the Stonesfield Ornithosaur, +and produced a picturesque effect (<a href="#Page_164">p. 164</a>); but +no restoration is possible without such attention to +the proportions of the bones as we have indicated.</p> +<p><span class='pagenum'><a name="Page_156" id="Page_156">[Pg 156]</a></span></p> + +<h4>OXFORD CLAY</h4> + +<p>A few bones of flying reptiles have been found in +the Lower Oxford Clay near Peterborough, and others +in the Upper Oxford Clay at St. Ives, in Huntingdonshire. +A single tail vertebra from the Middle Oxford +Clay, near Oxford, long since came under my own +notice, and shows that these animals belong to a +long-tailed type like Campylognathus. The cervical +vertebræ are remarkable for being scarcely longer than +the dorsal vertebræ; and the dorsal are at least half +as long again as is usual, having rather the proportion +of bones in the back of a crocodile.</p> + + +<h4>LITHOGRAPHIC SLATE</h4> + +<p>Long-tailed Pterodactyles are beautifully preserved +in the Lithographic Limestone of the south of Bavaria, +at Solenhofen, and the quarries in its neighbourhood, +often with the skeleton or a large part of it flattened +out in the plane of bedding of the rock. Fine skeletons +are preserved in the superb museum at Munich, +at Heidelberg, Bonn, Haarlem, and London, and are +all referred to the genus Rhamphorhynchus or to +Scaphognathus. It is a type with powerfully developed +wings and a long, stiff tail, very similar to +that of Dimorphodon, so that some naturalists refer +both to the same family. There is some resemblance.</p> + +<p>The type which is most like Dimorphodon is the +celebrated fossil at Bonn, sometimes called <i>Pterodactylus +crassirostris</i>, which in a restored form, with a +short tail, has been reproduced in many text-books. +No tail is preserved in the slab, and I ventured to +give the animal a tail for the first time in a restoration +(<a href="#Page_163">p. 163</a>) published by the <i>Illustrated London News</i> +in 1875, which accompanied a report of a Royal +<span class='pagenum'><a name="Page_157" id="Page_157">[Pg 157]</a></span> +Institution lecture. Afterwards, in 1882, Professor +Zittel, of Munich, published the same conclusion. +The reason for restoring the tail was that the animal +had the head constructed in the same way as +Pterodactyles with a long tail, and showed differences +from types in which the tail is short; and there +is no known short-tailed Pterodactyle, with wrist +and hand bones, such as characterise this animal. +The side of the face has a general resemblance to +the Pterodactyles from the Lias, for although the +framework is firmer, the four apertures in the head +are similarly placed. The nostril is rather small and +elongated, and ascends over the larger antorbital +vacuity. The orbit for the eye is the largest opening +in the head, so that these three apertures successively +increase in size, and are followed by the vertically +elongated post-orbital vacuity. The teeth are widely +spaced apart, and those in the skull extend some +distance backward to the end of the maxillary bone. +There are few teeth in the lower jaw, and they correspond +to the large anterior teeth of Dimorphodon, +there being no teeth behind the nasal opening. The +lower jaw is straight, and the extremities of the +jaws met when the mouth was closed. The breast +bone does not show the keel which is so remarkable +in Rhamphorhynchus, which may be attributed to +its under side being exposed, so as to exhibit the +pneumatic foramina.</p> + +<p>The ribs have double heads, more like those of a +Crocodile in the region of the back than is the case +with the bird-like ribs from Stonesfield. The second +joint in the wing finger may be longer than the first—a +character which would tend to the association of +this Pterodactyle with species from the Lias; a relation +<span class='pagenum'><a name="Page_158" id="Page_158">[Pg 158]</a></span> +to which attention was first drawn by Mr. E. T. Newton, +who described the Whitby skull.</p> + +<p>The Pterodactyles from the Solenhofen Slate which +possess long tails have a series of characters which show +affinity with the other long-tailed types. The jaws are +much more slender. The orbit of the eye in Rhamphorhynchus +is enormously large, and placed vertically +above the articulation for the lower jaw. Immediately +in front of the eye are two small and elongated openings, +the hinder of which, known as the antorbital +vacuity, is often slightly smaller than the nostril, which +is placed in the middle length of the head, or a little +further back, giving a long dagger-shaped jaw, which +terminates in a toothless spear. The lower jaw has +a corresponding sharp extremity. The teeth are +directed forward in a way that is quite exceptional. +Notwithstanding the massiveness and elongation of +the neck vertebræ, which are nearly twice as long as +those of the back, the neck is sometimes only about +half the length of the skull.</p> + +<p>All these long-tailed species from the Lithographic +Stone agree in having the sternum broad, with a long +strong keel, extending far forward. The coracoid +bones extend outward like those of a Crocodile, so +as to widen the chest cavity instead of being carried +forward as the bones are in Birds. These bones in this +animal were attached to the anterior extremity of the +sternum, so that the keel extended in advance of the +articulation as in other Pterodactyles. The breadth +of the sternum shows that, as in Mammals, the fore +part of the body must have been fully twice the +width of the region of the hip-girdle, where the +slenderer hind limbs were attached. The length +of the fore limb was enormous, for although the head +<span class='pagenum'><a name="Page_159" id="Page_159">[Pg 159]</a></span> +suggests an immense length relatively to the body, +nearly equal to neck and back together, the head is +not more than a third of the length of the wing +bones. The wing bones are remarkable for the short +powerful humerus with an expanded radial crest, +which is fully equal in width to half the length of the +bone. Another character is the extreme shortness +of the metacarpus, usually associated with immense +strength of the wing metacarpal bone.</p> + +<p>The hind limbs are relatively small and relatively +short. The femur is usually shorter than the humerus, +and the tibia is much shorter than the ulna. The +bones of the instep, instead of being held together +firmly as in the Lias genera, diverge from each other, +widening out, though it often happens that four of +the five metatarsals differ but little in length. The +fifth digit is always shorter.</p> + +<p>The hip-girdle of bones differs chiefly from other +types in the way in which those bones, which have +sometimes been likened to the marsupial bones, are +conditioned. They may be a pair of triangular bones +which meet in the middle line, so that there is an +outer angle like the arm of a capital Y. Sometimes +these triangular bones are blended into a curved, +bow-shaped arch, which in several specimens appears +to extend forward from near the place of articulation +of the femur. This is seen in fossil skeletons at +Heidelberg and Munich. It is possible that this +position is an accident of preservation, and that the +prepubic bones are really attached to the lower +border of the pubic bones.</p> + +<p>Immense as the length of the tail appears to be, +exceeding the skull and remainder of the vertebral +column, it falls far short of the combined length of the +<span class='pagenum'><a name="Page_160" id="Page_160">[Pg 160]</a></span> +phalanges of the wing finger. The power of flight +was manifestly greater in Rhamphorhynchus than in +other members of the group, and all the modifications +of the skeleton tend towards adaptation of the +animals for flying. The most remarkable modification +of structure at the extremity of the tail was made +known by Professor Marsh in a vertical, leaf-like +expansion in this genus, which had not previously +been observed (<a href="#Page_161">p. 161</a>). The vertebræ go on steadily +diminishing in length in the usual way, and then +the ossified structures which bordered the tail bones +and run parallel with the vertebræ in all the Rhamphorhynchus +family, suddenly diverge downward and +upward at right angles to the vertebræ, forming a +vertical crest above and a corresponding keel below; +and between these structures, which are identified +with the neural spines and chevron bones of ordinary +vertebræ, the membrane extends, giving the extremity +of the tail a rudder-like feature, which, from knowledge +of the construction of the tail of a child's kite, +may well be thought to have had influence in directing +and steadying the animal's movements. There +are many minor features in the shoulder-girdle, which +show that the coracoid, for example, was becoming +unlike that bone in the Lias, though it still continues +<span class='pagenum'><a name="Page_161" id="Page_161">[Pg 161]</a></span> +to have a bony union with the elongated shoulder-blade +of the back.</p> + + +<p><span class='pagenum'><a name="Page_162" id="Page_162">[Pg 162]</a></span></p> + +<div class="figcenter" style="width: 1024px;"> +<a name="Fig_56" id="Fig_56"></a> +<span class="caption">FIG. 56. RESTORATION OF THE SKELETON OF <i>RHAMPHORHYNCHUS PHYLLURUS</i></span> +<p class="center">From the Solenhofen Slate, partly based upon the skeleton with the wing membranes preserved</p> +<img src="images/i_188.jpg" width="1024" height="630" alt="FIG. 56." title="FIG. 56." /> +</div> + + + +<p><span class='pagenum'><a name="Page_163" id="Page_163">[Pg 163]</a></span></p> +<div class="figcenter" style="width: 640px;"> +<a name="Fig_57" id="Fig_57"></a> +<span class="caption">FIG. 57. RESTORATION OF THE SKELETON OF +<i>SCAPHOGNATHUS CRASSIROSTRIS</i></span> +<p class="center">Published in the <i>Illustrated London News</i> in 1875. In which a tail is +shown on the evidence of the structure of the head and hand</p> +<img src="images/i_190.jpg" width="640" height="408" alt="FIG. 57." title="FIG. 57." /> +</div> + + + + +<p>The great German delineator of these animals, Von +Meyer, admitted six different species. Mr. Newton +and Mr. Lydekker diminish the number to four. It is +not easy to determine these differences, or to say how +far the differences observed in the bones characterise +species or genera. It is certain that there is one +remarkable difference from other and older Pterodactyles, +in that the last or fourth bone in the wing +finger is usually slightly longer than the third bone, +which precedes it. There is a certain variability in +the specimens which makes discussion of their +characters difficult, and has led to some forms being +regarded as varieties, while others, of which less +material is available, are classed as species. I am +disposed to say that some of the confusion may +have resulted from specimens being wrongly named. +Thus, there is a Rhamphorhynchus called curtimanus, +or the form with the short hand. It is +represented by two types. One of these appears to +have the humerus short, the ulna and radius long, +and the finger bones long; the other has the humerus +longer, the ulna much shorter, and the finger bones +shorter. They are clearly different species, but the +second variety agrees in almost every detail with +a species named hirundinaceus, the swallow-like +Rhamphorhynchus. This identification shows, not +<span class='pagenum'><a name="Page_164" id="Page_164">[Pg 164]</a></span> +that the latter is a bad species, but that curtimanus +is a distinct species which had sometimes been confounded +with the other. While most of these +specimens show a small but steady decrease in the +length of the several wing finger bones, the species +called Gemmingi has the first three bones absolutely +equal and shorter than in the species curtimanus, +longimanus, or hirundinaceus. In the same way, +on the evidence of facts, I find myself unable to join +in discarding Professor Marsh's species phyllurus, +on account of the different proportions of its limb +bones. The humerus, metacarpus, and third phalange +<span class='pagenum'><a name="Page_165" id="Page_165">[Pg 165]</a></span> +of the wing finger in <i>Rhamphorhynchus phyllurus</i> +are exceptionally short as compared with other +species. Everyone agrees that the species called +longicaudus is a distinct one, so that it is chiefly in +slight differences in the proportions of constituent +parts of the skeleton that the types of the Rhamphorhynchus +are distinguished from each other. +I cannot quite concur with either Professor Zittel +(<a href="#Fig_58">Fig. 58, 3</a>) or Professor Marsh (<a href="#Fig_58">Fig. 58, 2</a>) in the +expansion which they give to the wing membrane +in their restorations; for although Professor Zittel +represents the tail as free from the hind legs, while +Professor Marsh connects them together, they both +concur in carrying the wing membrane from the +tip of the wing finger down to the extremity of the +ankle joint. I should have preferred to carry it no +further down the body than the lower part of the +back, there being no fossil evidence in favour of this +extension so far as specimens have been described. +Neither the membranous wings figured by Zittel nor +by Marsh would warrant so much body membrane as +the Rhamphorhynchus has been credited with. I +have based my restoration (<a href="#Page_161">p. 161</a>) of the skeleton +chiefly on <i>Rhamphorhynchus phyllurus.</i></p> + + +<div class="figcenter" style="width: 600px;"> +<a name="Fig_58" id="Fig_58"></a> +<span class="caption">FIG. 58. SIX RESTORATIONS</span> +<img src="images/i_191.jpg" width="600" height="620" alt="FIG. 58." title="FIG. 58." /> +<p class="noidt"> +1. Ramphocephalus. Stonesfield Slate. John Phillips, 1871<br /> +2. Rhamphorhynchus. O. C. Marsh, 1882<br /> +3. Rhamphorhynchus. V. Zittel, 1882<br /> +4. Ornithostoma. Williston, 1897<br /> +5. Dimorphodon. Buckland, 1836. Tail then unknown<br /> +6. Ornithocheirus. H. G. Seeley, 1865<br /> +</p> +</div> + + + +<h4>THE SHORT-TAILED TYPES</h4> + +<p>The Pterodactylia are less variable; and the variation +among the species is chiefly confined to relative +length of the head, length of the neck, and the +height of the body above the ground. The tail is +always so short as to be inappreciable. Many of the +specimens are fragmentary, and the characters of the +group are not easily determined without careful +comparisons and measurements. The bones of the +<span class='pagenum'><a name="Page_166" id="Page_166">[Pg 166]</a></span> +fore limb and wing finger are less stout than in +the Rhamphorhynchus type, while the femur is +generally a little longer than the humerus, and the +wing finger is short in comparison with its condition +in Rhamphorhynchus. These short-tailed Pterodactyles +give the impression of being active little +animals, having very much the aspect of birds, upon +four legs or two. The neck is about as long as the +lower jaw, the antorbital vacuity in the head is imperfectly +separated from the much larger nasal opening, +the orbit of the eye is large and far back, the +teeth are entirely in front of the nasal aperture, and +the post-orbital vacuity is minute and inconspicuous. +The sternum is much wider than long, and no specimens +give evidence of a manubrium. The finger +bones progressively decrease in length. The prepubic +bones have a partially expanded fan-like form, +and never show the triradiate shape, and are never +anchylosed. About fifteen different kinds of Pterodactyles +have been described from the Solenhofen +Slate, mostly referred to the genus Pterodactylus, +which comprises forms with a large head and long +snout. Some have been placed in a genus (Ornithocephalus, +or Ptenodracon) in which the head +is relatively short. The majority of the species +are relatively small. The skull in <i>Ornithocephalus +brevirostris</i> is only 1 inch long, and the animal +could not have stood more than 1½ inches to its back +standing on all fours, and but little over 2½ inches +standing as a biped, on the hind limbs.</p> + +<p>A restoration of the species called <i>Pterodactylus +scolopaciceps</i>, published in 1875 in the <i>Illustrated +London News</i> in the position of a quadruped, shows +an animal a little larger, with a body 2½ inches high +<span class='pagenum'><a name="Page_167" id="Page_167">[Pg 167]</a></span> +and 6 to 7 inches long, with the wing finger 4½ inches +long. Larger animals occur in the same deposit, and +in one named <i>Pterodactylus grandis</i> the leg bones +are a foot long; and such an animal may have been +nearly a foot in height to its back, standing as a +quadruped, though most of these animals had the +neck flexible and capable of being raised like the +neck of a Goose or a Deer (<a href="#Page_30">p. 30</a>), and bent down +like a Duck's when feeding.</p> + +<div class="figcenter" style="width: 877px;"> +<a name="Fig_59" id="Fig_59"></a> +<span class="caption">FIG. 59. RESTORATION OF THE SKELETON OF +<i>PTENODRACON BREVIROSTRIS</i></span> +<p class="center">From the Solenhofen Slate. The fourth joint of the wing finger appears to +be lost and has not been restored in the figure. (Natural size)</p> +<img src="images/i_194.jpg" width="877" height="768" alt="FIG. 59." title="FIG. 59." /> +</div> + + +<p>The type of the genus Pterodactylus is the form +originally described by Cuvier as<i> Pterodactylus longirostris</i> +(<a href="#Page_28">p. 28</a>). It is also known as <i>P. antiquus</i>, that +name having been given by a German naturalist after +Cuvier had invented the genus, and before he had +<span class='pagenum'><a name="Page_168" id="Page_168">[Pg 168]</a></span> +named the species. There are some remarkable +features in which Cuvier's animal is distinct from +others which have been referred to the same +genus. Thus the head is 4½ inches long, while +the entire length of the backbone to the extremity +of the tail is only 6½ inches, and one +vertebra in the neck is at least as long as six in +the back, so that the animal has the greater part +of its length in the head and neck, although the +neck includes so few vertebræ. Nearly all the teeth—which +are few in number, short and broad, not +exceeding a dozen in either jaw—are limited to the +front part of the beak, and do not extend anywhere +near the nasal vacuity. This is not the case with all.</p> + +<p>In the species named <i>P. Kochi</i>, which I have regarded +as the type of a distinct genus, there are +large teeth in the front of the jaw corresponding to +those of Pterodactylus, and behind these a smaller +series of teeth extending back under the nostril, +which approaches close to the orbit of the eye, +without any indication of a separate antorbital +vacuity. On those characters the genus Diopecephalus +was defined. It is closely allied to Pterodactylus; +both agree in having the ilium prolonged +forward more than twice as far as it is carried backward, +the anterior process covering about half a +dozen vertebræ, as in <i>Pterodactylus longirostris</i>. A +great many different types have been referred to +<i>Pterodactylus Kochi</i>, and it is probable that they +may eventually be distinguished from each other. +The species in which the upper borders of the orbits +approximate could be separated from those in which +the frontal interspace is wider.</p> + + +<div class="figcenter" style="width: 854px;"> +<a name="Fig_60" id="Fig_60"></a> +<span class="caption">FIG. 60. CYCNORHAMPHUS SUEVICUS FROM THE SOLENHOFEN SLATE +SHOWING THE SCATTERED POSITION OF THE BONES</span> +<p class="center"><i>Original in the Museum at Tübingen</i></p> +<img src="images/i_196.jpg" width="854" height="768" alt="FIG. 60." title="FIG. 60." /> +</div> + + +<p><span class='pagenum'><a name="Page_169" id="Page_169">[Pg 169]</a></span></p> +<div class="figcenter" style="width: 957px;"> +<a name="Fig_61" id="Fig_61"></a> +<span class="caption">FIG. 61. CYCNORHAMPHUS SUEVICUS<br /><br /> +RESTORATION SHOWING THE FORM OF THE BODY AND THE WING MEMBRANES</span> +<img src="images/i_198.jpg" width="957" height="768" alt="FIG. 61." title="FIG. 61." /> +</div> + + +<p>It is a remarkable feature in these animals that +the middle bones of the foot, termed instep bones +or metatarsals, are usually close together, so that +the toes diverge from a narrow breadth, as in <i>P. +longirostris</i>, <i>P. Kochi</i>, and other forms; but there +also appear to be splay-footed groups of Pterodactyles +like the species which have been named +<i>P. elegans</i> and <i>P. micronyx</i>, in which the metatarsus +widens out so that the bones of the toes do not +diverge, and that condition characterises the Ptenodracon +(<i>Pterodactylus brevirostris</i>), to which genus +these species may possibly be referred. Nearly all +who have studied these animals regard the singularly +short-nosed species <i>P. brevirostris</i> as forming a +separate genus. For that genus Sömmerring's descriptive +name Ornithocephalus, which he used for +Pterodactyles generally, might perhaps have been +retained. But the name Ptenodracon, suggested by +Mr. Lydekker, has been used for these types.</p> + + +<p>Some of the largest specimens preserved at Stuttgart +and Tübingen have been named <i>Pterodactylus +suevicus</i> and <i>P. Fraasii</i>. They do not approach the +species <i>P. grandis</i> in size, so far as can be judged +from the fragmentary remains figured by Von Meyer; +for what appears to be the third phalange of the +wing finger is 7½ inches long, while in these species +it is less than half that length, indicating an enormous +development of wing, relatively to the length +of the hind limb, which would probably refer the +species to another genus. <i>Pterodactylus suevicus</i> +differs from the typical Pterodactyles in having a +rounded, flattened under surface to the lower jaw, +instead of the common condition of a sharp keel +in the region of the symphysis. The beak also seems +flattened and swan-like, and the teeth are limited to +<span class='pagenum'><a name="Page_170" id="Page_170">[Pg 170]</a></span> +the front of the jaw. There appear to be some +indications of small nostrils, which look upward like +the nostrils of Rhamphorhynchus, but this may be +a deceptive appearance, and the nostrils are large +lateral vacuities, which are in the position of antorbital +vacuities, so that there would appear to be +only two vacuities in the side of the head in these +animals. The distinctive character of the skeleton in +this genus is found in the extraordinary length +of the metacarpus and in the complete ossification +of the smaller metacarpal bones throughout their +length. The metacarpal bones are much longer than +the bones of the fore-arm, and about twice the length +of the humerus. The first wing phalange is much +longer than the others, which successively and rapidly +diminish in length, so that the third is half the length +of the first. There are differences in the pelvis; for +the anterior process of the ilium is very short, in comparison +with its length in the genus Pterodactylus. +And the long stalk of the prepubic bone with its great +hammer-headed expansion transversely in front gives +those bones a character unlike other genera, so that +Cycnorhamphus ranks as a good genus, easily distinguished +from Cuvier's type, in which the four bones +of the wing are more equal in length, and the last is +more than half the length of the first; while the +metacarpus in that genus is only a little longer than +the humerus, and much shorter than the ulna. The +<i>Pterodactylus suevicus</i> has the neck vertebræ flat on +the under side, and relatively short as compared +with the more slender and narrower vertebræ of +<i>P. Fraasii</i>.</p> + +<div class="figcenter" style="width: 599px;"> +<a name="Fig_62" id="Fig_62"></a> +<span class="caption">FIG. 62. <i>CYCNORHAMPHUS SUEVICUS</i></span> +<p class="center">Skeleton restored from the bones in Fig. 60</p> +<img src="images/i_201.jpg" width="599" height="480" alt="FIG. 62." title="FIG. 62." /> +</div> + +<div class="figcenter" style="width: 768px;"> +<a name="Fig_63" id="Fig_63"></a> +<span class="caption">FIG. 63. RESTORATION OF SKELETON CYCNORHAMPHUS FRAASI<br /><br /> +SHOWING THE LIMBS ON THE RIGHT SIDE</span> +<p class="center"><i>From a specimen in the Museum at Stuttgart</i></p> +<img src="images/i_202.jpg" width="768" height="911" alt="FIG. 63." title="FIG. 63." /> +</div> + + + +<p><span class='pagenum'><a name="Page_171" id="Page_171">[Pg 171]</a></span></p> +<div class="figcenter" style="width: 768px;"> +<a name="Fig_64" id="Fig_64"></a> +<span class="caption">FIG. 64. CYCNORHAMPHUS FRAASI<br /><br /> +RESTORATION OF THE FORM OF THE BODY</span> +<img src="images/i_204.jpg" width="768" height="894" alt="FIG. 64." title="FIG. 64." /> +</div> + + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_172" id="Page_172">[Pg 172]</a></span></p> +<h2><a name="CHAPTER_XV" id="CHAPTER_XV"></a><small>CHAPTER XV</small><br /><br /> + +ORNITHOSAURS FROM THE UPPER +SECONDARY ROCKS</h2> + + +<p>When staying at Swanage, in Dorsetshire, many +years ago, I had the rare good fortune to obtain +from the Purbeck Beds the jaw of a Pterodactyle, +which had much in common in plan with the <i>Cycnorhamphus +Fraasii</i> from the Lithographic Slate, which +is preserved at Stuttgart. The tooth-bearing part of +this lower jaw is 8 inches long as preserved, extending +back 3 inches beyond the symphysis portion in which +the two sides are blended together. It is different +from Professor Fraas's specimen in having the teeth +carried much further back, and in the animal being +nearly twice as large. This fragment of the jaw is +little more than 1 foot long, which is probably less +than half its original length. A vertebra nearly +5 inches long, which is more than twice the length +of the longest neck bones in the Stuttgart fossil, is +the only indication of the vertebral column. Professor +Owen described a wing finger bone from these +Purbeck Beds, which is nearly 1 foot long. He terms +it the second of the finger. It may be the third, and +on the hypothesis that the animal had the proportions +of the Solenhofen fossil just referred to, the first wing +<span class='pagenum'><a name="Page_173" id="Page_173">[Pg 173]</a></span> +finger bone of the English Purbeck Pterodactyle +would have exceeded 2 feet in length, and would +give a length for the wing finger of about 5 feet +3 inches. For this animal the name Doratorhynchus +was suggested, but at present I am unable to distinguish +it satisfactorily from Cycnorhamphus, which +it resembles in the forms both of the neck bones and +of the jaw. Very small Pterodactyles are also found +in the English Purbeck strata, but the remains are +few, and scattered, like these larger bones.</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_65" id="Fig_65"></a> +<span class="caption">FIG. 65. THE LONGEST KNOWN NECK VERTEBRA</span> +<p class="center">From the Purbeck Beds of Swanage. (Half natural size)</p> +<img src="images/i_208a.jpg" width="640" height="174" alt="FIG. 65." title="FIG. 65." /> +</div> + + + + +<h4>ORNITHODESMUS LATIDENS</h4> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_66" id="Fig_66"></a> +<span class="caption">FIG. 66. CERVICAL VERTEBRA OF ORNITHODESMUS</span> +<p class="center">From the Wealden Beds of the Isle of Wight</p> +<img src="images/i_208b.jpg" width="640" height="418" alt="FIG. 66." title="FIG. 66." /> +</div> + + + +<p>The Wealden strata being shallow, fresh-water +deposits might have been expected to supply better +knowledge of Pterodactyles than has hitherto been +<span class='pagenum'><a name="Page_174" id="Page_174">[Pg 174]</a></span> +available. Jaws of Ornithocheirus sagittirostris have +been found in the beds at Hastings, and in other +parts of Sussex. Some fragments are as large as +anything known. The best-preserved remains have +come from the Isle of Wight, and were rewards to +the enthusiastic search of the Rev. W. Fox, of Brixton. +In the principal specimen the teeth were short and +wide, the head large and deep with large vacuities, +but the small brain case of that skull is bird-like. +The neck bones are 2½ inches long. In the upper +part of the back the bones are united together by +anchylosis, so that they form a structure in the back +like a sacrum, which does not give attachment to the +scapula, as in some Pterodactyles from the Chalk, but +the bones are simply blended, as in the frigate-bird, +allied to Pelicans and Cormorants. And then after a +few free vertebræ in the lower part of the back, succeeds +the long sacrum, formed in the usual way, of many +vertebræ. I described a sacrum of this type from the +Wealden Beds, under the name <i>Ornithodesmus</i>, referable +to another species, which in many respects was +so like the sacrum of a Bird that I could not at the +time separate it from the bird type. This genus has +a sternum with a strong deep keel, and the articulation +for the coracoid bones placed at the back of the +keel in the usual way, but with a relation to each +other seen in no genus hitherto known, for the +articular surfaces are wedge-shaped instead of being +ovate; and instead of being side by side, they obliquely +overlap, practically as in wading birds like the +Heron. I have never seen any Pterodactyle teeth so +flattened and shaped like the end of a lancet; and +from this character the form was known between +Mr. Fox and his friends as "latidens." The name<span class='pagenum'><a name="Page_175" id="Page_175">[Pg 175]</a></span> +Ornithodesmus is as descriptive of the sternum as of +the vertebral column. The wing bones, as far as +they are preserved, have the relatively great strength +in the fore limb which is found in many of the Pterodactyles +of the Cretaceous period, and are quite as +large as the largest from the Cambridge Greensand. +In the Sussex species named <i>P. sagittirostris</i> the +lower jaw articulation was inches wide.</p> + + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_67" id="Fig_67"></a> +<span class="caption">FIG. 67. STERNUM OF <i>ORNITHODESMUS</i></span> +<p class="center">Showing the overlapping facets for the coracoid bones (shaded) +behind the median keel</p> +<img src="images/i_210a.jpg" width="640" height="377" alt="FIG. 67." title="FIG. 67." /> +</div> + + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_68" id="Fig_68"></a> +<span class="caption">FIG. 68. FRONT OF THE KEEL OF THE STERNUM OF +<i>ORNITHODESMUS LATIDENS</i></span> +<p class="center">Showing also the articulation for the coracoid bone</p> +<img src="images/i_210b.jpg" width="640" height="416" alt="FIG. 68." title="FIG. 68." /> +</div> + +<p>A few Pterodactyles' bones have been discovered +<span class='pagenum'><a name="Page_176" id="Page_176">[Pg 176]</a></span> +in the Neocomian sands of England and Germany, +and other larger bones occur in the Gault of Folkestone +and the north of France; but never in such +association as to throw light on the aspect of the +skeleton.</p> + + +<h4>ORNITHOCHEIRUS</h4> + +<p>Within my own memory Pterodactyle remains +were equally rare from the Cambridge Greensand. +The late Professor Owen in one of his public lectures +produced the first few fragments received from +Cambridge, and with a knowledge which in its +scientific method seemed to border on the power of +creation, produced again the missing parts, so that +the bones told their story, which the work of waves +and mineral changes in the rock had partly obliterated. +Subsequently good fortune gave me the +opportunity during ten years to help my University +in the acquisition and arrangement of the finest +collection of remains of these animals in Europe. +Out of an area of a few acres, during a year or two, +came the thousand bones of Ornithosaurs, mostly +associated sets of remains, each a part of a separate +skeleton, described in my published catalogues, as +well as the best of those at York and in the British +Museum and other collections in London.</p> + +<p>The deposit which yields them, named Cambridge +Greensand, may or may not represent a long period +of time in its single foot of thickness; but the abundance +of fossils, obtained whenever the workmen were +adequately remunerated for preserving them, would +suggest that the Pterodactyles might have lived +like sea-birds or in colonies like the Penguins, if +it were not that the number of examples of each +species found is always small, and the many variations<span class='pagenum'><a name="Page_177" id="Page_177">[Pg 177]</a></span> +of structure suggested rather that the individuals +represent the life of many lands. The collections of +remains are mostly from villages in the immediate +vicinity of Cambridge, such as Chesterton, Huntingdon +Road, Coldham Common, Haslingfield, Barton, +Shillington, Ditton, Granchester, Harston, Barrington, +stretching south to Ashwell in Bedfordshire on +the one hand, as well as further north by Horningsea +into the fens. Each appears to be the associated +bones of a single individual. The remains mostly +belong to comparatively large animals. Some were +small, though none have been found so diminutive +as the smallest from the Solenhofen Slate. The +largest specimens with long jaws appear to have +had the head measuring not more than eighteen +inches in length, which is less than half the size of +the great toothless Pterodactyles from Kansas.</p> + + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_69" id="Fig_69"></a> +<span class="caption">FIG. 69. RESTORATION OF THE SKULL OF ORNITHOCHEIRUS</span> +<img src="images/i_212.jpg" width="640" height="221" alt="FIG. 69." title="FIG. 69." /> +<p class="center">The parts left white are in the Geological Museum at Cambridge. The shaded +parts have not been found. The two holes are the eye and the nostril<br /> +(From the Cambridge Greensand)</p> +</div> + +<p>The Cambridge specimens manifestly belong to at +least three genera. Something may be said of the +characters of the large animals which are included in +the genus Ornithocheirus. These fossils have many +points of structure in common with the great +American toothless forms which are of similar geological +age. The skull is remarkable for having the<span class='pagenum'><a name="Page_178" id="Page_178">[Pg 178]</a></span> +back of the head prolonged in a compressed median +crest, which rose above the brain case, and extended +upward and over the neck vertebræ, so as to indicate +a muscular power not otherwise shown in the group. +For about three inches behind the brain this wedge +of bone rested on the vertebræ, and probably overlapped +the first three neural arches in the neck.</p> + +<p>Another feature of some interest is the expansion +of the bone which comes below the eye. In Birds +this malar or cheek bone is a slender rod, but in +these Pterodactyles it is a vertical plate, which is +blended with the bone named the quadrate bone, +which makes the articulation with the lower jaw in +all oviparous animals.</p> + +<p>The beak varies greatly in length and in form, +though it is never quite so pointed as in the American +genus, for there is always a little truncation in front, +when teeth are seen projecting forward from a position +somewhat above the palate; the snout is often +massive and sometimes club-shaped. Except for these +variations of shape in the compressed snout, which is +characterised by a ridge in the middle of the palate, +and a corresponding groove in the lower jaw, and +the teeth, there is little to distinguish what is known +of the skull in its largest English Greensand fossils +from the skull remains which abound in the Chalk +of Kansas.</p> + +<p>This English genus Ornithocheirus, represented by +a great number of species, had the neural arch of +the neck bones expanded transversely over the body +of the vertebra in a way that characterises many +birds with powerful necks, and is seen in a few +Pterodactyles from Solenhofen.</p> + +<p>It is difficult to resist the conclusion that the neck<span class='pagenum'><a name="Page_179" id="Page_179">[Pg 179]</a></span> +vertebræ were not usually more than twice to three +times as long as those of the back, and it would +appear that the caudal vertebræ in the English +Cretaceous types were comparatively large, and +about twice as long as the dorsal vertebræ. Unless +there has been a singular succession of accidents in +the association of these vertebræ with the other remains, +Ornithocheirus had a tail of moderate length, +formed of a few vertebræ as long as those of the +neck, though more slender, quite unlike the tail in +either the long-tailed or short-tailed groups of Solenhofen +Pterodactyles, and longer than in the toothless +Pterodactyles of America.</p> + + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_70" id="Fig_70"></a> +<span class="caption">FIG. 70. CERVICAL VERTEBRA, ORNITHOCHEIRUS</span> +<p class="center">Under side, half natural size. (Cambridge Greensand)</p> +<img src="images/i_214.jpg" width="640" height="370" alt="FIG. 70." title="FIG. 70." /> +</div> + +<p>The singular articulation for the humerus at the +truncated extremity of the coracoid bone is a +character of this group, as is the articulation of the +scapulæ with the neural arches of the dorsal vertebræ, +at right angles to them (<a href="#Page_115">p. 115</a>), instead of running +over the ribs as in Birds and as in other Pterodactyles.</p> + +<p>The smaller Pterodactyles have their jaws less compressed +from side to side. The upper arm bone, the +humerus, instead of being truncated at its lower end +as in Ornithocheirus, is divided into two or three<span class='pagenum'><a name="Page_180" id="Page_180">[Pg 180]</a></span> +rounded articular surfaces. That for the radius, the +bone which carries the wrist, is a distinct and oblique +rounded facet, while the ulna has a rounded and +pulley-like articulation on which the hand may rotate. +These differences are probably associated with an +absence of the remarkable mode of union of the +scapulæ with the dorsal vertebræ. But I have +hesitated to give different names to these smaller +genera because no example of scapula has come +under my notice which is not truncated at the free +end. I do not think this European type can be +the Nyctodactylus of Professor Marsh, in which +sutures appear to be persistent between the bodies +of the vertebræ and their arches, because no examples +have been found at Cambridge with the neural arches +separated, although the scapula is frequently separated +from the coracoid in large animals.</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_71" id="Fig_71"></a> +<span class="caption">FIG. 71. UPPER AND LOWER JAWS OF AN ENGLISH PTERODACTYLE +FROM THE CHALK, AS PRESERVED</span> +<img src="images/i_215.jpg" width="640" height="301" alt="FIG. 71." title="FIG. 71." /> +</div> + + +<div class="figcenter" style="width: 618px;"> +<a name="Fig_72" id="Fig_72"></a> +<span class="caption">FIG. 72. THE PALATE OF THE ENGLISH TOOTHLESS +PTERODACTYLE, ORNITHOSTOMA</span> +<img src="images/i_216.jpg" width="618" height="480" alt="FIG. 72." title="FIG. 72." /> +</div> + + +<div class="figcenter" style="width: 480px;"> +<a name="Fig_73" id="Fig_73"></a> +<span class="caption">FIG. 73. TYPES OF THE AMERICAN TOOTHLESS +PTERODACTYLE, ORNITHOSTOMA</span> +<p class="center">Named by Marsh, Pteranodon</p> +<img src="images/i_217.jpg" width="480" height="489" alt="FIG. 73." title="FIG. 73." /> +</div> + + + + +<h4>ORNITHOSTOMA</h4> + +<p>The most interesting of all the English Pterodactyle +remains is the small fragment of jaw figured by +Sir Richard Owen in 1859, which is a little more than +<span class='pagenum'><a name="Page_181" id="Page_181">[Pg 181]</a></span> +two inches long and an inch wide, distinguished by a +concave palate with smooth rounded margins to the +jaws and a rounded ridge to the beak. It is the only +satisfactory fragment of the animal which has been +figured, and indicates a genus of toothless Pterodactyles, +for which the name Ornithostoma was first used +in 1871. After some years Professor Marsh found +toothless Pterodactyles in Kansas, and indicated +several species. There are remains to the number of +six hundred specimens of these American animals in +the Yale Museum alone; but very little was known of +them till Professor Williston, of Lawrence, in Kansas, +described the specimens from the Kansas University +Museum, when it became evident that the bones of +the skeleton are mostly formed on the same plan +as those of the Cambridge Greensand genus, Ornithocheirus. +They are not quite identical. Professor +Williston adopts for them the name Ornithostoma, +in preference to Pteranodon which Marsh had +<span class='pagenum'><a name="Page_182" id="Page_182">[Pg 182]</a></span> +suggested. Both animals have the dagger-shaped form +of jaw, with corresponding height and breadth of the +palate. The same flattened sides to the snout, converging +upwards to a rounded ridge, the same compressed +rounded margin to the jaw, which represents +the border in which teeth are usually implanted, and +in both the palate has the same smooth character +forming a single wide concave channel. Years previously +I had the pleasure of showing to Professor +Marsh the remarkable characters of the jaw, shoulder-girdle +bones, and scapulæ in the Greensand Pterodactyles +while the American fossils were still undiscovered. +I subsequently made the restoration of the +shoulder-girdle (<a href="#Page_115">p. 115</a>). Professor Williston states to +<span class='pagenum'><a name="Page_183" id="Page_183">[Pg 183]</a></span> +me that the shoulder-girdle bones in American examples +of Ornithostoma have a close resemblance to +those of Ornithocheirus figured in 1891, as is evident +from remains now shown in the British Museum. It +appears that the Kansas bones are almost invariably +crushed flat, so that their articular ends are distorted. +The neck vertebræ are relatively stout as in Ornithocheirus. +The hip-girdle of the American Ornithostoma +can be closely paralleled in some English +specimens of Ornithocheirus, though each prepubic +bone is triangular in the American fossils as in +<i>P. rhamphastinus</i>. They are united into a transverse +bar as in Rhamphorhynchus, unknown in the English +fossils. The femur has the same shape as in Ornithocheirus; +and the long tibia terminates in a pulley. +There is no fibula. The sternum in both has a +manubrium, or thick keel mass, prolonged in front +of its articular facets for the coracoid bones, which +are well separated from each other. Four ribs +articulate with its straight sides. The animal has +four toes and the fifth is rudimentary; there are no +claws to the first and second.</p> + +<div class="figcenter" style="width: 800px;"> +<a name="Fig_74" id="Fig_74"></a> +<span class="caption">FIG. 74. RESTORATION OF THE SKELETON OF +<i>ORNITHOSTOMA INGENS</i> (<span class="smcap">Marsh</span>)</span> +<img src="images/i_218.jpg" width="800" height="311" alt="FIG. 74." title="FIG. 74." /> +<p class="center">From the Niobrara Cretaceous of Western Kansas. Made by Professor Williston. +The original has a spread of wing of about 19 feet 4 inches. Fragments of +larger individuals are preserved at Munich</p> +</div> + + +<p><span class='pagenum'><a name="Page_184" id="Page_184">[Pg 184]</a></span>In +the restoration which Professor Williston has +made the wing metacarpal is long, and in the shortest +specimen measures 1 foot 7 inches, and in the longest +1 foot 8 inches. This is exactly equal to the length +of the first phalange of the wing finger. The second +wing finger bone is 3 inches shorter, the third is little +more than half the length of the first, while the fourth +is only 6¾ inches long, showing a rapid shortening of +the bones, a condition which may have characterised +all the Cretaceous Pterodactyles. The short +humerus, about 1 foot long, and the fore-arm, which +is scarcely longer, are also characteristic proportions +of Ornithostoma or Pteranodon, as known from the +American specimens. Professor Williston gives no +details of the remarkable tail, beyond saying that the +tail is small and short, and that the vertebræ are flat +at the ends, without transverse processes. In the restoration +the tail is shorter than in the short-tailed +species from the Lithographic Slate, and unlike the +tail in Ornithocheirus.</p> + +<p> </p> + +<p>This is the succession of Pterodactyles in geological +time. Their history is like that of the human +race. In the most ancient nations man's life comes +upon us already fully organised. The Pterodactyles +begin, so far as isolated bones are concerned, in the +Rhætic strata; perhaps in the Muschelkalk or middle +division of the Trias. And from the beginning +of the Secondary time they live on with but little +diversity in important and characteristic structures, +and so far as habit goes, the great Pterodactyles +of the Upper Chalk of England cannot be said to +be more highly organised than the earlier stiff-tailed +genera of the Lias or the Oolites. There is nothing<span class='pagenum'><a name="Page_185" id="Page_185">[Pg 185]</a></span> +like evolution. No modification such as that which +derives the one-toed horse or the two-toed ox from +ancestors with a larger number of digits. On the +other hand, there is little, if any, evidence of degeneration. +The later Pterodactyles do not appear +to have lost much, although the tail in some of the +Solenhofen genera may be degenerate when compared +with the long tail of Dimorphodon; but the +short-tailed types are found side by side with the +long-tailed Rhamphorhynchus. The absence of teeth +may be regarded as degeneration, for they have +presumably become lost, in the same way that Birds +now existing have lost the teeth which characterised +the fossil birds—Ichthyornis of the American Greensand, +and Archæopteryx of the Upper Oolites of +Bavaria. But just as some of the earlier Pterodactyles +have no teeth at the extremity of the jaw, such +as Dorygnathus and Rhamphorhynchus, so the loss +of teeth may have extended backward till the jaws +became toothless. The specimens hitherto known +give no evidence of such a change being in progress. +But just as the division of Mammals termed Edentata +usually wants only the teeth which characterise the +front of the jaw, yet others, like the Great Ant-eater +of South America named Myrmecophaga, have the +jaws as free from teeth as the toothless Pterodactyles +or living Birds, and show that in that order the teeth +have no value in separating these animals into subordinate +groups any more than they have among the +Monotremata, where one type has teeth and the other +is toothless.<span class='pagenum'><a name="Page_186" id="Page_186">[Pg 186]</a></span></p> + +<p>The following table gives a summary of the Geological +History and succession in the Secondary +Rocks of the principal genera of Flying Reptiles.</p> + + +<div class='center'> +<table border="1" cellpadding="4" cellspacing="0" summary=""> +<tr><th rowspan='2'>GEOLOGICAL FORMATIONS.</th><th colspan='2'>NAMES OF THE GENERA.</th></tr> +<tr><th>British and European.</th><th>North American.</th></tr> +<tr> + <td align='left'>Upper Chalk<br /><br />Lower Chalk<br />Upper Greensand<br />Gault</td> + <td align='left'><br /><br />} Ornithocheirus<br />} Ornithostoma</td> + <td align='left'>} Ornithostoma<br />} (<i>Pteranodon</i>)<br />} Nyctodactylus<br /><br /> </td> +</tr> +<tr> + <td align='left'>Lower Greensand<br />Wealden<br />Purbeck</td> + <td align='left'>Ornithodesmus<br />Doratorhynchus</td> + <td align='left'></td> +</tr> +<tr> + <td align='left'>Portland<br />Kimeridge Clay and Solenhofen Slate<br />Coralline Oolite<br />Oxford Clay</td> + <td align='left'>{ Pterodactylus<br />{ Ptenodracon<br />{ Cycnorhamphus<br />{ Diopecephalus<br />{ Rhamphorhynchus<br />{ Scaphognathus</td> + <td> </td> +</tr> +<tr> + <td align='left'>Great Oolite and Stonesfield Slate<br />Inferior Oolite</td> + <td align='left'>Rhamphocephalus</td> + <td> </td> +</tr> +<tr> + <td align='left'>Upper Lias<br /><br />Lower Lias</td> + <td align='left'>{ Campylognathus<br />{ Dorygnathus<br />Dimorphodon</td> + <td align='left'> </td> +</tr> +<tr> + <td align='left'>Rhætic<br /><br />Muschelkalk</td> + <td align='left'>bones<br /><br />? bones</td> + <td align='left'> </td> +</tr> +</table></div> + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_187" id="Page_187">[Pg 187]</a></span></p> +<h2><a name="CHAPTER_XVI" id="CHAPTER_XVI"></a><small>CHAPTER XVI</small><br /><br /> + +CLASSIFICATION OF THE +ORNITHOSAURIA</h2> + + +<p>When an attempt is made to determine the +place in nature of an extinct group of animals +and the relation to each other of the different types +included within its limits, so as to express those facts +in a classification, attention is directed in the first +place to characters which are constant, and persist +through the whole of its constituent genera. We +endeavour to find the structural parts of the skeleton +which are not affected by variation in the dentition, +or the proportions of the extremities, or length of +the tail, which may define families or genera, or +species.</p> + +<p>It has already been shown that while in many +ways the Ornithosaurian animals are like Birds, they +have also important resemblances to Reptiles. They +are often named Pterosauria. The wing finger gives +a distinctive character which is found in neither one +class of existing animals nor the other, and is common +to all the Pterodactyles at present known. They have +been named Ornithosauria as a distinct minor division +of back-boned animals, which may be regarded as +neither Reptiles nor Birds in the sense in which those<span class='pagenum'><a name="Page_188" id="Page_188">[Pg 188]</a></span> +terms are used to define a Lizard or Ostrich among +animals which still exist. It is not so much that they +mark a transition from Reptile to Bird, as that they +are a group which is parallel to Birds, and more +manifestly holds an intermediate place than Birds do +between Reptiles and Mammals. In plan of structure +Bird and Reptile have more in common than was +at one time suspected. The late Professor Huxley +went so far as to generalise on those coincidences +in parts of the skeleton, and united Birds and Reptiles +into one group, which he named Sauropsida, to express +the coincidences of structure between the Lizard +and the Bird tribes. The idea is of more value than +the term in which it is expressed, because Reptiles +are not, as we have seen, a group of animals which +can be defined by any set of characters as comprehensive +as those which express the distinctive features +of Birds. From the anatomist's point of view Birds +are a smaller group, and while some Reptiles have +affinity with them, it is rather the extinct than the +living groups which indicate that relation. Other +Reptiles have affinities of a more marked kind with +Mammals, and there are points in the Ornithosaurian +skeleton which are distinctly Mammalian. So that +when the Monotreme Mammals are united with +South African reptiles known as Theriodontia, which +resemble them, in a group termed Theropsida to +express their mammalian resemblances, it is evident +that there is no one continuous chain of life or gradation +in complexity of structure of animals.</p> + +<p>We have to determine whether the Ornithosauria incline +towards the Sauropsidan or Bird-Reptile alliance, +or to the Mammal-Reptile or Theropsidan alliance. +There can be no doubt that the predominant ten<span class='pagenum'><a name="Page_189" id="Page_189">[Pg 189]</a></span>dency +is to the former, with a minor affinity towards +the latter.</p> + +<p>The Ornithosauria are one of a series of groups +of animals, living and extinct, which have been +combined in an alliance named the Ornithomorpha. +That group includes at least five great divisions +of animals, which circle about birds, known as +Ornithosauria, Crocodilia, Saurischia, Aves, Ornithischia, +and Aristosuchia. Their relations to each other +are not evident in an enumeration, but may be shown +in some degree in a diagram (see <a href="#Page_190">p. 190</a>).</p> + + +<h4>THE ORNITHOMORPHA</h4> + +<p>The Ornithomorpha arranged in this way show +that the three middle groups—carnivorous Saurischia, +Aristosuchia, herbivorous Ornithischia—which are +usually united as Dinosauria, intervene between +Birds and Ornithosaurs; and that the Crocodilia +and Ornithosauria are parallel groups which are connected +with Birds, by the group of Dinosaurs, which +resembles Birds most closely.</p> + +<p>The Ornithomorpha is only one of a series of large +natural groups of animals into which living and +extinct terrestrial vertebrata may be arranged. And +the succeeding diagram may contribute to make +evident the relations of Ornithosauria to the other +terrestrial vertebrata (see <a href="#Page_191">p. 191</a>).</p> + +<p>Herein it is seen that while the Ornithomorpha +approach towards Mammalia through the Ornithosauria, +and less distinctly through the Crocodilia, +they approach more directly to the Sauromorpha, +through the Plesiosaurs and Hatteria; while +that group also approaches more directly to the +Mammals through the Plesiosaurs and Anomodonts.</p> +<p><span class='pagenum'><a name="Page_190" id="Page_190">[Pg 190]</a></span></p> + +<h3><span class="smcap">Diagram of the Affinities of the Orders of Animals +comprised in the Ornithomorpha.</span></h3> + +<div class="figcenter" style="width: 600px;"> +<img src="images/i_225.jpg" width="600" height="644" alt="" title="" /> +<span class="caption">After a diagram in the <i>Philosophical Transactions of the +Royal Society</i>, 1892.</span> +</div> + +<p>The Aristosuchia is imperfectly known, and therefore +to some extent a provisional group. It is a +small group of animals.</p> +<p><span class='pagenum'><a name="Page_191" id="Page_191">[Pg 191]</a></span></p> + +<h3><span class="smcap">Diagram showing the Relations of the Ornithomorpha +to the chief large groups of Terrestrial Vertebrata,<br /> +and their affinities with each other.</span></h3> + +<div class="figcenter" style="width: 499px;"> +<img src="images/i_226.jpg" width="499" height="480" alt="" title="" /> +<span class="caption">After a diagram in the <i>Philosophical Transactions of the +Royal Society</i>, 1892.</span> +</div> + + + +<p>Cordylomorpha are Ichthyosaurs and the Labyrinthodont +group. Herpetomorpha include Lacertilia, +Homœosauria, Dolichosauria, Chameleonoidea, +Ophidia, Pythonomorpha.</p> + +<p><span class='pagenum'><a name="Page_192" id="Page_192">[Pg 192]</a></span></p> +<p>The Sauromorpha comprises the groups of extinct +and living Reptiles named Chelonia, Rhynchocephala, +Sauropterygia, Anomodontia, Nothosauria, and Protorosauria. +These details may help to explain the +place which has been given to the Ornithosauria in +the classification of animals.</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_75" id="Fig_75"></a> +<span class="caption">FIG. 75. COMPARISON OF SIX GENERA</span> +<p class="center">The skulls are seen on the left side in the order of the names below them</p> +<img src="images/i_227.jpg" width="640" height="458" alt="FIG. 75." title="FIG. 75." /> +</div> + + + +<p>Turning to the Pterodactyles themselves, Von +Meyer divided them naturally into short-tailed and +long-tailed. The short-tailed indicated by the name +Pterodactylus he further divided into long-nosed and +short-nosed. The short-nosed genus has since been +named Ptenodracon (<a href="#Fig_59">Fig. 59, p. 167</a>). The long-tailed +group was divided into two types—the Rhamphorhynchus +of the Solenhofen Slate (<a href="#Fig_56">Fig. 56, p. 161</a>) +and the English form now known as Dimorphodon +(<a href="#Fig_52">Fig. 52, p. 150</a>), which had been described from the +Lias.</p> + +<p>The Cretaceous Pterodactyles form a distinct<span class='pagenum'><a name="Page_193" id="Page_193">[Pg 193]</a></span> +family. So that, believing the tail to have been short +in that group (<a href="#Fig_58">Fig. 58</a>), there are two long-tailed as +well as two short-tailed families, which were defined +from their typical genera Pterodactylus, Ornithocheirus, +Rhamphorhynchus, and Dimorphodon.</p> + +<p>The differences in structure which these animals +present are, first: the big-headed forms from the Lias +like Dimorphodon, agree with the Rhamphorhynchus +type from Solenhofen in having a vacuity in the skull +defined by bone, placed between the orbit of the eye +and the nostril. With those characters are correlated +the comparatively short bones which correspond to +the back of the hand termed metacarpals, and the +tail is long, and stiffened down its length with ossified +tendons. These characters separate Ornithosaurs +with long tails from those with short tails.</p> + +<p>The short-tailed types represented by Pterodactylus +and Ornithocheirus have no distinct antorbital vacuity +in the skull defined by bone. The metacarpal bones +of the middle hand are exceptionally elongated, and +the tail, which was flexible in both, appears to have +been short. These differences in the skeleton warrant +a primary division of flying reptiles into two principal +groups.</p> + +<p>The short-tailed group, which was recognised by +De Blainville as intermediate between Birds and +Reptiles, may take the name Pterodactylia, which +he suggested as a convenient, distinctive name. It +may probably be inconvenient to enlarge its significance +to comprise not only the true Pterodactyles +originally defined as Pterosauria, but the newer +Ornithostoma and Ornithocheirus which have been +grouped as Ornithocheiroidea.</p> + +<p>The second order, in which the wing membrane<span class='pagenum'><a name="Page_194" id="Page_194">[Pg 194]</a></span> +appears to have had a much greater extent, in being +carried down the hind limbs, where the outermost +digit and metatarsal are modified for its support, has +been named Pterodermata, to include the types +which are arranged around Rhamphorhynchus and +Dimorphodon.</p> + +<p>Both these principal groups admit of subdivision +by many characters in the skeleton, the most remarkable +of which is afforded by the pair of bones carried +in front of the pubes, and termed prepubic bones. +In the Pterodactyle family the bones in front of the +pubes are always separate from each other, always +directed forward, and have a peculiar fan-shaped +form with concave sides like the bone which holds a +similar position in a Crocodile. In the Ornithocheirus +family the prepubic bones appear to have been originally +triangular, but were afterwards united so as +to form a strong continuous bar which extends transversely +across the abdomen in advance of the pubic +bones. This at least is the distinctive character in the +genus Ornithostoma according to Professor Williston, +which in many ways closely resembles Ornithocheirus.</p> + +<p>The two families in the long-tailed order named +Pterodermata are separated from each other by a +similar difference in their prepubic bones. In Dimorphodon +those bones are separate from each other, +and remain distinct through life, meeting in the +middle line of the body in a wide plate. On the +other hand, in Rhamphorhynchus the prepubic bones, +which are at first triangular and always slender, +become blended together into a slight transverse bar, +which only differs from that attributed to Ornithostoma +in its more slender bow-shaped form.</p> + + + +<p>Thus if other characters of the skeleton are +<span class='pagenum'><a name="Page_195" id="Page_195">[Pg 195]</a></span> +ignored and a classification based upon the structure of +the pelvis and prepubic bones, there would be some +ground for associating the long-tailed Rhamphorhynchus +from the Upper Oolites which is losing the +teeth in the front of its jaw with the Cretaceous Ornithostoma, +which has the teeth completely wanting; +while the long-tailed Dimorphodon would come into +closer association with the short-tailed Pterodactylus. +The drum-stick bone or tibia in Dimorphodon, with +its slender fibula, like that of a Bird, also resembles +a Bird in the rounded and pulley-shaped terminal +end which makes the joint corresponding to the +middle of the ankle bones in man. The same condition +of a terminal pulley joint is found in the +Cretaceous Pterodactyles. But in the true Pterodactyles +and in Rhamphorhynchus there usually is +no pulley-shaped termination to the lower end of +the drum-stick, for the tarsal bones remain separate +from each other, and form two rows of ossifications, +showing the same differences as separate Dinosaurs +into the divisions which have been referred to, from +their Bird-like pelvis and tibio-tarsus, as Ornithischia +in the one case, and Saurischia in the other from +their bones being more like those of living Lizards.</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_76" id="Fig_76"></a> +<span class="caption">FIG. 76. LEFT SIDE OF PELVIS OF ORNITHOSTOMA</span> +<p class="center">(After Williston)</p> +<img src="images/i_230.jpg" width="640" height="296" alt="FIG. 76." title="FIG. 76." /> +</div> + + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_196" id="Page_196">[Pg 196]</a></span></p> +<h2><a name="CHAPTER_XVII" id="CHAPTER_XVII"></a><small>CHAPTER XVII</small><br /><br /> + +FAMILY RELATIONS OF PTERODACTYLES +TO ANIMALS WHICH +LIVED WITH THEM</h2> + + +<p>Enough has been said of the general structure +of Pterodactyles and the chief forms which +they assumed while the Secondary rocks were accumulating, +to convey a clear idea of their relations +to the types of vertebrate animals which still survive +on the earth. We may be unable to explain the +reasons for their existence, and for their departure +from the plan of organisation of Reptiles and Birds. +But the evidence has not been exhausted which may +elucidate their existence. Sometimes, in problems of +this kind, which involve comparison of the details of +the skeleton in different animals, it is convenient to +imagine the possibility of changes and transitions +which are not yet supported by the discovery of +fossil remains. If, for example, the Pterodactyle be +conceived of as divested of the wing finger, which is +its most distinctive character, or that finger is supposed +to be replaced by an ordinary digit, like the three-clawed +digits of the hand which we have regarded +as applied to the ground, where, it may be asked, +would the animal type be found which approximates<span class='pagenum'><a name="Page_197" id="Page_197">[Pg 197]</a></span> +most closely to a Pterodactyle which had been thus +modified? There are two possible replies to such a +question, suggested by the form of the foot. For +the old Bird Archæopteryx has three such clawed +digits, but no wing finger. And some Dinosaurs also +have the hand with three digits terminating in claws, +which are quite comparable to the clawed digits of +Pterodactyles.</p> + +<p>The truth expressed in the saying that no man by +taking thought can add a cubit to his stature is of +universal application in the animal world, in relation +to the result upon the skeleton of the exercise of a +function by the individual. Yet such is the relation +in proportions of the different parts of the animal to +the work which it performs, so marked is the evidence +that growth has extended in direct relation to use of +organs and active life, and that structures have become +dwarfed from overwork, or have wasted away from +disuse—seen throughout all vertebrate animals, that +we may fairly attribute to the wing finger some correlated +influence upon the proportions of the animal, +as a consequence of the dependence of the entire +economy upon each of its parts. Therefore if an +allied animal did not possess a wing finger, and did +not fly, it might not have developed the lightness of +bone, or the length of limb which Pterodactyles +possess.</p> + +<p>The mere expansion of the parachute membrane +seen in so-called flying animals, both Mammals and +Reptiles, which are devoid of wings, is absolutely +without effect in modifying the skeleton. But when +in the Bat a wing structure is met with which may +be compared to a gigantic extension of the web foot +of the so-called Flying Frog, the bones of the fingers<span class='pagenum'><a name="Page_198" id="Page_198">[Pg 198]</a></span> +and the back of the hand elongate and extend under +the stimulus of the function of flight in the same +way as the legs elongate in the more active hoofed +animals, with the function of running. Therefore it +is not improbable that the limbs shared to some +extent in growth under stimulus of exercise which +developed the wing finger. And if an animal can be +found among fossils so far allied as to indicate a +possible representative of the race from which these +Flying Dragons arose, it might be expected to be at +least shorter legged, and possibly more distinctly +Reptilian in the bones of the shoulder-girdle which +support the muscles used in flight. It may readily +be understood that the kinds of life which were most +nearly allied to Pterodactyles are likely to have +existed upon the earth with them, and that flight was +only one of the modes of progression which became +developed in relation to their conditions of existence. +The principal assemblage of terrestrial animals +available for such comparison is the Dinosauria. They +may differ from Pterodactyles as widely as the Insectivora +among Mammals differ from Bats, but not +in a more marked way. Comparisons will show that +there are resemblances between the two extinct groups +which appeal to both reason and imagination.</p> + +<p>Dinosaurs are conveniently divided by characters +of the pelvis first into the order Saurischia, which +includes the carnivorous Megalosaurus and the Cetiosaurus, +with the pelvis on the Reptile plan; and +secondly the order Ornithischia, represented by Iguanodon, +with the pelvis on the Bird plan. It may be +only a coincidence, but nevertheless an interesting +one, that the characters of those two great groups of +reptiles, which also extend throughout the Secondary<span class='pagenum'><a name="Page_199" id="Page_199">[Pg 199]</a></span> +rocks, are to some extent paralleled in parts of the +skeleton of the two divisions of Pterodactyles. This +may be illustrated by reference to the skull, pelvis, +hind limb, and the pneumatic condition of the bones.</p> + +<div class="figcenter" style="width: 524px;"> +<a name="Fig_77" id="Fig_77"></a> +<span class="caption">FIG. 77. COMPARISON OF THE SKULL OF THE +DINOSAUR ANCHISAURUS WITH THE ORNITHOSAUR DIMORPHODON</span> +<img src="images/i_234.jpg" width="524" height="480" alt="FIG. 77." title="FIG. 77." /> +</div> + + +<p>The Saurischian Dinosauria have an antorbital +vacuity in the side of the skull between the nasal +opening and the eye, as in the long-tailed Ornithosaurs +named Pterodermata. In some of the older +genera of these carnivorous Dinosaurs of the Trias, +the lateral vacuities of the head are as large as in +Dimorphodon. But in some at least of the Iguanodont, +or Ornithischian Dinosaurs, there is no antorbital +vacuity, and the side of the face in that +respect resembles the short-tailed Pterodactylia. +The skull of a carnivorous Dinosaur possesses teeth +which, though easily distinguished from those of +Pterodactyles, can be best compared with them. The<span class='pagenum'><a name="Page_200" id="Page_200">[Pg 200]</a></span> +most striking difference is in the fact that in the +Dinosaur the nostrils are nearly terminal, while in +the Pterodactyle they are removed some distance +backward. This result is brought about by growth +taking place, in the one case at the front margin of +the maxillary bone so as to carry the nostril forward, +and in the other case at the back margin of the premaxillary +bone. Thus an elongated part of the jaw +is extended in front of the nostril. Hence there is +a different proportion between the premaxillary and +maxillary bones in the two groups of animals, which +corresponds to the presence of a beak in a bird, and +its absence in living reptiles. It is not known whether +the extremity of the Pterodactyle's beak is a single +bone, the intermaxillary bone, such as forms the +corresponding toothless part of the jaw in the South +African reptile Dicynodon, or whether it is made +by the pair of bones called premaxillaries which +form the extremity of the jaw in most Dinosaurs. +Too much importance may perhaps be attached +to such differences which are partly hypothetical, +because the extinct Ichthyosaurus, which has an exceptionally +long snout, has the two premaxillary +bones elongated so as to extend backward to the +nostrils. A similar elongation of those bones is seen +in Porpoises, which also have a long snout; and the +bones are carried back from the front of the head to +the nostrils, which are sometimes known as blowholes. +But the Porpoise has those premaxillary +bones not so much in advance of the bones which +carry teeth named maxillary, as placed in the interspace +between them. The nostrils, however, are not +limited to the extremity of the head in all Dinosaurs. +If this region of the beak in Dimorphodon be compared +<span class='pagenum'><a name="Page_201" id="Page_201">[Pg 201]</a></span> +with the corresponding part of a Dinosaur +from the Permian rocks, or Trias, the relation of the +nostril to the bones forming the beak may be better +understood.</p> + + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_78" id="Fig_78"></a> +<span class="caption">FIG. 78. COMPARISON OF THE SKULL OF THE +DINOSAUR ORNITHOSUCHUS WITH THE ORNITHOSAUR DIMORPHODON</span> +<img src="images/i_236.jpg" width="640" height="470" alt="FIG. 78." title="FIG. 78." /> +</div> + +<p>In the sandstone of Elgin, usually named Trias, a +small Dinosaur is found, which has been named Ornithosuchus, +from the resemblance of its head to that +of a Bird. Seen from above, the head has a remarkable +resemblance to the condition in Rhamphorhynchus, +in the sharp-pointed beak and positions +of the orbits and other openings. In side view the +orbits have the triangular form seen in Dimorphodon, +and the preorbital vacuities are large, as in that genus, +while the lateral nostrils, which are smaller, are further +forward in the Dinosaur. The differences from Dimorphodon +are in the articulation for the jaw being +carried a little backward, instead of being vertical as +in the Pterodactyle, and the bone in front of the +nose is smaller. Notwithstanding probable differences<span class='pagenum'><a name="Page_202" id="Page_202">[Pg 202]</a></span> +in the palate, the approximation, which extends to +the Crocodile-like vacuity in the lower jaw, is such +that by slight modification in the skull the differences +would be substantially obliterated by which the skull +of such an Ornithosaur is technically distinguished +from such a Dinosaur.</p> + +<p>The back of the skull is clearly seen in the Whitby +Pterodactyle, and its structure is similar to the corresponding +part of such Dinosaurs as Anchisaurus or +Atlantosaurus, without the resemblance quite amounting +to identity, but still far closer than is the resemblance +between the same region in the heads of +Crocodiles, Lizards, Serpents, Chelonians. Few of +these fossil Dinosaur skulls are available for comparison, +and those differ among themselves. The +coincidences rather suggest a close collateral relation +than prove the elaboration of one type from the +other. They may have had a common ancestor.</p> + +<p>The Trias rocks near Stuttgart have yielded Dinosaurs +as unlike Pterodactyles as could be imagined, +resembling heavily armoured Crocodiles, in such types +as the genus Belodon. Its jaws are compressed from +side to side, as in many Pterodactyles, and the nostrils +are at least as far backward as in Rhamphorhynchus. +Belodon has preorbital vacuities and postorbital vacuities, +but the orbit of the eye is never large, as in +Pterodactyles. It might not be worth while dwelling +on such points in the skull if it were not that the +pelvis in Belodon is a basin formed by the blending +of the expanded plates of the ischium and the pubis, +into a sheet of bone which more nearly resembles +the same region in Pterodactyles than does the +ischio-pubic region in other Dinosaurian animals +like Cetiosaurus.<span class='pagenum'><a name="Page_203" id="Page_203">[Pg 203]</a></span></p> + +<p>The backbone in a few Dinosaurs is suggestive of +Pterodactyles. In such genera as have been named +Cœlurus and Calamospondylus, in which the skeleton +is only partially known, the neck vertebræ become +elongated, so as to compare with the long-necked +Pterodactyles. The cervical rib is often very similar +to that type, and blended with the vertebra, as in +Pterodactyles and Birds. The early dorsal vertebræ +of Pterodactyles might almost be mistaken for those +of Dinosaurs. The tail vertebræ of a Pterodactyle +are usually longer than in long-tailed Dinosauria.</p> + +<p>In the limbs and the bony girdles which support +them there is more resemblance between Pterodactyles +and Dinosaurs than might have been anticipated, +considering their manifest differences in +habit. Thus all Dinosaurs have the hip bone named +ilium prolonged in front of the articulation for the +femur as well as behind it, almost exactly as in +Pterodactyles and Birds (see <a href="#Page_95">p. 95</a>). There is some +difference in the pubis and ischium which is more +conspicuous in form than in direction of the bones. +There is a Pterodactyle imperfectly preserved, named +<i>Pterodactylus dubius</i>, in which the ischium is directed +backward and the pubis downward, and the bones +unite below the acetabular cavity for the head of the +femur to work in, but do not appear to be otherwise +connected. In Rhamphorhynchus the connexion +between these two thickened bars is made by a thin +plate of bone. In such a Dinosaur as the American +carnivorous Ceratosaurus the two bars of the pubis +and ischium remain separate and diverging, and +there is no film of bone extending over the interspace +between them. The development of such a +bony condition would make a close approximation +<span class='pagenum'><a name="Page_204" id="Page_204">[Pg 204]</a></span> +between the Ornithosaurian pelvis and that of those +Dinosaurs which closely resemble Pterodactyles in +skull and teeth.</p> + + +<div class="figcenter" style="width: 492px;"> +<a name="Fig_79" id="Fig_79"></a> +<span class="caption">FIG. 79. LEFT SIDE OF PELVIS</span> +<img src="images/i_239.jpg" width="492" height="480" alt="FIG. 79." title="FIG. 79." /> +<p class="center">A Pterodactyle is shown between a carnivorous Dinosaur above and +a herbivorous Dinosaur below</p> +</div> + + +<p>Another pelvic character of some interest is the +blending of the pubis and ischium of the right and +left sides in the middle line of the body. There are +some genera of Dinosaurs like the English Aristosuchus +from the Weald, and the American genera +Cœlurus, Ceratosaurus, and others, in which the +pubic bones, instead of uniting at their extremities, +are pinched together from side to side, and unite +down the lower part of their length, terminating +in an expanded end like a shoe, which is seen to be +a separate ossification, and probably formed by a pair +<span class='pagenum'><a name="Page_205" id="Page_205">[Pg 205]</a></span> +of ossifications joined in the median line. This small +bone, which is below the pubes, and in these animals +becomes blended with them, we may regard as a pair +of prepubic bones like those of Pterodactyles and +Crocodiles, except that they have lost the stalk-like +portions, which in those animals are developed to +compensate for the diminished length of the pubic +bones. The prepubic bones may also be developed +in Iguanodon, in which a pair of bones of similar +form remains throughout life in advance of the +pubes, as in Pterodactyles. In those Dinosauria +with the Bird-like type of pelvis the pubic bone +is exceptionally developed, sending one process +backward and another process forward, so that +there is a great gap between these diverging limbs +to the bone. In the region behind the sternum to +which the ribs were attached, and in front of the +pelvis, is a pair of bones in Iguanodon shaped like +the prepubic bones of Dimorphodon. They have +sometimes been interpreted as a hinder part of the +sternum, but may more probably be regarded as a +pair of prepubic bones articulating each with the +anterior process of the pubis (see <a href="#Fig_80">Fig. 80</a>). The small +bones found at the extremities of the pubes in such +carnivorous Dinosaurs as Aristosuchus are blended +by bony union with the pubes. The bones in Iguanodon +are placed behind the sternal region without +any attachment for sternal ribs, and the expanded +processes converge forwards from the stalk and unite +exactly like the prepubic bones of Ornithosaurs. +While this character, on the one hand, may link +Pterodactyles with the Dinosaurs, on the other hand +it may be a link between both those groups and the +Crocodiles, in which the front pair of bones of the +<span class='pagenum'><a name="Page_206" id="Page_206">[Pg 206]</a></span> +pelvis has also appeared to be representative of the +prepubic bones of Flying Reptiles (see <a href="#Fig_32">Fig. 32, p. 98</a>).</p> + +<div class="figcenter" style="width: 640px;"> +<a name="Fig_80" id="Fig_80"></a> +<span class="caption">FIG. 80. DIAGRAM OF THE PELVIS SEEN FROM BELOW IN +AN ORNITHOSAUR AND A DINOSAUR</span> +<img src="images/i_241.jpg" width="640" height="441" alt="FIG. 80." title="FIG. 80." /> +</div> + +<p>The resemblances between Pterodactyles and Dinosaurs +in the hind limb are not of less interest, though +it is rather in the older Pterodactyles such as Dimorphodon, +Pterodactylus, and Rhamphorhynchus that +the resemblance is closest with the slender carnivorous +Dinosaurs. They never have the head of +the thigh bone, femur, separated from its shaft by a +constricted neck, as in the Pterodactyles from the +Chalk. In many ways the thigh bone of Dinosaurs +tends towards being Avian; while that of Pterodactyles +inclines towards being Mammalian, but with a +tendency to be Bird-like in the older types, and to be +Mammal-like in the most recent representatives of +the group in the Chalk.</p> + +<p>The bones of the leg in Ornithosaurs, known as +tibia and fibula, are remarkable for the circumstance +first that they resemble Birds in the fibula being slender +<span class='pagenum'><a name="Page_207" id="Page_207">[Pg 207]</a></span> +and only developed in its upper part towards the +femur, and secondly that in a genus like Dimorphodon +this drum-stick bone has the two upper bones of +the ankle blended with the tibia, so as to form a +rounded pulley joint which is indistinguishable from +that of a Bird (see <a href="#Page_102">p. 102</a>). There is a large number +of Dinosaurs in which this remarkable distinctive +character of Birds is also found. Only, Dinosaurs +like Iguanodon, for instance, have the slender fibula +as long as the tibia, and contributing to unite with the +separate ankle bones of the similarly rounded pulley +at the lower end. There are no Birds in which the +tarsal bones remain separated and distinct throughout +life. But in Pterodactylus from Solenhofen, as +in a number of Dinosaurs, especially the carnivorous +genera, the bones of the tarsus remain distinct throughout +life, and never acquired such forms as would have +enabled the ankle bone, termed astragalus, to embrace +the extremity of the tibia, as it does in Iguanodon. +Thus the resemblance of the Ornithosaur drum-stick +is almost as close to Dinosaurs as to Birds.</p> + +<p>There is great similarity between Dinosaurs and +Pterodactyles seen in the region of the instep, known +as the metatarsus. These bones are usually four in +number, parallel to each other, and similar in form. +They are commonly longer than in Dinosaurs; but +among some of the carnivorous Dinosaurs their +length approximates to that seen in Pterodactyles. +In neither group are the bones blended together by +bony union, while they are always united in Birds, as +in Oxen and similar even-hoofed mammals. Dinosaurs +agree with Pterodactyles in maintaining the metatarsal +bones separate, but they differ from them and agree +with Birds frequently, in having the number of meta<span class='pagenum'><a name="Page_208" id="Page_208">[Pg 208]</a></span>tarsal +bones reduced to three, as in Iguanodon, though +Dinosaurs often have as many as five digits developed.</p> + +<p>The toe bones, the phalanges of these digits of the +hind limb, are usually longer in Pterodactyles than in +Dinosaurs, but they resemble carnivorous Dinosaurs +in the forms of their sharp terminal bones for the claws, +which are similarly compressed from side to side.</p> + +<p>So diverse are the functions of the fore limb in +Dinosaurs and Pterodactyles, and so remarkably does +the length of the metacarpal region of the back of the +hand vary in the long-tailed and short-tailed Ornithosaurs, +that there is necessarily a less close correspondence +in that region of the skeleton between these two +groups of animals; for the Pterodactyle fore limb is +modified in relation to a function which can only be +paralleled among Birds and Bats; and yet neither +of those groups of animals approximates closely in +this region of the skeleton to the Flying Reptile. +Under all the modifications of structure which may +be attributed to differences of function, some resemblance +to Dinosaurs may be detected, which is +best evident in the upper arm bone, humerus; is +slight in the fore-arm bones, ulna and radius; and +becomes lost towards the extremity of the limb.</p> + +<p>If the tendency of the thigh bone to resemble a +Mammalian type of femur (<a href="#Page_100">p. 100</a>) is a fundamental, +deep-seated character of the skeleton, it might be anticipated +that a trace of Mammalian character would +also be found in the humerus. For what the character +is worth, the head of the humerus does show a closer +approximation to a Monotreme Mammal than is seen +in Birds, and is to some extent paralleled in those +South African reptiles which approximate to Mammals +most closely. Not the least remarkable of the many<span class='pagenum'><a name="Page_209" id="Page_209">[Pg 209]</a></span> +astonishing resemblances of these light aerial creatures +to the more heavy bodied Dinosaurs is the circumstance +that the humerus in both groups makes a not +dissimilar approach to that of certain Mammals.</p> + +<p>These illustrations may be accepted as demonstrating +a relationship between the Ornithosaurs and +Dinosaurs now compared, which can only be explained +as results of influence of a common parentage +upon the forms of the bones. But more interesting +than resemblances of that kind is the similarity that +may be traced in the way in which air is introduced +into cavities in the bones in both groups. In some +of the imperfectly known Dinosaurs, like Aristosuchus, +Cœlurus, and Thecospondylus, the bone texture is as +thin as in Pterodactyles, and the vertebræ are excavated +by pneumatic cavities, which are amazing in +size when compared with the corresponding structures +in birds, for the vertebra is often hollowed out so that +nothing remains but a thin external film like paper +for its thickness. In the Dinosaurian genus Cœlurus +this condition is as well marked in the tail and back +as it is in the neck. The essential difference from +Birds appears to be that in the larger carnivorous +Dinosaurs the pneumatic condition of the bones is +confined to the vertebral column; while Birds and +Pterodactyles have the pneumatic condition more +conspicuously developed in the limb bones. The +pneumatic skeleton, however, appears to be absent +from the herbivorous types like Iguanodon and all +Dinosaurs which have the Bird-like form of pelvis, +and are most Bird-like in the forms of bones of the +hind limb. It is possible that some of the carnivorous +Dinosaurs also possessed limb bones with pneumatic +cavities. Many of those bones are hollow with very<span class='pagenum'><a name="Page_210" id="Page_210">[Pg 210]</a></span> +thin walls. If their cavities were connected with the +lungs the foramina are inconspicuous and unlike the +immense holes seen in the sides of the vertebræ.</p> + +<p>According to the late Professor Marsh, the limbs +of Cœlurus and its allies, which at present are imperfectly +known, are in some cases pneumatic. Therefore +there is a closer fundamental resemblance between +some carnivorous Dinosaurs and Pterodactyles +than might have been anticipated. But the skull of +Cœlurus is unknown, and the fragments of the +skeleton hitherto published are insufficient to do +more than show that the two types were near in +kindred, though distinct in habit. Each has elaborated +a skeleton which owes much to the common +stock which transmitted the vital organs, and the tendency +of the bones to take special forms; but which +also owes more than can be accurately measured to +the action of muscles in shaping the bones and the +influence of the mechanical conditions of daily life +upon the growth of the bones in both of these orders +of animals. Enough is known to prove that all Dinosaurs +cannot be regarded as Ornithosaurs which have +not acquired the power of flight; though the evidence +would lead us to believe that the primitive Ornithosaur +was a four-footed animal, before the wing finger +became developed in the fore limb as a means of +extending a patagial membrane, like the membrane +which in the hind limb of Dimorphodon has bent the +outermost digit of the foot upward and outward to +support the corresponding organ of flight extending +down the hind legs.</p> + +<p>It may thus be seen that the characters of Ornithosaurs +which have already been spoken of as Reptilian, +as distinguished from the resemblances to Birds, may<span class='pagenum'><a name="Page_211" id="Page_211">[Pg 211]</a></span> +now with more accuracy be regarded as Dinosaurian. +The Dinosaurs, like Pterodactyles, must be regarded +as intermediate in some respects between +Reptiles and Birds. The resemblances enumerated +would alone constitute a partial transition from the +Reptile to the Bird, although no Dinosaurs have +organs of flight; many are heavily armoured with +plates of bone, and few, if any, approximate in the +technical parts of the skeleton to the Bird class, +except in the hind limbs. Yet Dinosaurs have +sometimes been regarded as standing to Birds in +the relation of ancestors, or as parallel to an +ancestral stock.</p> + +<p>Before an attempt can be made to estimate the +mutual relation of the Flying Reptiles to Dinosaurs +on the one hand, and to Birds on the other, it may +be well to remember that the resemblance of such +a Dinosaur as Iguanodon to a Bird in its pelvis and +hind limb is not more remarkable than that of +Pterodactyles to Birds in the shoulder-girdle and +bones of the fore limb. The keeled sternum, the +long, slender coracoid bones and scapulæ, are absolutely +Bird-like in most Ornithosaurs; and that region +of the skeleton only differs from Birds in the absence +of a furculum which represents the clavicles, and is +commonly named the "merry-thought." The elongated +bones of the fore-arm and the hand, terminating +in three sharp claws, are characters in which the +fossil bird Archæopteryx resembles the Pterodactyle +Rhamphorhynchus, a resemblance which extends to +a similar elongation of the tail. It is remarkable +that the resemblance should be so close, since Archæopteryx +affords the only bird's skeleton known to be +contemporary which can be compared with the Solen<span class='pagenum'><a name="Page_212" id="Page_212">[Pg 212]</a></span>hofen +Flying Reptiles. The resemblance may possibly +be closer than has been imagined. The back of the +head of Archæopteryx is imperfectly preserved in +the region of the quadrate bone, malar arch, and +temporal vacuity. And till these are better known +it cannot be affirmed that the back of the head is +more Reptilian in Pterodactyles than in the oldest +Birds. The side of the head in Archæopteryx is +distinguished by the nostril being far forward, the +vacuity in front of the orbit being as large as in +the Pterodactyle Scaphognathus from Solenhofen +and other long-tailed Pterodactyles.</p> + + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_213" id="Page_213">[Pg 213]</a></span></p> +<h2><a name="CHAPTER_XVIII" id="CHAPTER_XVIII"></a><small>CHAPTER XVIII</small><br /><br /> + +HOW PTERODACTYLES MAY +HAVE ORIGINATED</h2> + + +<p>Ornithosauria have many characters inseparably +blended together which are otherwise +distinctive of Reptiles, Birds, and Mammals, and associated +with peculiar structures which are absent +from all other animals. They are not quite alone in +this incongruous combination of different types of +animals in the same skeleton. Dinosaurs, which were +contemporary with Ornithosaurs, approximate to them +in blending characters of Birds with the structure of +a Reptile and something of a Mammal in one animal. +If an Ornithosaur is Reptilian in its backbone, in the +articular ends of each vertebra having the cup in +front and ball behind in the manner of Crocodiles, +Serpents, and many Lizards, a Dinosaur like Iguanodon, +which had the reversed condition of ball in +front and cup behind in its early vertebræ, may be +more Mammalian than Avian in a corresponding +resemblance of the bones to the neck in hoofed +Mammals. But while Pterodactyles are sometimes +Mammalian in having the head of the thigh bone +moulded as in carnivorous Mammals and Man, the +corresponding bone in a Dinosaur is more like that<span class='pagenum'><a name="Page_214" id="Page_214">[Pg 214]</a></span> +of a Bird. And while the Pterodactyle shoulder-girdle +is often absolutely Bird-like, that region in +Dinosaurs can only be paralleled among Reptiles.</p> + +<p>Such combinations of diverse characters are not +limited to animals which are extinct. There were +not wanting scientific men who regarded the Platypus +of Australia, when first sent to Europe, as an +ingenious example of Eastern skill, in which an +animal had been compounded artificially by blending +the beak of a Bird with the body of a Mammal. +Fuller knowledge of that remarkable animal has +continuously intensified wonder at its combination +of Mammal, Bird, and Reptile in a single animal. +It has broken down the theoretical divisions between +the higher Vertebrata, demonstrating that a +Mammal may lay eggs like a Reptile or Bird, that +the skull may include the reptilian characters of the +malar arch and pre-frontal and post-frontal bones, +otherwise unknown in Mammals and Birds. The +groups of Mammals, Birds, and Reptiles now surviving +on the earth prove to be less sharply defined +from each other when the living and extinct types +are considered together. But in Pterodactyles, +Mammal Bird and Reptile lose their identity, as three +colours would do when unequally mixed together.</p> + +<p>This mingling of characteristics of different animals +is not to be attributed to interbreeding, but is the +converse of the combination of characters found in +hybrid animals. It is no exaggeration to say that +there is a sense in which Mammal, Bird, Reptile, and +the distinctive structures of the Ornithosaur, have +simultaneously developed from one egg, in the body +of one animal.</p> + +<p>The differences between those vertebrate types of<span class='pagenum'><a name="Page_215" id="Page_215">[Pg 215]</a></span> +animals consist chiefly in the way in which their +organisation is modified, by one strain of characters +being eliminated so that another becomes predominant, +while a distinctive set of structures is elaborated +in each class of animals. The earlier geological history +of the higher Vertebrata is very imperfectly +known, but the evidence tends to the inference that +the older representatives of the several classes approximate +to each other more closely than do their +surviving representatives, so that in still earlier ages +of time the distinction between them had not become +recognisable. The relation of the great groups of +animals to each other, among Vertebrata, is essentially +a parallel relation, like the colours of the solar spectrum, +or the parallel digits of the hand. It was +natural, when only the surviving life on the earth was +known, to imagine that animals were connected in a +continuous chain by successive descent, but Mammals +have given no evidence of approximation to Birds; +and Birds discover no evidence that their ancestors +were Reptiles, in the sense in which that word is used +to define animals which now exist on the earth. +When the variation which animals attain in their +maturity and exhibit in development from the egg +was first realised, it was imagined that Nature, by +slow summing up and accumulation of differences +which were observed, would so modify one animal +type that it would pass into another. There is little +evidence to support belief that the changes between +the types of life have been wrought in that way. +The history of fossil animals has not shown transitions +of this kind from the lower to higher Vertebrata, +but only intermediate, parallel groups of animals, +analogous to those which survive, and distinct from<span class='pagenum'><a name="Page_216" id="Page_216">[Pg 216]</a></span> +them in the same way as surviving groups are distinct +from each other. The circumstance that Mammals, +Birds, and Reptiles are all known low down in the +Secondary epoch of geological time, is favourable to +the idea of their history being parallel rather than +successive. Such a conception is supported by the +theory of elimination of characters from groups of +animals as the basis of their differentiation. This loss +appears always to be accompanied by a corresponding +gain of characters, which is more remarkable in +the soft, vital organs than in the skeleton. The gain +in higher Vertebrates in the bones is chiefly in the +perfection of joints at their extremities; but the gain +in brain, lungs, heart, and other soft parts is an +elaboration of those structures and an increase in +amount of tissue.</p> + +<p>The resemblances of Ornithosaurs to Mammals are +the least conspicuous of their characters. Those seen +in the upper arm bone and thigh bone are manifestly +not derived from Mammals. They cannot be explained +as adaptations of the bones to conditions of +existence, because there is no community of habit to +be inferred between Pterodactyles and Mammals, in +which the bones are in any way comparable.</p> + +<p>Other fossil animals show that a fundamentally +Reptilian structure is capable of developing in the +Mammalian direction in the skull, backbone, shoulder-girdle, +hip-girdle, and limbs, so as to be uniformly +Mammalian in its tendencies. This is proved by +tracing the North American Texas fossils named +Labyrinthodonts, through the South African Theriodonts, +towards the Monotremata and other Mammalia. +Just as those animals have obliterated all traces of +the Bird from their skeletons, Birds have obliterated<span class='pagenum'><a name="Page_217" id="Page_217">[Pg 217]</a></span> +the distinctive characters of Mammals. The Ornithosaur +has partially obliterated both. With a skull and +backbone marked by typical characters of the Reptile, +it combines the shoulder-girdle and hip-girdle of a +Bird, with characters in the limbs which suggest both +those types in combination with Mammals.</p> + +<p>The bones have been compared in the skeleton of +each order of existing Reptiles, and found to show +side by side with their peculiar characters not only +resemblances to the other Reptilia, but an appreciable +number of Mammalian and Avian characters in their +skeletons. The term "crocodile," for example, indicates +an animal in which the skeleton is dominated +by one set of peculiar characters. Crocodiles retain +enough of the characteristics of several other orders +of reptiles to show that an animal sprung from the +old Crocodile stock might diverge widely from existing +Crocodiles by intensifying what might be termed +its dormant characters in the Crocodile skeleton. +Comparing animals together bone by bone it is +possible to value the modifications of form which +they put on, and the resemblances between them, +so as to separate the inherited wealth of an animal's +affinities with ancestors or collateral groups, from +the peculiar characters which have been acquired +as an increase based upon its typical bony possessions +or osteological capital. There is no part of the Pterodactyle +skeleton which is more distinctly modified +than the head of the upper arm bone, which fits +into the socket between the coracoid bone and the +shoulder-blade. The head of the humerus, as the +articular part is named, is somewhat crescent-shaped, +convex on its inner border, and a little concave on +its outer border, and therefore unlike the ball-shaped<span class='pagenum'><a name="Page_218" id="Page_218">[Pg 218]</a></span> +head of the upper arm bone in Man and the higher +Mammals. It is much more nearly paralleled in the +little group of Monotremata allied to the living +Ornithorhynchus. In that sense the head of the +humerus in a Pterodactyle has some affinity with the +lowest Mammalia, which approach nearest to Reptiles. +The character might pass unregarded if it were not +found in more striking development in fossil Reptiles +from Cape Colony, which from having teeth like +Mammals are named Theriodontia. In several of +those South African reptiles the upper arm bone +approaches closer to the humerus in Ornithosaurs +than to Ornithorhynchus. Such coincidences of +structure are sometimes dismissed from consideration +and placed beyond investigation by being termed +adaptive modifications; but there can be no hope +of finding community of habit between the burrowing +Monotreme, the short-limbed Theriodont, and +the flying Pterodactyle which might have caused +this articular part of the upper arm bone to acquire +a form so similar in animals constructed so differently. +If the resemblance in the humerus to Monotremes +in this respect is not to be attributed to +burrowing, neither can the crescent form of its upper +articulation be attributed to flight; for in Birds the +head of the bone is compressed, but always convex, +and Bats fly without any approach to the Pterodactyle +form in the head of the humerus. This +apparently trivial character may from such comparisons +be inferred to be something which the way +of life of the animal does not sufficiently account for. +These deepest-seated parts of the limbs are slow to +adapt themselves to changing circumstances of existence, +and retain their characters with moderate<span class='pagenum'><a name="Page_219" id="Page_219">[Pg 219]</a></span> +variation of the bones in each of the orders or classes +of animals. It therefore is safer to regard Mammalian +characters, as well as the resemblances which +Pterodactyles show to other kinds of animals, as due +to inheritance from a time when there was a common +stock from which none of these animals which have +been considered had been distinctly elaborated.</p> + +<p>A few characters of Ornithosaurs are regarded as +having been acquired, because they are not found +in any other animals, or have been developed only +in a portion of the group. The most obvious of +these is the elongated wing finger; but in some +genera, like Dimorphodon, there is also a less elongation +of the fifth digit of the foot, and perhaps in +all genera there is a backward development of the +first digit of the hand, which is without a claw, and +therefore unlike the clawed digit of a Bat. An +acquired character of another kind, which is limited +to the Cretaceous genera, is seen in the shoulder-blade +being directed transversely outward, so that +its truncated end articulates by a true joint with the +early vertebræ of the back, and defended the cavity +inclosed by the ribs by a strong bony external arch. +And finally, as the animals later in time acquire short +tails, and relatively longer limbs, the bones of the +back of the hand, termed metacarpals, acquire +greater and distinctive length, which is not seen in +the long-tailed types like Rhamphorhynchus.</p> + +<p>These and such-like acquired characters distinguish +the class of animals from all groups with +which it may be compared, and mark the possible +limits of variation of the skeleton within the +boundary of the order. But no further variation of +these parts of the skeleton could make a transition<span class='pagenum'><a name="Page_220" id="Page_220">[Pg 220]</a></span> +to another order of animals, or explain how the +Pterodactyles came into existence, because the characters +which separate orders and classes of animals +from each other differ in kind from those which +separate smaller groups, named genera and species, +of which the order is made up. The accumulation +of the characters of genera will not sum up into the +characters of an order or class.</p> + +<p>In making the division of Vertebrate animals into +classes the skeleton is often almost ignored. Its +value is entirely empirical and based upon the +observed association of the various forms of bones +with the more important characters of the brain and +other vital organs. What is understood as a Mammalian +or Avian character in the skeleton is the form +of bone which is found in association with the soft +vital organs which constitute an animal a Mammal +or a Bird.</p> + +<p>The characters which theoretically define a Mammal +appear to be the enormous overgrowth of the cerebral +hemispheres of the brain by which the cerebrum +comes into contact with the cerebellum, as among +Birds. This character distinguishes both groups of +animals from all Reptiles, recent and fossil. But in +examining the mould of the interior of the brain +case it is rare to have the bones fitting so closely +to the brain as to prove that the lateral expansion +below the cerebrum and cerebellum is formed by +the optic lobes of the brain. Otherwise the brain +of a Pterodactyle might be as like to the brain of +Ornithorhynchus as it is like that of a Bird (<a href="#Fig_19">Fig. 19</a>). +But it is precisely in this condition of arrangement +of the parts of the brain that the specimens appear +to be most clear. The lateral mass of brain in<span class='pagenum'><a name="Page_221" id="Page_221">[Pg 221]</a></span> +specimens of Ornithosaurs from the Lower Secondary +rocks appears to be transversely divided into back +and front parts, which may be thought to correspond +to the structures in a Mammal brain named +<i>corpora quadrigemina</i>, but to be placed as the optic +lobes are placed in Birds, and to have relatively +greater dimensions than in Mammals. No evidence +has been observed of this transverse division of the +optic lobes of the brain in Pterodactyles from the +Chalk and Cretaceous rocks, and so far as the evidence +goes this part of the brain was shaped as in birds, +but rather smaller.</p> + +<p>The brain is the only soft organ in which a Mammalian +character could be evidenced. The uniformity +in character of the brain throughout the group in +Mammals is remarkable, in reference to the circumstance +that the reproduction varies in type; the lowest, +or Monotreme division, being oviparous. If there is +no necessary connexion between the Mammalian +brain and the prevalent condition under which the +young are produced alive, it may be affirmed also +that there is no necessary connexion between the +form of the brain and the form of the bones, since +the brain cavity in Theriodont reptiles shows no +resemblance to that of a Mammal, while the bones +are in so many respects only paralleled among +Monotremata and Mammalia. The variety of forms +which the existing Mammalian orders of animals +assume, shows the astonishing range of structure of +the skeleton which may coexist with the Mammalian +brain. And therefore we are led to the conclusion +that any other fundamental modification of brain—such +as distinguishes the class of Birds—might also +be associated with forms and structures of the skele<span class='pagenum'><a name="Page_222" id="Page_222">[Pg 222]</a></span>ton +which would vary in similar ways. In other +words, if for convenience we define a Mammal by +its form of brain, structure of the heart and lungs, +and provision for nutrition of the young, without +regard to the covering of the skin, which varies +between the scales of a pangolin and the practically +naked skin of the whale—a bird might be also +defined by its peculiar conditions of brain and lungs, +without reference to the feathered condition of the +skin, though the feathered condition extends backward +in time to the Upper Secondary rocks, as seen +in the Archæopteryx.</p> + +<p>The Avian characters of Pterodactyles are the predominant +parts of their organisation, for the conditions +of the brain and lungs shown by the moulds +of the brain case and the thin hollow bones with +conspicuous pneumatic foramina, give evidence of +a community of vital structures with Birds, which +is supported by characters of the skeleton. If any +classificational value can be associated with the distribution +of the pneumatic foramina as tending to +establish membership of the same class for animals +fashioned on the same plan of soft organs, the +evidence is not weakened when a community of +structures is found to extend among the bones to +such distinctive parts of the skeleton as the sternum, +shoulder-girdle, bones of the fore-arm and fore-leg; +for in all these regions the Pterodactyle bones are +practically indistinguishable from those of Birds. +This is the more remarkable because other parts of +the skeleton, such as the humerus and pelvis, show +a partial resemblance to Birds, while the parts which +are least Avian, like the neck bones, have no tendency +to vary the number of the vertebræ, in the<span class='pagenum'><a name="Page_223" id="Page_223">[Pg 223]</a></span> +way which is common among Birds, following more +closely the formula of the seven cervical vertebræ of +Mammals.</p> + +<p>It would therefore appear from the vital community +of structures with Birds, that Pterodactyles +and Birds are two parallel groups, which may be +regarded as ancient divergent forks of the same +branch of animal life, which became distinguished +from each other by acquiring the different condition +of the skin, and the structures which were developed +in consequence of the bony skeleton ministering to +flight in different ways; and with different habit of +terrestrial progression, this extinct group of animals +acquired some modifications of the skeleton which +Birds have not shown. There is nothing to suggest +that Pterodactyles are a branch from Birds, but their +relation to Birds is much closer, so far as the skeleton +goes, than is their relation with the flightless Dinosaurs, +with which Birds and Pterodactyles have many +characters in common.</p> + +<p>On the theory of elimination of character which +I have used to account for the disappearance of some +Mammalian characters from the Pterodactyle, that +loss is seen chiefly in the removal of the parts which +have left a Reptilian articulation of the lower jaw +with the skull, and the articulation of the vertebræ +throughout the vertebral column by a modified cup-and-ball +form of joint. The furculum of the Bird is +always absent from the Pterodactyle. No specimen +has shown recognisable clavicles or collar-bones. +Judged by the standard of existing life, Pterodactyles +belong to the same group as Birds, on the +evidence of brain and lungs, but they belong to +a different group on account of the dissimilar<span class='pagenum'><a name="Page_224" id="Page_224">[Pg 224]</a></span> +modifications of the skeleton and apparent absence +of feathers from the skin.</p> + +<p>The most impressive facts in the Pterodactyle +skeleton, in view of these affinities, are the structures +which it has in common with Reptiles. Some structures +are fundamental, like the cup-and-ball articulation +of the vertebræ, which is never found in birds +or mammals. Although not quite identical with the +condition in any Reptile, this structure is approximately +Lizard-like or Crocodile-like in the cup-and-ball +character. It shows that the deepest-seated part +of the skeleton is Reptile-like, though it may not be +more Reptilian than is the vertebral column of a +Mammal, if comparison is made between Mammals +and extinct groups of animals known as Reptiles, +such as Dinosaurs and Theriodontia.</p> + +<p>The orders of animals which have been included +under the name Reptilia comprise such different +structural conditions of the parts of the skeleton +which may be termed reptilian in Ornithosaurs, that +there is good reason for regarding the cup-and-ball +articulation as quite a distinctive Reptilian specialisation, +in the same sense that the saddle-shaped articulation +between the bodies of adjacent vertebræ in +a bird is an Avian specialisation. From the theoretical +point of view the Ornithosaur acquired its Reptilian +characters simultaneously with its Avian and Mammalian +characters.</p> + +<p>There is nothing in the structure of the skeleton +of the Dinosauria, to which Ornithosaurs approximate +in several parts of the body, which would help to +explain the cup-and-ball articulation of the backbone, +if the Flying Reptile were supposed to be an offshoot +from the carnivorous Dinosaurs.<span class='pagenum'><a name="Page_225" id="Page_225">[Pg 225]</a></span></p> + +<p>The elimination of Reptile characters from so much +of the skeleton, and the substitution for them of the +characters of Birds and Mammals, would be of exceptional +interest if there had been any ground for +regarding the flying animal as more nearly related to +a Reptile than to a Bird. But if the evidence from +the form of the brain and nature of the pneumatic +organs seen in the limb bones accounts for the Avian +features of the skeleton, the Reptilian condition of the +vertebral column helps to show a capacity for variation, +and that the fixity of type and structure, which +the skeleton of the modern Bird has attained, is not +necessarily limited to or associated with the vital +organs of Birds.</p> + +<p>The variation of the cup-and-ball articulation in +the neck of a Chelonian, which makes the third +vertebra cupped behind, the fourth bi-convex, the +fifth cupped in front, and the sixth flattened behind, +shows that too much importance may be attached +to the mode of union of these bones in Serpents, +Crocodiles, and those Lizards which have the cup in +front; for while in Lizards the anterior cup, oblique and +depressed, is found in most of its groups, the Geckos +show no trace of the cup-and-ball structure, and in +that respect resemble the Hatteria of New Zealand.</p> + +<p>If, therefore, the cup-and-ball articulation of vertebræ +in Ornithosauria has any significance as a mark +of affinity to Reptiles, it could only be in approximation +to those living Reptiles which possess the same +character, and would have it on the hypothesis that +both have preserved the structure by descent from an +earlier type of animal. This hypothesis is negatived +by the fact that the cup-and-ball articulation is unknown +in the older fossil Reptiles.<span class='pagenum'><a name="Page_226" id="Page_226">[Pg 226]</a></span></p> + +<p>Although the articulation for the lower jaw with +the skull in Ornithosaurs is only to be paralleled +among Reptiles, the structure is adapted to a brain +case which is practically indistinguishable from that +of a Bird, except for the postorbital arch.</p> + +<p>The hypothesis of descent, therefore, becomes impossible, +in any intelligible form, in explanation +of distinctive character of the skeleton. The hypothesis +of elimination may also seem to be insufficient, +unless the potential capacity for new development be +recognised as concurrent, and as capable of modifying +each region of the skeleton, or hard parts of the +animal, in the same way that the soft organs may be +modified. From which we infer that all structures, +which distinguish the several grades of organisation +in modern classifications, soft parts and hard parts +alike, may come into existence together, in so far +as they are compatible with each other, in any class +or ordinal division of animals.</p> + +<p>Although the young Mammal passes through a +stage of growth in which the brain may be said to be +Reptilian, there is no good ground for inferring that +Mammal or Bird type of skeleton was developed later +in time than that of Reptiles. The various types of +Fishes have the brains in general so similar to those +of Reptiles that it is more intelligible for all the +vertebrate forms of brain to have differentiated at +the same time, under the law of elimination of characters, +than that there should be any other bond of +union between the classes of animals.</p> + +<p>If we ask what started the Ornithosauria into +existence, and created the plan of construction of +that animal type, I think science is justified in boldly +affirming that the initial cause can only be sought<span class='pagenum'><a name="Page_227" id="Page_227">[Pg 227]</a></span> +under the development of patagial membranes, such +as have been seen in various animals ministering to +flight. Such membranes, in an animal which was +potentially a Bird in its vital organs, have owed development +to the absence of quill feathers. Thus +the wing membrane may be the cause for the chief +differences of the skeleton by which Ornithosaurs +are separated from Birds, for the stretch of wing in +one case is made by the skin attached to the bones, +and in the other case by feathers on the skin so +attached as to necessitate that the wing bones have +different proportions from Ornithosaurs.</p> + +<p>It is a well-known observation that each great +epoch of geological time has had its dominant forms +of animal life, which, so far as the earth's history is +known now, came into existence, lived their time, +and were seen no more. In the same way the +smaller groups of species and genera included in an +ordinal group of animals or class have abounded, +giving a tone to the life of each geological formation, +until the vitality of the animal is exhausted, and the +species becomes extinct or ceases to preponderate. +This process is seen to be still modifying the life on +the earth, when some kinds of animals and plants +are introduced to new conditions. Plants appear to +wage successful war more easily than animals. The introduction +of the Cactus in some parts of Cape Colony +has locally modified both the fauna and flora, just +as the Anacharis introduced into England spread +from Cambridge over the whole country, and became +for many years the predominant form of plant life +in the streams. The Rabbit in Australia is a historic +pest. Something similar to this physical fertility +and increase appears to take place under new cir<span class='pagenum'><a name="Page_228" id="Page_228">[Pg 228]</a></span>cumstances +in certain organs within the bodies of +animals, by the development of structures previously +unknown. A familiar example is seen in the internal +anatomy of the Trout introduced into New Zealand, +where the number of pyloric appendages about the +stomach has become rapidly augmented, while the +size and the form of the animal have changed. The +rapidity with which some of these changes have been +brought about would appear to show that Nature is +capable of transforming animals more rapidly than +might have been inferred from their uniform life +under ordinary circumstances. Growth of the vital +organs in this way may modify the distinctive form +of any vital organ, brain or lungs, and thus as a consequence +of modification of the internal structures due +to changes of food and habit, bring a new group of +animals into existence. And just as the group of +animals ceases to predominate after a time, so there +comes a limit to the continued internal development +of vital structures as their energy fails, for each organ +behaves to some extent like an independent organism.</p> + +<p>Under such explanations of the mutual relations of +the parts of animals, and groups of animals, time +ceases to be a factor of primary importance in their +construction or elaboration. The supposed necessity +for practically unlimited time to produce changes in +the vital organs which separate animals into great +orders or classes is a nightmare, born of hypothesis, +and may be profitably dismissed. The geological +evidence is too imperfect for dogmatism on speculative +questions; but the nature of the affinities of +Ornithosaurs to other animals has been established +on a basis of comparison which has no need of +theory to justify the facts. It is not improbable that<span class='pagenum'><a name="Page_229" id="Page_229">[Pg 229]</a></span> +the primary epoch of time, even as known at present, +may be sufficiently long to contain the parent races +from which Ornithosaurs and all their allies have +arisen.</p> + +<p>In thus stating the relation of Ornithosaurs to +other animals the Flying Reptile has been traced +home to kindred, though not to its actual parents or +birthplace. There is no geological history of the +rapid or gradual development of the wing finger, and +although the wing membrane may be accepted as +its cause of existence, the wing finger is powerfully +developed in the oldest known Pterodactyles as in +their latest representatives.</p> + +<p>Pterodactyles show singularly little variation in +structure in their geological history. We chronicle +the loss of the tail and loss of teeth. There is also +the loss of the outermost wing digit from the hind +foot as a supporter of the wing membrane. But the +other variations are in the length of the metacarpus, +or of the neck, or head. One of the fundamental +laws of life necessitates that when an animal type +ceases to adapt its organisation and modify its +structures to suit the altered circumstances forced +upon it by revolutions of the earth's surface its life's +history becomes broken. It must bend or break.</p> + +<p>The final disappearance of these animals from the +earth's history in the Chalk may yet be modified +by future discoveries, but the Flying Reptiles have +vanished, in the same way as so many other groups +of animals which were contemporary with them in +the Secondary period of time. Such extinctions +have been attributed to catastrophes, like the submergence +of land, so that the habitations of animals +became an area gradually decreasing in size, which<span class='pagenum'><a name="Page_230" id="Page_230">[Pg 230]</a></span> +at last disappeared. It appears also to be a law of +life, illustrated by many extinct groups of animals, +that they endure for geological ages, and having +fought their battle in life's history, grow old and unable +to continue the fight, and then disappear from +the earth, giving place to more vigorous types adapted +to live under new conditions.</p> + +<p>The extinct Pterodactyles hold a relation to Birds +in the scheme of life not unlike that which Monotremata +hold to other Mammals. Both are remarkable +for the variety of their affinities and resemblances +to Reptiles. The Ornithosauria have long passed +away; the Monotremes are nearing extinction. Both +appear to be supplanted by parallel groups which +were their contemporaries. Birds now fill the earth +in a way that Flying Reptiles never surpassed; but +their flight is made in a different manner, and the +wing is extended to support the animal in the air, +chiefly by appendages to the skin.</p> + +<p>If these fossils have taught that Ornithosaurs have +a community of soft vital organs with Dinosaurs and +Birds, they have also gone some way towards proving +that causes similar to those which determined the +structural peculiarities of their bony framework, +originated the special forms of respiratory organs +and brain which lifted them out of association with +existing Reptiles.</p> + +<p> </p> + +<p>These old flying animals sleep through geological +ages, not without honour, for the study of their story +has illuminated the mode of origin of animals which +survive them, and in cleaving the rocks to display +their bones we have opened a new page of the book +of life.</p> + + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_231" id="Page_231">[Pg 231]</a></span></p> +<h2><a name="APPENDIX" id="APPENDIX"></a>APPENDIX</h2> + + +<p>The best public collections of Ornithosaurian remains in +England are in the British Museum (Natural History); +Museum of Practical Geology, Royal College of Surgeons; +the University Museum, Oxford; Geological Museum, +Cambridge; and the Museum of the Philosophical Society +at York.</p> + +<p>Detailed descriptions and original figures of the principal +specimens mentioned or referred to may be found in +the following writings:—</p> + +<div class="blockquot"><p class="noidt"> +H. v. Meyer, <i>Reptilien aus dem Lithograph</i>. <i>Schiefer</i>. 1859. Folio.<br /> +v. Quenstedt, <i>Pterodactylus suevicus</i>. 1855. 4to.<br /> +Goldfuss, <i>Nova Acta Leopold</i>. XV.<br /> +v. Munster, <i>Nova Acta Leopold</i>. XV.<br /> +A. Wagner, <i>Abhandl. Bayerischen Akad.</i>, vi., viii.<br /> +Cuvier, <i>Annales du Museum</i>, xiii. 1809.<br /> +<span style="margin-left: 1em;">" <i>Ossemens fossiles</i>, v. 1824.</span><br /> +Buckland, <i>Geol. Trans.</i>, ser. 2, iii.<br /> +R. Owen, <i>Palæontographical Society</i>. 1851, 1859, 1860, 1870, 1874.<br /> +K. v. Zittel, <i>Palæontographica</i>, xxix. 1882.<br /> +T. C. Winkler, <i>Mus. Teyler Archives</i>. 1874, 1883.<br /> +Oscar Fraas, <i>Palæontographica</i>, xxv. 1878.<br /> +Anton Fritsch, <i>Böhm. Gesell. Sitzber</i>. 1881.<br /> +R. Lydekker, <i>Catalogue of Fossil Reptilia in British Museum</i>, I. 1888.<br /> +O. C. Marsh, <i>Amer. Jour. Science</i>. 1882, 1884.<br /> +S. W. Williston, <i>Kansas University Quarterly</i>. 1893, 1896.<br /> +<span class='pagenum'><a name="Page_232" id="Page_232">[Pg 232]</a></span>E. T. Newton, <i>Phil. Trans. Royal Soc.</i> 1888, 1894.<br /> +H. G. Seeley, <i>Ornithosauria</i>. 8vo. 1870.<br /> +<span style="margin-left: 2em;">" <i>Annals and Mag. Natural Hist.</i> 1870, 1871, 1890, 1891.</span><br /> +<span style="margin-left: 2em;">" <i>Linn. Society</i>. 1874, 1875.</span><br /> +<span style="margin-left: 2em;">" <i>Geol. Mag.</i> 1881.</span><br /> +Felix Pleininger, <i>Palæontographica</i>. 1894, 1901.<br /> +</p></div> + + + +<hr style="width: 65%;" /> +<p><span class='pagenum'><a name="Page_233" id="Page_233">[Pg 233]</a></span></p> +<h2><a name="INDEX" id="INDEX"></a>INDEX</h2> + + +<div class="blockquot"><p class="noidt"> +<b>A</b><br /> +<br /> +Abdominal ribs, <a href="#Page_85">85</a>, <a href="#Page_154">154</a><br /> +<br /> +Accumulation of characters, <a href="#Page_220">220</a><br /> +<br /> +Acetabulum, <a href="#Page_95">95</a><br /> +<br /> +Acquired characters, <a href="#Page_219">219</a><br /> +<br /> +Adjacent land, <a href="#Page_136">136</a><br /> +<br /> +Air cells, <a href="#Page_10">10</a>, <a href="#Page_48">48</a><br /> +<br /> +Albatross, <a href="#Page_23">23</a>, <a href="#Page_36">36</a>, <a href="#Page_176">176</a><br /> +<br /> +Alligator, brain, <a href="#Page_53">53</a>;<br /> +<span style="margin-left: 1em;">pelvis, <a href="#Page_98">98</a></span><br /> +<br /> +American Greensand, <a href="#Page_185">185</a><br /> +<br /> +— ornithosaurs, <a href="#Page_87">87</a>, <a href="#Page_126">126</a><br /> +<br /> +Amphibia, <a href="#Page_4">4</a>, <a href="#Page_191">191</a><br /> +<br /> +Anabas, <a href="#Page_17">17</a><br /> +<br /> +Anacharis, <a href="#Page_227">227</a><br /> +<br /> +Anchisaurus, <a href="#Page_199">199</a><br /> +<br /> +Angle of lower jaw, <a href="#Page_75">75</a><br /> +<br /> +Ankle bones, <a href="#Page_103">103</a>, <a href="#Page_195">195</a>, <a href="#Page_207">207</a><br /> +<br /> +Anomodonts, <a href="#Page_192">192</a><br /> +<br /> +Ant-eater of Africa, <a href="#Page_142">142</a>;<br /> +<span style="margin-left: 1em;">India, <a href="#Page_40">40</a>;</span><br /> +<span style="margin-left: 1em;">South America, <a href="#Page_40">40</a>, <a href="#Page_185">185</a></span><br /> +<br /> +Apteryx, lungs, <a href="#Page_48">48</a>;<br /> +<span style="margin-left: 1em;">pelvis, <a href="#Page_95">95</a></span><br /> +<br /> +Aquatic mammals, <a href="#Page_141">141</a><br /> +<br /> +Aramis, scapular arch, <a href="#Page_113">113</a><br /> +<br /> +Archæopteryx, <a href="#Page_58">58</a>, <a href="#Page_76">76</a>, <a href="#Page_104">104</a>, <a href="#Page_130">130</a>, <a href="#Page_197">197</a>, <a href="#Page_211">211</a><br /> +<br /> +Aristosuchus, <a href="#Page_129">129</a>, <a href="#Page_190">190</a>, <a href="#Page_205">205</a>, <a href="#Page_209">209</a><br /> +<br /> +Armadillo, <a href="#Page_40">40</a>, <a href="#Page_141">141</a><br /> +<br /> +Articulation of the jaw, <a href="#Page_12">12</a>, <a href="#Page_75">75</a><br /> +<br /> +Ashwell, <a href="#Page_177">177</a><br /> +<br /> +Atlantosaurus, <a href="#Page_202">202</a><br /> +<br /> +Atlas and axis, <a href="#Page_80">80</a>, <a href="#Page_81">81</a><br /> +<br /> +Aves, <a href="#Page_190">190</a><br /> +<br /> +Avian characters, <a href="#Page_220">220</a>, <a href="#Page_222">222</a><br /> +<br /> +<br /> +<b>B</b><br /> +<br /> +Backbone, <a href="#Page_78">78</a>, <a href="#Page_84">84</a><br /> +<br /> +Banz, <a href="#Page_148">148</a><br /> +<br /> +Barbastelle, <a href="#Page_25">25</a><br /> +<br /> +Barrington, <a href="#Page_177">177</a><br /> +<br /> +Barton, <a href="#Page_177">177</a><br /> +<br /> +Bat, <a href="#Page_38">38</a>, <a href="#Page_110">110</a>, <a href="#Page_197">197</a>;<br /> +<span style="margin-left: 1em;">sternum of, <a href="#Page_107">107</a>;</span><br /> +<span style="margin-left: 1em;">metacarpus, <a href="#Page_128">128</a></span><br /> +<br /> +Bavaria, <a href="#Page_156">156</a>, <a href="#Page_185">185</a><br /> +<br /> +Beak, horny, <a href="#Page_74">74</a>, <a href="#Page_178">178</a><br /> +<br /> +Bear, skull of, <a href="#Page_12">12</a>;<br /> +<span style="margin-left: 1em;">femur, <a href="#Page_100">100</a></span><br /> +<br /> +Bel and the Dragon, <a href="#Page_15">15</a><br /> +<br /> +Belodon, <a href="#Page_202">202</a><br /> +<br /> +Bird, <a href="#Page_80">80</a>, <a href="#Page_110">110</a>, <a href="#Page_120">120</a><br /> +<br /> +— resemblances, <a href="#Page_63">63</a>, <a href="#Page_65">65</a>, <a href="#Page_71">71</a>, <a href="#Page_95">95</a>, <a href="#Page_102">102</a>, <a href="#Page_108">108</a>, <a href="#Page_113">113</a>, <a href="#Page_119">119</a>, <a href="#Page_120">120</a>, <a href="#Page_211">211</a><br /> +<br /> +Bird-reptile, <a href="#Page_188">188</a><br /> +<br /> +Bird wing, <a href="#Page_128">128</a>, <a href="#Page_130">130</a><br /> +<br /> +Birds in flight, <a href="#Page_22">22</a>;<br /> +<span style="margin-left: 1em;">with teeth, <a href="#Page_76">76</a></span><br /> +<br /> +Black-headed bunting, <a href="#Page_47">47</a><br /> +<br /> +Blainville, D. de, <a href="#Page_30">30</a>, <a href="#Page_193">193</a><br /> +<br /> +Blood, temperature of, <a href="#Page_56">56</a><br /> +<br /> +Bohemia, <a href="#Page_34">34</a><br /> +<br /> +Bonaparte, Prince Charles, <a href="#Page_30">30</a><br /> +<br /> +Bones of birds, variation in, <a href="#Page_41">41</a><br /> +<br /> +— of reptiles, variation in, <a href="#Page_42">42</a><br /> +<br /> +— about the brain, <a href="#Page_69">69</a><br /> +<br /> +— in the back, <a href="#Page_84">84</a><br /> +<br /> +Bone texture, <a href="#Page_59">59</a>, <a href="#Page_209">209</a><br /> +<br /> +Bonn Museum, <a href="#Page_32">32</a>, <a href="#Page_85">85</a>, <a href="#Page_156">156</a><br /> +<br /> +Brain and breathing organs, <a href="#Page_55">55</a><br /> +<br /> +Brain cavity, in birds and reptiles, <a href="#Page_52">52</a>;<br /> +<span style="margin-left: 1em;">in mammals, <a href="#Page_221">221</a>, <a href="#Page_226">226</a>;</span><br /> +<span style="margin-left: 1em;">in Solenhofen pterodactyles, <a href="#Page_54">54</a>, <a href="#Page_220">220</a></span><br /> +<br /> +Brazil, <a href="#Page_34">34</a><br /> +<br /> +Breathing organs, <a href="#Page_8">8</a><br /> +<br /> +Bridgewater Treatise, <a href="#Page_143">143</a><br /> +<br /> +British Museum, <a href="#Page_133">133</a>, <a href="#Page_183">183</a><br /> +<br /> +Brixton, Isle of Wight, <a href="#Page_55">55</a>, <a href="#Page_174">174</a><br /> +<br /> +Buckland, Dean, <a href="#Page_143">143</a>, <a href="#Page_148">148</a>, <a href="#Page_231">231</a><br /> +<br /> +Burrowing limb, <a href="#Page_38">38</a><br /> +<span class='pagenum'><a name="Page_234" id="Page_234">[Pg 234]</a></span><br /> +<br /> +<b>C</b><br /> +<br /> +Cactus, <a href="#Page_227">227</a><br /> +<br /> +Calamospondylus, <a href="#Page_203">203</a><br /> +<br /> +Cambridge Greensand, <a href="#Page_33">33</a>, <a href="#Page_89">89</a>, <a href="#Page_176">176</a><br /> +<br /> +— Museum, <a href="#Page_177">177</a><br /> +<br /> +Camel, <a href="#Page_83">83</a><br /> +<br /> +Campylognathus, <a href="#Page_68">68</a>, <a href="#Page_71">71</a>, <a href="#Page_135">135</a>;<br /> +<span style="margin-left: 1em;">size of, <a href="#Page_149">149</a></span><br /> +<br /> +Canary, <a href="#Page_47">47</a><br /> +<br /> +Carnivorous dinosaurs, <a href="#Page_129">129</a><br /> +<br /> +Carpus, <a href="#Page_122">122</a><br /> +<br /> +Caudal fin, <a href="#Page_91">91</a>, <a href="#Page_161">161</a><br /> +<br /> +— vertebræ, <a href="#Page_89">89</a>, <a href="#Page_92">92</a>, <a href="#Page_203">203</a><br /> +<br /> +Ceratodus, <a href="#Page_4">4</a>, <a href="#Page_5">5</a>, <a href="#Page_9">9</a>, <a href="#Page_17">17</a><br /> +<br /> +Ceratosaurus, <a href="#Page_203">203</a>, <a href="#Page_204">204</a><br /> +<br /> +Cervical rib, <a href="#Page_81">81</a><br /> +<br /> +Cetacea, <a href="#Page_40">40</a><br /> +<br /> +Cetiosaurus, <a href="#Page_198">198</a>, <a href="#Page_203">203</a><br /> +<br /> +Chalinolobus, <a href="#Page_25">25</a><br /> +<br /> +Chalk, pterodactyles in, <a href="#Page_136">136</a>;<br /> +<span style="margin-left: 1em;">of Kansas, <a href="#Page_103">103</a>, <a href="#Page_132">132</a></span><br /> +<br /> +Chameleon, <a href="#Page_17">17</a>, <a href="#Page_51">51</a>, <a href="#Page_70">70</a>;<br /> +<span style="margin-left: 1em;">scapula, <a href="#Page_112">112</a>;</span><br /> +<span style="margin-left: 1em;">sternum, <a href="#Page_107">107</a></span><br /> +<br /> +Chameleonoidea, <a href="#Page_191">191</a><br /> +<br /> +Cheek bones, <a href="#Page_178">178</a><br /> +<br /> +Chelonia, <a href="#Page_86">86</a>, <a href="#Page_112">112</a>, <a href="#Page_193">193</a><br /> +<br /> +Chesterton, <a href="#Page_177">177</a><br /> +<br /> +Chlamydosaurus, <a href="#Page_21">21</a><br /> +<br /> +<i>Chrysochloris capensis</i>, <a href="#Page_121">121</a><br /> +<br /> +Classification, <a href="#Page_192">192</a>;<br /> +<span style="margin-left: 1em;">on pelvis characters, <a href="#Page_195">195</a>;</span><br /> +<span style="margin-left: 1em;">of dinosaurs, <a href="#Page_198">198</a></span><br /> +<br /> +Clavicles, <a href="#Page_111">111</a>, <a href="#Page_112">112</a><br /> +<br /> +Claw, <a href="#Page_105">105</a>, <a href="#Page_116">116</a>, <a href="#Page_183">183</a>, <a href="#Page_208">208</a><br /> +<br /> +Cœlurus, <a href="#Page_203">203</a>, <a href="#Page_209">209</a><br /> +<br /> +Coldham Common, <a href="#Page_177">177</a><br /> +<br /> +Collar bone, <a href="#Page_111">111</a><br /> +<br /> +Collini, <a href="#Page_27">27</a><br /> +<br /> +Comparison with dinosaurs, <a href="#Page_198">198</a>;<br /> +<span style="margin-left: 1em;">of pelvis, <a href="#Page_204">204</a>, <a href="#Page_206">206</a>;</span><br /> +<span style="margin-left: 1em;">of skulls, <a href="#Page_192">192</a>, <a href="#Page_199">199</a>, <a href="#Page_201">201</a></span><br /> +<br /> +Cope, Professor, <a href="#Page_31">31</a>, <a href="#Page_34">34</a><br /> +<br /> +Coracoid, <a href="#Page_109">109</a>, <a href="#Page_112">112</a>, <a href="#Page_113">113</a><br /> +<br /> +Cordylomorpha, <a href="#Page_191">191</a><br /> +<br /> +Cormorant, <a href="#Page_70">70</a>, <a href="#Page_174">174</a>;<br /> +<span style="margin-left: 1em;">sternum, <a href="#Page_108">108</a></span><br /> +<br /> +Corpora quadrigemina, <a href="#Page_221">221</a><br /> +<br /> +Crisp, Dr., on pneumatic skeleton, <a href="#Page_47">47</a><br /> +<br /> +Crocodile, characters of, <a href="#Page_217">217</a>;<br /> +<span style="margin-left: 1em;">heart, <a href="#Page_56">56</a>;</span><br /> +<span style="margin-left: 1em;">lung, <a href="#Page_9">9</a>;</span><br /> +<span style="margin-left: 1em;">shoulder-girdle, <a href="#Page_111">111</a>;</span><br /> +<span style="margin-left: 1em;">skull, <a href="#Page_46">46</a>;</span><br /> +<span style="margin-left: 1em;">vertebræ, <a href="#Page_79">79</a></span><br /> +<br /> +Crocodilia, <a href="#Page_190">190</a><br /> +<br /> +Curlew, <a href="#Page_68">68</a><br /> +<br /> +Cuvier, <a href="#Page_1">1</a>, <a href="#Page_27">27</a>, <a href="#Page_28">28</a>, <a href="#Page_54">54</a>, <a href="#Page_76">76</a>, <a href="#Page_77">77</a>, <a href="#Page_130">130</a>, <a href="#Page_231">231</a><br /> +<br /> +Cycnorhamphus, <a href="#Page_70">70</a>, <a href="#Page_94">94</a>, <a href="#Page_171">171</a>, <a href="#Page_173">173</a>, <a href="#Page_204">204</a><br /> +<br /> +<i>Cycnorhamphus Fraasii</i>, <a href="#Page_80">80</a>, <a href="#Page_96">96</a>, <a href="#Page_169">169</a><br /> +<br /> +— <i>suevicus</i>, <a href="#Page_169">169</a>, <a href="#Page_170">170</a><br /> +<br /> +Cypselus, <a href="#Page_42">42</a><br /> +<br /> +<br /> +<b>D</b><br /> +<br /> +<i>Dacelo gigantea</i>, <a href="#Page_63">63</a><br /> +<br /> +Darwin, <a href="#Page_3">3</a><br /> +<br /> +Davy, Dr. John, <a href="#Page_142">142</a><br /> +<br /> +Deuterosaurus, <a href="#Page_97">97</a><br /> +<br /> +Dicynodon, <a href="#Page_200">200</a><br /> +<br /> +<i>Dicynodon lacerticeps</i>, <a href="#Page_71">71</a><br /> +<br /> +Digits, of ostrich, <a href="#Page_23">23</a>;<br /> +<span style="margin-left: 1em;">of pterodactyle, <a href="#Page_128">128</a></span><br /> +<br /> +Digits with claws, <a href="#Page_130">130</a>;<br /> +<span style="margin-left: 1em;">foot bones in, <a href="#Page_105">105</a></span><br /> +<br /> +Dimorphodon, <a href="#Page_63">63</a>, <a href="#Page_64">64</a>, <a href="#Page_66">66</a>, <a href="#Page_67">67</a>, <a href="#Page_73">73</a>, <a href="#Page_74">74</a>, <a href="#Page_83">83</a>, <a href="#Page_90">90</a>, <a href="#Page_102">102</a>, <a href="#Page_113">113</a>, <a href="#Page_143">143</a>, <a href="#Page_192">192</a>, <a href="#Page_194">194</a>, <a href="#Page_199">199</a>, <a href="#Page_201">201</a>, <a href="#Page_206">206</a><br /> +<br /> +Dinosauria, <a href="#Page_6">6</a>, <a href="#Page_77">77</a>, <a href="#Page_84">84</a>, <a href="#Page_87">87</a>, <a href="#Page_95">95</a>, <a href="#Page_129">129</a>, <a href="#Page_144">144</a>, <a href="#Page_198">198</a>, <a href="#Page_209">209</a><br /> +<br /> +Dinosaurs from Lias, <a href="#Page_135">135</a>, <a href="#Page_192">192</a>;<br /> +<span style="margin-left: 1em;">from Elgin, <a href="#Page_201">201</a>, <a href="#Page_207">207</a>;</span><br /> +<span style="margin-left: 1em;">Stuttgart, <a href="#Page_202">202</a>;</span><br /> +<span style="margin-left: 1em;">Trias dinosaurs, <a href="#Page_199">199</a>, <a href="#Page_200">200</a></span><br /> +<br /> +Diopecephalus, <a href="#Page_168">168</a><br /> +<br /> +Diving birds, <a href="#Page_23">23</a>, <a href="#Page_83">83</a>, <a href="#Page_102">102</a><br /> +<br /> +Dolichosauria, <a href="#Page_191">191</a><br /> +<br /> +Dolphin, <a href="#Page_107">107</a><br /> +<br /> +Doratorhynchus, <a href="#Page_173">173</a><br /> +<br /> +Dorygnathus, <a href="#Page_74">74</a>, <a href="#Page_148">148</a><br /> +<br /> +Dragons, <a href="#Page_3">3</a>, <a href="#Page_15">15</a>, <a href="#Page_17">17</a><br /> +<br /> +Drumstick bone, <a href="#Page_103">103</a>, <a href="#Page_195">195</a><br /> +<br /> +Duck, <a href="#Page_22">22</a>, <a href="#Page_83">83</a><br /> +<br /> +<br /> +<b>E</b><br /> +<br /> +Echidna, <a href="#Page_75">75</a>, <a href="#Page_76">76</a>, <a href="#Page_95">95</a>, <a href="#Page_100">100</a><br /> +<br /> +Edentata, <a href="#Page_185">185</a><br /> +<br /> +Edentulous beak, <a href="#Page_153">153</a><br /> +<br /> +Eichstädt, <a href="#Page_32">32</a><br /> +<br /> +Elephant, head of, <a href="#Page_46">46</a><br /> +<br /> +Enumeration of characters, <a href="#Page_223">223</a>, <a href="#Page_225">225</a><br /> +<br /> +Ephesus, winged figure, <a href="#Page_16">16</a><br /> +<br /> +Epiphysis to first phalange, <a href="#Page_123">123</a><br /> +<br /> +Exocœtus, <a href="#Page_18">18</a><br /> +<span class='pagenum'><a name="Page_235" id="Page_235">[Pg 235]</a></span><br /> +Extinctions, <a href="#Page_129">129</a><br /> +<br /> +Eye hole, <a href="#Page_144">144</a>;<br /> +<span style="margin-left: 1em;">sclerotic bones in, <a href="#Page_65">65</a></span><br /> +<br /> +<br /> +<b>F</b><br /> +<br /> +Farren, William, <a href="#Page_34">34</a><br /> +<br /> +Femur, <a href="#Page_100">100</a><br /> +<br /> +Fibula, <a href="#Page_102">102</a>, <a href="#Page_183">183</a>, <a href="#Page_206">206</a><br /> +<br /> +Fifth outer digit, <a href="#Page_132">132</a>;<br /> +<span style="margin-left: 1em;">in foot, <a href="#Page_145">145</a></span><br /> +<br /> +Figure from temple at Ephesus, <a href="#Page_16">16</a><br /> +<br /> +First phalange, <a href="#Page_151">151</a><br /> +<br /> +Fish-eating crocodile, <a href="#Page_137">137</a><br /> +<br /> +Flight, organs of, <a href="#Page_17">17</a>;<br /> +<span style="margin-left: 1em;">in bats, <a href="#Page_25">25</a></span><br /> +<br /> +Flying limb, <a href="#Page_38">38</a><br /> +<br /> +Flying fishes, <a href="#Page_18">18</a>, <a href="#Page_57">57</a>;<br /> +<span style="margin-left: 1em;">foxes, <a href="#Page_26">26</a>;</span><br /> +<span style="margin-left: 1em;">frogs, <a href="#Page_19">19</a>, <a href="#Page_197">197</a>;</span><br /> +<span style="margin-left: 1em;">gecko, <a href="#Page_21">21</a>, <a href="#Page_24">24</a>;</span><br /> +<span style="margin-left: 1em;">lizards, <a href="#Page_20">20</a>;</span><br /> +<span style="margin-left: 1em;">reptiles, <a href="#Page_37">37</a>, <a href="#Page_46">46</a>;</span><br /> +<span style="margin-left: 1em;">squirrel, <a href="#Page_24">24</a></span><br /> +<br /> +Foot, <a href="#Page_104">104</a>;<br /> +<span style="margin-left: 1em;">digits in, <a href="#Page_105">105</a>, <a href="#Page_146">146</a></span><br /> +<br /> +Fore leg, <a href="#Page_102">102</a>, <a href="#Page_206">206</a><br /> +<br /> +— limb, <a href="#Page_38">38</a>, <a href="#Page_107">107</a>, <a href="#Page_116">116</a>, <a href="#Page_120">120</a><br /> +<br /> +Four claws, <a href="#Page_147">147</a><br /> +<br /> +Fox, Rev. W., <a href="#Page_55">55</a>, <a href="#Page_174">174</a><br /> +<br /> +Fraas, Professor Oscar, <a href="#Page_172">172</a>, <a href="#Page_231">231</a><br /> +<br /> +Frigate bird, vertebræ of, <a href="#Page_86">86</a>, <a href="#Page_174">174</a><br /> +<br /> +Frog, lungs of, <a href="#Page_8">8</a><br /> +<br /> +Furculum, <a href="#Page_114">114</a><br /> +<br /> +<br /> +<b>G</b><br /> +<br /> +Gaudry, Professor A., <a href="#Page_31">31</a><br /> +<br /> +Gavial, <a href="#Page_136">136</a><br /> +<br /> +Gecko, <a href="#Page_21">21</a>, <a href="#Page_23">23</a><br /> +<br /> +Genera, comparison of, <a href="#Page_192">192</a><br /> +<br /> +Geological distribution, <a href="#Page_186">186</a><br /> +<br /> +Gills, <a href="#Page_4">4</a><br /> +<br /> +Giraffe, <a href="#Page_38">38</a>, <a href="#Page_39">39</a><br /> +<br /> +Glossy starling, <a href="#Page_47">47</a><br /> +<br /> +Golden eagle, <a href="#Page_120">120</a><br /> +<br /> +— mole, <a href="#Page_121">121</a><br /> +<br /> +Goldfuss, <a href="#Page_30">30</a>, <a href="#Page_231">231</a><br /> +<br /> +Granchester, <a href="#Page_177">177</a><br /> +<br /> +Great ant-eater, <a href="#Page_40">40</a>, <a href="#Page_185">185</a><br /> +<br /> +Guillemot, <a href="#Page_102">102</a><br /> +<br /> +Gull, <a href="#Page_22">22</a><br /> +<br /> +<br /> +<b>H</b><br /> +<br /> +Haarlem, Teyler Museum at, <a href="#Page_32">32</a><br /> +<br /> +Habits, probable, <a href="#Page_134">134</a>, <a href="#Page_176">176</a>, <a href="#Page_198">198</a><br /> +<br /> +Hairless skins, <a href="#Page_141">141</a><br /> +<br /> +Hand in mammals, <a href="#Page_38">38</a><br /> +<br /> +Harston, <a href="#Page_177">177</a><br /> +<br /> +Haslingfield, <a href="#Page_177">177</a><br /> +<br /> +Hastings, <a href="#Page_174">174</a><br /> +<br /> +Hatteria lung, <a href="#Page_9">9</a>, <a href="#Page_27">27</a>;<br /> +<span style="margin-left: 1em;">brain, <a href="#Page_53">53</a>;</span><br /> +<span style="margin-left: 1em;">skull, <a href="#Page_70">70</a>, <a href="#Page_77">77</a>;</span><br /> +<span style="margin-left: 1em;">ribs, <a href="#Page_86">86</a>;</span><br /> +<span style="margin-left: 1em;">a reptile type, <a href="#Page_13">13</a></span><br /> +<br /> +Head, characters of, <a href="#Page_76">76</a><br /> +<br /> +Heidelberg Museum, <a href="#Page_32">32</a>, <a href="#Page_54">54</a>, <a href="#Page_159">159</a><br /> +<br /> +Herpetomorpha, <a href="#Page_191">191</a><br /> +<br /> +Heron, <a href="#Page_65">65</a>, <a href="#Page_174">174</a><br /> +<br /> +Hesperornis, <a href="#Page_76">76</a><br /> +<br /> +Hind foot, <a href="#Page_104">104</a>, <a href="#Page_135">135</a><br /> +<br /> +— limb, <a href="#Page_93">93</a>, <a href="#Page_99">99</a>, <a href="#Page_159">159</a>, <a href="#Page_206">206</a><br /> +<br /> +Hip-girdle in whale tribe, <a href="#Page_39">39</a>, <a href="#Page_159">159</a><br /> +<br /> +Homœosauria, <a href="#Page_191">191</a><br /> +<br /> +Horningsea, <a href="#Page_177">177</a><br /> +<br /> +Horse, metacarpus of, <a href="#Page_127">127</a>;<br /> +<span style="margin-left: 1em;">vertebræ of, <a href="#Page_79">79</a></span><br /> +<br /> +Humerus, <a href="#Page_46">46</a>, <a href="#Page_117">117</a>, <a href="#Page_217">217</a><br /> +<br /> +Huxley, Professor, <a href="#Page_31">31</a>, <a href="#Page_89">89</a>, <a href="#Page_154">154</a>, <a href="#Page_188">188</a><br /> +<br /> +Hyo-mandibular arch, <a href="#Page_13">13</a><br /> +<br /> +Hypothesis of descent, <a href="#Page_226">226</a><br /> +<br /> +Hyrax, <a href="#Page_101">101</a><br /> +<br /> +<br /> +<b>I</b><br /> +<br /> +Ichthyornis, <a href="#Page_76">76</a><br /> +<br /> +Ichthyosaurus, <a href="#Page_6">6</a>, <a href="#Page_191">191</a><br /> +<br /> +Iguanodon, <a href="#Page_209">209</a>;<br /> +<span style="margin-left: 1em;">pelvis, <a href="#Page_206">206</a></span><br /> +<br /> +Ilium, <a href="#Page_93">93</a>, <a href="#Page_95">95</a>, <a href="#Page_96">96</a>, <a href="#Page_98">98</a>, <a href="#Page_204">204</a><br /> +<br /> +Instep, <a href="#Page_105">105</a>, <a href="#Page_207">207</a><br /> +<br /> +Inherited characters, <a href="#Page_217">217</a><br /> +<br /> +Interclavicle, <a href="#Page_111">111</a><br /> +<br /> +Ischium, <a href="#Page_93">93</a>, <a href="#Page_96">96</a>, <a href="#Page_203">203</a>, <a href="#Page_204">204</a><br /> +<br /> +Isle of Wight, <a href="#Page_174">174</a><br /> +<br /> +<br /> +<b>J</b><br /> +<br /> +Jaw, in birds, <a href="#Page_12">12</a>;<br /> +<span style="margin-left: 1em;">in fishes, <a href="#Page_13">13</a>;</span><br /> +<span style="margin-left: 1em;">in mammals, <a href="#Page_12">12</a>;</span><br /> +<span style="margin-left: 1em;">in reptiles, <a href="#Page_13">13</a>;</span><br /> +<span style="margin-left: 1em;">in pterodactyles, <a href="#Page_63">63</a>;</span><br /> +<span style="margin-left: 1em;">suspension of, <a href="#Page_11">11</a>, <a href="#Page_74">74</a>, <a href="#Page_76">76</a></span><br /> +<br /> +— lower, <a href="#Page_75">75</a><br /> +<br /> +<br /> +<b>K</b><br /> +<br /> +Kansas, Chalk of, <a href="#Page_72">72</a>, <a href="#Page_103">103</a>, <a href="#Page_115">115</a>;<br /> +<span style="margin-left: 1em;">University Museum of, <a href="#Page_181">181</a></span><br /> +<br /> +Kelheim, <a href="#Page_32">32</a><br /> +<br /> +Keuper, <a href="#Page_33">33</a><br /> +<br /> +Kimeridge Clay, <a href="#Page_132">132</a><br /> +<br /> +Kingfisher, <a href="#Page_63">63</a><br /> +<br /> +Kiwi, <a href="#Page_23">23</a><br /> +<span class='pagenum'><a name="Page_236" id="Page_236">[Pg 236]</a></span><br /> +<br /> +<b>L</b><br /> +<br /> +Labyrinthodontia, <a href="#Page_191">191</a><br /> +<br /> +Lachrymal bones, <a href="#Page_67">67</a><br /> +<br /> +Laramie rocks, <a href="#Page_34">34</a><br /> +<br /> +Largest ornithosaur, <a href="#Page_133">133</a><br /> +<br /> +Lateral vacuities in skull, <a href="#Page_147">147</a><br /> +<br /> +Lawrence in Kansas, <a href="#Page_181">181</a><br /> +<br /> +Lengths of bones, <a href="#Page_146">146</a><br /> +<br /> +Lepidosiren, <a href="#Page_17">17</a><br /> +<br /> +Lias, <a href="#Page_33">33</a><br /> +<br /> +Lithographic Slate, <a href="#Page_35">35</a>, <a href="#Page_156">156</a><br /> +<br /> +Lizards, <a href="#Page_20">20</a>, <a href="#Page_21">21</a>, <a href="#Page_27">27</a>, <a href="#Page_123">123</a><br /> +<br /> +Llama, neck of, <a href="#Page_79">79</a>, <a href="#Page_83">83</a><br /> +<br /> +Loach, swim bladder of, <a href="#Page_52">52</a><br /> +<br /> +Lower jaw, <a href="#Page_12">12</a>, <a href="#Page_74">74</a>, <a href="#Page_76">76</a>, <a href="#Page_149">149</a><br /> +<br /> +Lumbar vertebræ, <a href="#Page_89">89</a><br /> +<br /> +Lungs, <a href="#Page_47">47</a>;<br /> +<span style="margin-left: 1em;">in apteryx, <a href="#Page_48">48</a>;</span><br /> +<span style="margin-left: 1em;">in chameleon, <a href="#Page_51">51</a>;</span><br /> +<span style="margin-left: 1em;">in ostrich, <a href="#Page_49">49</a>;</span><br /> +<span style="margin-left: 1em;">in reptiles, <a href="#Page_8">8</a>, <a href="#Page_9">9</a>, <a href="#Page_51">51</a></span><br /> +<br /> +Lydekker, R., <a href="#Page_160">160</a>, <a href="#Page_169">169</a>, <a href="#Page_231">231</a><br /> +<br /> +Lyme Regis, <a href="#Page_33">33</a><br /> +<br /> +<br /> +<b>M</b><br /> +<br /> +Macrocercus, palate of, <a href="#Page_71">71</a><br /> +<br /> +Malar bone, <a href="#Page_67">67</a><br /> +<br /> +Mallard, <a href="#Page_22">22</a><br /> +<br /> +Mammal, <a href="#Page_8">8</a>, <a href="#Page_12">12</a>, <a href="#Page_24">24</a>, <a href="#Page_79">79</a>, <a href="#Page_53">53</a>, <a href="#Page_95">95</a><br /> +<br /> +Mammalia, <a href="#Page_38">38</a>, <a href="#Page_141">141</a><br /> +<br /> +Mammalian characters, <a href="#Page_12">12</a>, <a href="#Page_220">220</a><br /> +<br /> +Mammoth, <a href="#Page_141">141</a><br /> +<br /> +Manis, <a href="#Page_40">40</a>, <a href="#Page_57">57</a>, <a href="#Page_142">142</a><br /> +<br /> +Manubrium of sternum, <a href="#Page_108">108</a>, <a href="#Page_109">109</a>, <a href="#Page_183">183</a><br /> +<br /> +Marrow bones in a bird, <a href="#Page_134">134</a><br /> +<br /> +Marsh, Professor O. C., <a href="#Page_31">31</a>, <a href="#Page_72">72</a>, <a href="#Page_90">90</a>, <a href="#Page_115">115</a>, <a href="#Page_121">121</a>, <a href="#Page_131">131</a>, <a href="#Page_140">140</a>, <a href="#Page_160">160</a>, <a href="#Page_165">165</a>, <a href="#Page_180">180</a>, <a href="#Page_181">181</a>, <a href="#Page_210">210</a>, <a href="#Page_231">231</a><br /> +<br /> +Marsupial, <a href="#Page_70">70</a>, <a href="#Page_94">94</a>, <a href="#Page_99">99</a><br /> +<br /> +Megalosaurus, <a href="#Page_129">129</a>, <a href="#Page_198">198</a><br /> +<br /> +Merganser, <a href="#Page_108">108</a><br /> +<br /> +Merry-thought, <a href="#Page_114">114</a><br /> +<br /> +Metacarpus, <a href="#Page_116">116</a>, <a href="#Page_124">124</a>, <a href="#Page_126">126</a>, <a href="#Page_128">128</a>, <a href="#Page_130">130</a><br /> +<br /> +Metatarsal bones, <a href="#Page_104">104</a>, <a href="#Page_207">207</a>, <a href="#Page_208">208</a><br /> +<br /> +Meyer, Hermann von, <a href="#Page_31">31</a>, <a href="#Page_45">45</a>, <a href="#Page_46">46</a>, <a href="#Page_85">85</a>, <a href="#Page_105">105</a>, <a href="#Page_108">108</a>, <a href="#Page_121">121</a>, <a href="#Page_160">160</a>, <a href="#Page_192">192</a>, <a href="#Page_231">231</a><br /> +<br /> +Moa of New Zealand, <a href="#Page_35">35</a><br /> +<br /> +Mole, humerus, <a href="#Page_38">38</a>;<br /> +<span style="margin-left: 1em;">sternum, <a href="#Page_107">107</a></span><br /> +<br /> +Monotremes, <a href="#Page_70">70</a>, <a href="#Page_94">94</a>, <a href="#Page_111">111</a>, <a href="#Page_121">121</a>, <a href="#Page_185">185</a>, <a href="#Page_218">218</a><br /> +<br /> +Mososaurus, <a href="#Page_77">77</a><br /> +<br /> +Movement of the leg, <a href="#Page_101">101</a><br /> +<br /> +Mugger, <a href="#Page_137">137</a><br /> +<br /> +Munich Museum, <a href="#Page_32">32</a>, <a href="#Page_159">159</a><br /> +<br /> +Munster, von, <a href="#Page_231">231</a><br /> +<br /> +Muschelkalk, <a href="#Page_184">184</a><br /> +<br /> +Museum, <a href="#Page_32">32</a>, <a href="#Page_156">156</a>, <a href="#Page_231">231</a>, <a href="#Page_159">159</a>;<br /> +<span style="margin-left: 1em;">Natural History, <a href="#Page_133">133</a>, <a href="#Page_231">231</a></span><br /> +<br /> +Myrmecophaga, <a href="#Page_185">185</a><br /> +<br /> +<br /> +<b>N</b><br /> +<br /> +Names of genera, <a href="#Page_183">183</a><br /> +<br /> +Natural History Museum, <a href="#Page_38">38</a>, <a href="#Page_231">231</a><br /> +<br /> +Neck, <a href="#Page_79">79</a>;<br /> +<span style="margin-left: 1em;">in Dimorphodon, <a href="#Page_145">145</a>;</span><br /> +<span style="margin-left: 1em;">in Giraffe, <a href="#Page_39">39</a>;</span><br /> +<span style="margin-left: 1em;">in Llama, <a href="#Page_79">79</a>;</span><br /> +<span style="margin-left: 1em;">in Pterodactyles, <a href="#Page_80">80</a>;</span><br /> +<span style="margin-left: 1em;">in Whales, <a href="#Page_39">39</a></span><br /> +<br /> +Newton, E. T., <a href="#Page_55">55</a>, <a href="#Page_70">70</a>, <a href="#Page_158">158</a>, <a href="#Page_160">160</a>, <a href="#Page_201">201</a>, <a href="#Page_232">232</a><br /> +<br /> +New Zealand Bat, <a href="#Page_25">25</a><br /> +<br /> +— — Hatteria, <a href="#Page_68">68</a><br /> +<br /> +Niobrara rock, <a href="#Page_183">183</a><br /> +<br /> +Nostril, bones round the, <a href="#Page_62">62</a>;<br /> +<span style="margin-left: 1em;">small, <a href="#Page_147">147</a></span><br /> +<br /> +Notarium, <a href="#Page_87">87</a>, <a href="#Page_115">115</a><br /> +<br /> +Nothosauria, <a href="#Page_192">192</a><br /> +<br /> +Nusplingen, <a href="#Page_32">32</a><br /> +<br /> +Nyctodactylus, <a href="#Page_115">115</a>, <a href="#Page_180">180</a><br /> +<br /> +<br /> +<b>O</b><br /> +<br /> +Obliteration of characters, <a href="#Page_216">216</a><br /> +<br /> +Opercular bones, <a href="#Page_13">13</a><br /> +<br /> +Ophidia, <a href="#Page_52">52</a>, <a href="#Page_191">191</a><br /> +<br /> +Optic lobes, <a href="#Page_53">53</a>, <a href="#Page_221">221</a><br /> +<br /> +Organs of flight, <a href="#Page_17">17</a><br /> +<br /> +Ornithischia, <a href="#Page_190">190</a>, <a href="#Page_198">198</a><br /> +<br /> +Ornithocephalus, <a href="#Page_166">166</a><br /> +<br /> +Ornithocheirus, atlas and axis, <a href="#Page_81">81</a>;<br /> +<span style="margin-left: 1em;">brain, <a href="#Page_55">55</a>, <a href="#Page_69">69</a>;</span><br /> +<span style="margin-left: 1em;">carpus, <a href="#Page_124">124</a>;</span><br /> +<span style="margin-left: 1em;">cervical vertebra, <a href="#Page_83">83</a>, <a href="#Page_179">179</a>;</span><br /> +<span style="margin-left: 1em;">claw phalange, <a href="#Page_129">129</a>;</span><br /> +<span style="margin-left: 1em;">coracoid, <a href="#Page_109">109</a>;</span><br /> +<span style="margin-left: 1em;">femur, <a href="#Page_100">100</a>;</span><br /> +<span style="margin-left: 1em;">pelvis, <a href="#Page_98">98</a>;</span><br /> +<span style="margin-left: 1em;">pubic bones, <a href="#Page_194">194</a>;</span><br /> +<span style="margin-left: 1em;">sternum, <a href="#Page_109">109</a>;</span><br /> +<span style="margin-left: 1em;">shoulder-girdle, <a href="#Page_115">115</a>;</span><br /> +<span style="margin-left: 1em;">remains, <a href="#Page_176">176</a>;</span><br /> +<span style="margin-left: 1em;">teeth, <a href="#Page_74">74</a>, <a href="#Page_76">76</a>;</span><br /> +<span style="margin-left: 1em;">absence of teeth, <a href="#Page_138">138</a></span><br /> +<br /> +<i>Ornithocheirus machærorhynchus</i>, <a href="#Page_139">139</a>;<br /> +<span style="margin-left: 1em;"><i>microdon</i>, <a href="#Page_139">139</a></span><br /> +<span class='pagenum'><a name="Page_237" id="Page_237">[Pg 237]</a></span><br /> +Ornithocheiroidea, <a href="#Page_193">193</a><br /> +<br /> +Ornithodesmus, neck bones, <a href="#Page_173">173</a>, <a href="#Page_175">175</a>;<br /> +<span style="margin-left: 1em;">coracoid, <a href="#Page_109">109</a>, <a href="#Page_116">116</a>;</span><br /> +<span style="margin-left: 1em;">dorsal vertebræ, <a href="#Page_86">86</a>;</span><br /> +<span style="margin-left: 1em;">remains of <i>O. latidens</i>, <a href="#Page_173">173</a>;</span><br /> +<span style="margin-left: 1em;"><i>O. sagittirostris</i>, <a href="#Page_175">175</a></span><br /> +<br /> +Ornithomorpha, <a href="#Page_189">189</a><br /> +<br /> +Ornithorhynchus, <a href="#Page_40">40</a>, <a href="#Page_53">53</a>, <a href="#Page_95">95</a>, <a href="#Page_117">117</a><br /> +<br /> +Ornithosauria, <a href="#Page_30">30</a>, <a href="#Page_31">31</a>, <a href="#Page_50">50</a>, <a href="#Page_52">52</a>, <a href="#Page_58">58</a>, <a href="#Page_72">72</a>, <a href="#Page_89">89</a>, <a href="#Page_95">95</a>, <a href="#Page_104">104</a>, <a href="#Page_108">108</a>, <a href="#Page_125">125</a>, <a href="#Page_132">132</a>, <a href="#Page_133">133</a>, <a href="#Page_143">143</a>, <a href="#Page_187">187</a>, <a href="#Page_190">190</a>, <a href="#Page_192">192</a>, <a href="#Page_216">216</a><br /> +<br /> +Ornithostoma, <a href="#Page_66">66</a>, <a href="#Page_69">69</a>, <a href="#Page_72">72</a>, <a href="#Page_180">180</a>;<br /> +<span style="margin-left: 1em;">lower jaw, <a href="#Page_75">75</a>, <a href="#Page_76">76</a>;</span><br /> +<span style="margin-left: 1em;">pelvis, <a href="#Page_98">98</a>;</span><br /> +<span style="margin-left: 1em;">sternum, <a href="#Page_110">110</a>;</span><br /> +<span style="margin-left: 1em;">phalange, <a href="#Page_122">122</a>;</span><br /> +<span style="margin-left: 1em;">size, <a href="#Page_133">133</a>;</span><br /> +<span style="margin-left: 1em;">skull, <a href="#Page_181">181</a>, <a href="#Page_182">182</a></span><br /> +<br /> +Ornithosuchus, <a href="#Page_201">201</a><br /> +<br /> +Orycteropus, <a href="#Page_96">96</a><br /> +<br /> +<i>Ossa innominata</i>, <a href="#Page_93">93</a><br /> +<br /> +Ossified ligaments, <a href="#Page_150">150</a><br /> +<br /> +Ostrich, <a href="#Page_23">23</a>, <a href="#Page_45">45</a>, <a href="#Page_49">49</a>, <a href="#Page_113">113</a>, <a href="#Page_129">129</a><br /> +<br /> +Owen, Sir R., <a href="#Page_31">31</a>, <a href="#Page_36">36</a>, <a href="#Page_46">46</a>, <a href="#Page_48">48</a>, <a href="#Page_110">110</a>, <a href="#Page_117">117</a>, <a href="#Page_143">143</a>, <a href="#Page_172">172</a>, <a href="#Page_176">176</a>, <a href="#Page_180">180</a>, <a href="#Page_231">231</a><br /> +<br /> +Owl, <a href="#Page_46">46</a>, <a href="#Page_53">53</a><br /> +<br /> +Oxford Clay, <a href="#Page_33">33</a>, <a href="#Page_156">156</a><br /> +<br /> +— University Museum, <a href="#Page_154">154</a><br /> +<br /> +Ox, vertebra of, <a href="#Page_79">79</a>;<br /> +<span style="margin-left: 1em;">metacarpus, <a href="#Page_127">127</a></span><br /> +<br /> +<br /> +<b>P</b><br /> +<br /> +Palate, bones of, <a href="#Page_71">71</a><br /> +<br /> +Pangolin, <a href="#Page_142">142</a><br /> +<br /> +Pappenheim, <a href="#Page_32">32</a><br /> +<br /> +Parallel groups, <a href="#Page_215">215</a><br /> +<br /> +Parrot, <a href="#Page_71">71</a><br /> +<br /> +Patagial membranes, <a href="#Page_227">227</a><br /> +<br /> +Pelican, <a href="#Page_174">174</a><br /> +<br /> +Pelvis, <a href="#Page_88">88</a>, <a href="#Page_94">94-98</a>, <a href="#Page_151">151</a>, <a href="#Page_195">195</a>, <a href="#Page_202">202</a>, <a href="#Page_204">204</a>, <a href="#Page_206">206</a><br /> +<br /> +Penguin, <a href="#Page_41">41</a>, <a href="#Page_42">42</a>, <a href="#Page_104">104</a>, <a href="#Page_176">176</a><br /> +<br /> +Periophthalmus, <a href="#Page_17">17</a><br /> +<br /> +Peterborough, bones from, <a href="#Page_113">113</a>, <a href="#Page_156">156</a><br /> +<br /> +Phalanges, <a href="#Page_129">129</a>, <a href="#Page_132">132</a>;<br /> +<span style="margin-left: 1em;">wing finger, <a href="#Page_155">155</a></span><br /> +<br /> +Phillips, Professor John, <a href="#Page_155">155</a><br /> +<br /> +Pigeon, <a href="#Page_119">119</a><br /> +<br /> +Platydactylus, <a href="#Page_21">21</a><br /> +<br /> +Platypus, <a href="#Page_214">214</a><br /> +<br /> +Plesiosaurus, <a href="#Page_6">6</a>, <a href="#Page_73">73</a>, <a href="#Page_75">75</a>, <a href="#Page_93">93</a>, <a href="#Page_189">189</a><br /> +<br /> +Pleininger, <a href="#Page_149">149</a>, <a href="#Page_232">232</a><br /> +<br /> +Pneumatic foramina, <a href="#Page_45">45</a>, <a href="#Page_83">83</a>, <a href="#Page_88">88</a>, <a href="#Page_132">132</a>, <a href="#Page_209">209</a><br /> +<br /> +Pond, Mr., <a href="#Page_34">34</a><br /> +<br /> +Porcupine, <a href="#Page_40">40</a><br /> +<br /> +Porpoise, <a href="#Page_38">38</a>, <a href="#Page_73">73</a>, <a href="#Page_141">141</a>, <a href="#Page_200">200</a><br /> +<br /> +Premaxillary bones, <a href="#Page_77">77</a>, <a href="#Page_200">200</a>, <a href="#Page_205">205</a><br /> +<br /> +Prepubic bones, <a href="#Page_94">94</a>, <a href="#Page_96">96-98</a>, <a href="#Page_194">194</a>, <a href="#Page_204">204</a>, <a href="#Page_205">205</a><br /> +<br /> +Protorosauria, <a href="#Page_192">192</a><br /> +<br /> +<i>Ptenodracon brevirostris</i>, <a href="#Page_64">64</a>, <a href="#Page_99">99</a>, <a href="#Page_167">167</a>, <a href="#Page_169">169</a>, <a href="#Page_192">192</a><br /> +<br /> +Pterodactyle aspects, <a href="#Page_35">35</a>;<br /> +<span style="margin-left: 1em;">avian characters, <a href="#Page_222">222</a>;</span><br /> +<span style="margin-left: 1em;">beak, <a href="#Page_200">200</a>;</span><br /> +<span style="margin-left: 1em;">brain, <a href="#Page_53">53</a>;</span><br /> +<span style="margin-left: 1em;">coracoid, <a href="#Page_113">113</a>;</span><br /> +<span style="margin-left: 1em;">discovery, <a href="#Page_27">27</a>, <a href="#Page_33">33</a>;</span><br /> +<span style="margin-left: 1em;">foot, <a href="#Page_104">104</a>;</span><br /> +<span style="margin-left: 1em;">fore limb, <a href="#Page_117">117</a>;</span><br /> +<span style="margin-left: 1em;">history in Germany, <a href="#Page_31">31</a>, <a href="#Page_148">148</a>;</span><br /> +<span style="margin-left: 1em;">hand, <a href="#Page_130">130</a>;</span><br /> +<span style="margin-left: 1em;">hind limb, <a href="#Page_100">100</a>;</span><br /> +<span style="margin-left: 1em;">long tails, <a href="#Page_156">156</a>;</span><br /> +<span style="margin-left: 1em;">palate, <a href="#Page_71">71</a>;</span><br /> +<span style="margin-left: 1em;">sacrum, <a href="#Page_89">89</a>;</span><br /> +<span style="margin-left: 1em;">short tails, <a href="#Page_165">165</a>;</span><br /> +<span style="margin-left: 1em;">size, <a href="#Page_35">35</a>, <a href="#Page_133">133</a>;</span><br /> +<span style="margin-left: 1em;">sacrum, <a href="#Page_89">89</a>;</span><br /> +<span style="margin-left: 1em;">skull, <a href="#Page_192">192</a>;</span><br /> +<span style="margin-left: 1em;">teeth, <a href="#Page_73">73</a>;</span><br /> +<span style="margin-left: 1em;">vertebræ, <a href="#Page_80">80</a></span><br /> +<br /> +Pterodactyles from Kansas Chalk, <a href="#Page_177">177</a>, <a href="#Page_181">181</a><br /> +<br /> +— from Lias Clay, <a href="#Page_135">135</a>, <a href="#Page_147">147</a>, <a href="#Page_152">152</a><br /> +<br /> +— from Neocomian Sand, <a href="#Page_176">176</a><br /> +<br /> +— from Oxford Clay, <a href="#Page_155">155</a><br /> +<br /> +— from Purbeck beds, <a href="#Page_173">173</a><br /> +<br /> +— from Solenhofen Slate, <a href="#Page_156">156</a>, <a href="#Page_158">158</a><br /> +<br /> +— from Stonesfield Slate, <a href="#Page_153">153</a>, <a href="#Page_158">158</a><br /> +<br /> +Pterodactylia, <a href="#Page_30">30</a>, <a href="#Page_165">165</a>, <a href="#Page_193">193</a>, <a href="#Page_199">199</a><br /> +<br /> +<i>Pterodactylus antiquus</i>, <a href="#Page_167">167</a>;<br /> +<span style="margin-left: 1em;"><i>brevirostris</i>, <a href="#Page_99">99</a>, <a href="#Page_167">167</a>, <a href="#Page_169">169</a>;</span><br /> +<span style="margin-left: 1em;"><i>crassirostris</i>, <a href="#Page_156">156</a>;</span><br /> +<span style="margin-left: 1em;"><i>dubius</i>, <a href="#Page_87">87</a>, <a href="#Page_96">96</a>, <a href="#Page_97">97</a>, <a href="#Page_203">203</a>;</span><br /> +<span style="margin-left: 1em;"><i>elegans</i>, <a href="#Page_169">169</a>;</span><br /> +<span style="margin-left: 1em;"><i>Fraasii</i>, <a href="#Page_169">169</a>;</span><br /> +<span style="margin-left: 1em;"><i>grandipelvis</i>, <a href="#Page_87">87</a>, <a href="#Page_90">90</a>;</span><br /> +<span style="margin-left: 1em;"><i>grandis</i>, <a href="#Page_102">102</a>, <a href="#Page_167">167</a>, <a href="#Page_169">169</a>;</span><br /> +<span style="margin-left: 1em;"><i>Kochi</i>, <a href="#Page_12">12</a>, <a href="#Page_61">61</a>, <a href="#Page_87">87</a>, <a href="#Page_90">90</a>, <a href="#Page_168">168</a>, <a href="#Page_169">169</a>;</span><br /> +<span style="margin-left: 1em;"><i>longirostris</i>, <a href="#Page_28">28</a>, <a href="#Page_90">90</a>, <a href="#Page_96">96</a>, <a href="#Page_101">101</a>, <a href="#Page_103">103</a>, <a href="#Page_105">105</a>, <a href="#Page_167">167</a>, <a href="#Page_169">169</a>;</span><br /> +<span style="margin-left: 1em;"><i>micronyx</i>, <a href="#Page_105">105</a>, <a href="#Page_169">169</a>;</span><br /> +<span style="margin-left: 1em;"><i>rhamphastinus</i>, <a href="#Page_183">183</a>;</span><br /> +<span style="margin-left: 1em;"><i>scolopaciceps</i>, <a href="#Page_105">105</a>, <a href="#Page_166">166</a>;</span><br /> +<span style="margin-left: 1em;"><i>spectabilis</i>, <a href="#Page_83">83</a>;</span><br /> +<span style="margin-left: 1em;"><i>suevicus</i>, <a href="#Page_169">169</a></span><br /> +<br /> +Pterodermata, <a href="#Page_194">194</a>, <a href="#Page_199">199</a><br /> +<br /> +Pteroid bone of first digit, <a href="#Page_121">121</a><br /> +<br /> +Pteromys, <a href="#Page_24">24</a><br /> +<br /> +Pterosauria, <a href="#Page_187">187</a>, <a href="#Page_193">193</a><br /> +<br /> +Pterygoid bones, <a href="#Page_72">72</a>, <a href="#Page_147">147</a><br /> +<br /> +Pythonomorpha, <a href="#Page_191">191</a><br /> +<br /> +<br /> +<b>Q</b><br /> +<br /> +Quadrate bone, <a href="#Page_12">12</a>, <a href="#Page_68">68</a>, <a href="#Page_77">77</a><br /> +<br /> +Quenstedt, <a href="#Page_231">231</a><br /> +<span class='pagenum'><a name="Page_238" id="Page_238">[Pg 238]</a></span><br /> +<br /> +<b>R</b><br /> +<br /> +Rabbit, <a href="#Page_227">227</a><br /> +<br /> +Radius, <a href="#Page_119">119</a>, <a href="#Page_120">120</a><br /> +<br /> +Redshanks, <a href="#Page_22">22</a><br /> +<br /> +Relation between head and tail, <a href="#Page_157">157</a>, <a href="#Page_193">193</a><br /> +<br /> +Reptile, <a href="#Page_6">6</a>, <a href="#Page_79">79</a>, <a href="#Page_80">80</a><br /> +<br /> +Resin, <a href="#Page_136">136</a><br /> +<br /> +Restorations—<br /> +<span style="margin-left: 1em;">Campylognathus, palate of, <a href="#Page_71">71</a></span><br /> +<span style="margin-left: 1em;">Dimorphodon, <a href="#Page_143">143</a>, <a href="#Page_147">147</a>, <a href="#Page_164">164</a></span><br /> +<span style="margin-left: 1em;">Ornithocheirus, <a href="#Page_164">164</a></span><br /> +<span style="margin-left: 1em;">Ornithostoma, <a href="#Page_164">164</a>, <a href="#Page_183">183</a></span><br /> +<span style="margin-left: 1em;">Ptenodracon, <a href="#Page_167">167</a></span><br /> +<span style="margin-left: 1em;">Pterodactylus, <a href="#Page_29">29</a>, <a href="#Page_30">30</a></span><br /> +<span style="margin-left: 1em;">Rhamphocephalus, <a href="#Page_164">164</a></span><br /> +<span style="margin-left: 1em;">Rhamphorhynchus, <a href="#Page_161">161</a>, <a href="#Page_164">164</a></span><br /> +<span style="margin-left: 1em;">Scaphognathus, <a href="#Page_163">163</a></span><br /> +<br /> +Rhacophorus, <a href="#Page_19">19</a><br /> +<br /> +Rhætic beds, <a href="#Page_184">184</a><br /> +<br /> +Rhamphocephalus, <a href="#Page_113">113</a>, <a href="#Page_136">136</a>, <a href="#Page_153">153</a><br /> +<br /> +Rhamphorhynchus, <a href="#Page_118">118</a>, <a href="#Page_192">192</a>;<br /> +<span style="margin-left: 1em;">foot, <a href="#Page_104">104</a>;</span><br /> +<span style="margin-left: 1em;">hind limb, <a href="#Page_99">99</a>;</span><br /> +<span style="margin-left: 1em;">pelvis, <a href="#Page_95">95</a>;</span><br /> +<span style="margin-left: 1em;">sacrum, <a href="#Page_88">88</a>;</span><br /> +<span style="margin-left: 1em;">skull, <a href="#Page_54">54</a>, <a href="#Page_63">63-6</a>, <a href="#Page_69">69</a>;</span><br /> +<span style="margin-left: 1em;">sternum, <a href="#Page_108">108</a>;</span><br /> +<span style="margin-left: 1em;">tail, <a href="#Page_91">91</a>;</span><br /> +<span style="margin-left: 1em;">teeth, <a href="#Page_73">73</a>;</span><br /> +<span style="margin-left: 1em;">tibia and fibula, <a href="#Page_103">103</a>;</span><br /> +<span style="margin-left: 1em;">web-footed, <a href="#Page_105">105</a></span><br /> +<br /> +<i>Rhamphorhynchus curtimanus</i>, <a href="#Page_163">163</a>;<br /> +<span style="margin-left: 1em;"><i>hirundinaceus</i>, <a href="#Page_163">163</a>;</span><br /> +<span style="margin-left: 1em;"><i>longimanus</i>, <a href="#Page_164">164</a>;</span><br /> +<span style="margin-left: 1em;"><i>phyllurus</i>, <a href="#Page_91">91</a>, <a href="#Page_165">165</a></span><br /> +<br /> +Rhinoceros, <a href="#Page_40">40</a>, <a href="#Page_141">141</a><br /> +<br /> +Rhopoladon, <a href="#Page_97">97</a><br /> +<br /> +Rhynchocephala, <a href="#Page_192">192</a><br /> +<br /> +Roc, <a href="#Page_36">36</a><br /> +<br /> +Rochester, <a href="#Page_136">136</a><br /> +<br /> +Running limb, <a href="#Page_38">38</a><br /> +<br /> +Ryle, Bishop, <a href="#Page_17">17</a><br /> +<br /> +<br /> +<b>S</b><br /> +<br /> +Sacrum, <a href="#Page_87">87</a>, <a href="#Page_88">88</a><br /> +<br /> +St. George, <a href="#Page_15">15</a><br /> +<br /> +St. Ives, <a href="#Page_156">156</a><br /> +<br /> +Sarcorhamphus, <a href="#Page_102">102</a><br /> +<br /> +Saurians, <a href="#Page_27">27</a><br /> +<br /> +Saurischia, <a href="#Page_190">190</a>, <a href="#Page_195">195</a>, <a href="#Page_198">198</a>, <a href="#Page_199">199</a><br /> +<br /> +Sauromorpha, <a href="#Page_191">191</a>, <a href="#Page_192">192</a><br /> +<br /> +Sauropsida, <a href="#Page_188">188</a><br /> +<br /> +Sauropterygia, <a href="#Page_192">192</a><br /> +<br /> +Scaphognathus, <a href="#Page_64">64</a>, <a href="#Page_85">85</a>, <a href="#Page_140">140</a>, <a href="#Page_152">152</a>, <a href="#Page_192">192</a>, <a href="#Page_212">212</a><br /> +<br /> +<i>Scaphognathus crassirostris</i>, <a href="#Page_73">73-5</a>, <a href="#Page_83">83</a><br /> +<br /> +Scapular arch, <a href="#Page_111">111</a>, <a href="#Page_113">113</a><br /> +<br /> +Scelidosaurus, <a href="#Page_135">135</a><br /> +<br /> +Sclerotic circle, <a href="#Page_65">65</a><br /> +<br /> +Seals, <a href="#Page_41">41</a><br /> +<br /> +Sedgwick, Professor Adam, v, <a href="#Page_46">46</a><br /> +<br /> +Shillington, <a href="#Page_77">77</a><br /> +<br /> +Shoebill, <a href="#Page_67">67</a><br /> +<br /> +Shoe-shaped prepubic bones, <a href="#Page_204">204</a>, <a href="#Page_205">205</a><br /> +<br /> +Short-tailed pterodactyles, <a href="#Page_165">165</a>, <a href="#Page_193">193</a><br /> +<br /> +Shoulder-girdle, <a href="#Page_107">107</a>, <a href="#Page_111">111</a>, <a href="#Page_114">114</a>, <a href="#Page_115">115</a>, <a href="#Page_183">183</a><br /> +<br /> +Siberia, <a href="#Page_141">141</a><br /> +<br /> +Simultaneous origin of characters, <a href="#Page_214">214</a>, <a href="#Page_224">224</a><br /> +<br /> +Skin covering, <a href="#Page_40">40</a>, <a href="#Page_41">41</a>, <a href="#Page_58">58</a>, <a href="#Page_139">139</a>, <a href="#Page_140">140</a><br /> +<br /> +Skulls, <a href="#Page_68">68</a><br /> +<br /> +Sloth, <a href="#Page_112">112</a><br /> +<br /> +Snipe, <a href="#Page_47">47</a>, <a href="#Page_68">68</a><br /> +<br /> +Solenhofen Slate, <a href="#Page_28">28</a>, <a href="#Page_32">32</a>, <a href="#Page_88">88</a>, <a href="#Page_153">153</a>, <a href="#Page_156">156</a><br /> +<br /> +Sömmerring, <a href="#Page_29">29</a><br /> +<br /> +South African reptiles, <a href="#Page_188">188</a>, <a href="#Page_208">208</a>, <a href="#Page_216">216</a><br /> +<br /> +Spotted fly-catcher, <a href="#Page_47">47</a><br /> +<br /> +Squamosal bone, <a href="#Page_12">12</a>, <a href="#Page_13">13</a><br /> +<br /> +Sternal ribs, <a href="#Page_110">110</a><br /> +<br /> +Sternum, <a href="#Page_107">107</a>, <a href="#Page_158">158</a><br /> +<br /> +Stonesfield Slate, <a href="#Page_33">33</a>, <a href="#Page_88">88</a>, <a href="#Page_153">153</a><br /> +<br /> +Structures common to reptiles, <a href="#Page_224">224</a><br /> +<br /> +Stuttgart Museum, <a href="#Page_32">32</a>, <a href="#Page_172">172</a>, <a href="#Page_203">203</a><br /> +<br /> +Swanage, <a href="#Page_172">172</a><br /> +<br /> +Swan, neck of, <a href="#Page_80">80</a>, <a href="#Page_113">113</a><br /> +<br /> +Swift, <a href="#Page_50">50</a><br /> +<br /> +Swimming limb, <a href="#Page_38">38</a><br /> +<br /> +Synotus, <a href="#Page_25">25</a><br /> +<br /> +Syrinx, <a href="#Page_48">48</a><br /> +<br /> +<br /> +<b>T</b><br /> +<br /> +Tail, description of, <a href="#Page_90">90</a>;<br /> +<span style="margin-left: 1em;">in Cretaceous Pterodactyles, <a href="#Page_193">193</a></span><br /> +<span style="margin-left: 1em;">— long, <a href="#Page_156">156</a>;</span><br /> +<span style="margin-left: 1em;">short, <a href="#Page_166">166</a>;</span><br /> +<span style="margin-left: 1em;">in Dimorphodon, <a href="#Page_145">145</a>;</span><br /> +<span style="margin-left: 1em;">in Ornithocheirus, <a href="#Page_179">179</a></span><br /> +<br /> +Tanystrophœus, long vertebræ in, <a href="#Page_79">79</a><br /> +<br /> +Tarsal bones, <a href="#Page_102">102</a>, <a href="#Page_207">207</a><br /> +<br /> +Tarso-metatarsus, <a href="#Page_128">128</a><br /> +<span class='pagenum'><a name="Page_239" id="Page_239">[Pg 239]</a></span><br /> +Teeth, <a href="#Page_73">73</a>, <a href="#Page_137">137</a>, <a href="#Page_138">138</a>;<br /> +<span style="margin-left: 1em;">in porpoise, <a href="#Page_40">40</a></span><br /> +<br /> +Temperature of blood, <a href="#Page_56">56</a><br /> +<br /> +Temporal arches, <a href="#Page_68">68</a><br /> +<br /> +— bone, <a href="#Page_12">12</a><br /> +<br /> +— fossa, <a href="#Page_67">67</a><br /> +<br /> +Teredo, <a href="#Page_137">137</a><br /> +<br /> +Texas fossils, <a href="#Page_216">216</a><br /> +<br /> +Thecospondylus, <a href="#Page_209">209</a><br /> +<br /> +Theriodont pelvis, <a href="#Page_97">97</a><br /> +<br /> +— reptiles, <a href="#Page_75">75</a>;<br /> +<span style="margin-left: 1em;">of Russia, <a href="#Page_96">96</a>, <a href="#Page_97">97</a>;</span><br /> +<span style="margin-left: 1em;">of South Africa, <a href="#Page_96">96</a>, <a href="#Page_117">117</a></span><br /> +<br /> +Theropsida, <a href="#Page_188">188</a><br /> +<br /> +Thigh bone, <a href="#Page_100">100</a>, <a href="#Page_206">206</a>, <a href="#Page_211">211</a><br /> +<br /> +Three claws, <a href="#Page_146">146</a>, <a href="#Page_197">197</a><br /> +<br /> +Tibia, <a href="#Page_102">102</a>, <a href="#Page_195">195</a>;<br /> +<span style="margin-left: 1em;">in Iguanodon, <a href="#Page_207">207</a></span><br /> +<br /> +Toothless mammals, <a href="#Page_40">40</a><br /> +<br /> +— pterodactyles, <a href="#Page_138">138</a>, <a href="#Page_181">181</a>;<br /> +<span style="margin-left: 1em;">beak of pterodactyles, <a href="#Page_150">150</a></span><br /> +<br /> +Transition from reptiles to birds, <a href="#Page_211">211</a><br /> +<br /> +Tree frogs, <a href="#Page_21">21</a><br /> +<br /> +Trias dinosaurs, <a href="#Page_199">199</a><br /> +<br /> +Triceratops, pelvis of, <a href="#Page_204">204</a><br /> +<br /> +Trout, <a href="#Page_139">139</a>;<br /> +<span style="margin-left: 1em;">of New Zealand, <a href="#Page_228">228</a></span><br /> +<br /> +Tuatera, <a href="#Page_13">13</a><br /> +<br /> +Tübingen Museum, <a href="#Page_32">32</a><br /> +<br /> +Tundras, <a href="#Page_141">141</a><br /> +<br /> +Tunny, <a href="#Page_57">57</a><br /> +<br /> +Turtles, neck bones, <a href="#Page_79">79</a><br /> +<br /> +<br /> +<b>U</b><br /> +<br /> +Ulna, description of, <a href="#Page_119">119</a><br /> +<br /> +Uncinate process of ribs, <a href="#Page_85">85</a><br /> +<br /> +Unlimited time, <a href="#Page_228">228</a><br /> +<br /> +Upper arm bone, <a href="#Page_117">117</a><br /> +<br /> +— Greensand, remains in, <a href="#Page_136">136</a><br /> +<br /> +— Lias of Whitby, <a href="#Page_147">147</a><br /> +<br /> +— Oolites, <a href="#Page_185">185</a>, <a href="#Page_195">195</a><br /> +<br /> +<br /> +<b>V</b><br /> +<br /> +Variation of bones in mammals, <a href="#Page_38">38</a><br /> +<br /> +— in Pterodactyles, <a href="#Page_229">229</a><br /> +<br /> +Variation of bones in vertebræ, <a href="#Page_225">225</a><br /> +<br /> +Vertebræ, caudal, <a href="#Page_89">89</a>, <a href="#Page_92">92</a>, <a href="#Page_203">203</a><br /> +<br /> +— cervical, <a href="#Page_173">173</a>, <a href="#Page_179">179</a>, <a href="#Page_203">203</a><br /> +<br /> +— dorsal, <a href="#Page_86">86</a><br /> +<br /> +Vertebral articulation, <a href="#Page_82">82</a>, <a href="#Page_224">224</a><br /> +<br /> +— column, <a href="#Page_78">78</a><br /> +<br /> +Vulture, neck vertebræ of, <a href="#Page_80">80</a>;<br /> +<span style="margin-left: 1em;">tibia and fibula of, <a href="#Page_102">102</a></span><br /> +<br /> +Vomer, <a href="#Page_147">147</a><br /> +<br /> +Vomerine bones, <a href="#Page_72">72</a><br /> +<br /> +<br /> +<b>W</b><br /> +<br /> +Wagler, <a href="#Page_29">29</a><br /> +<br /> +Wagner, Andreas, <a href="#Page_30">30</a>, <a href="#Page_148">148</a>, <a href="#Page_231">231</a><br /> +<br /> +Walker, J. F., <a href="#Page_54">54</a><br /> +<br /> +Wealden beds, Pterodactyles in, <a href="#Page_55">55</a>, <a href="#Page_84">84</a>;<br /> +<span style="margin-left: 1em;">bones in, <a href="#Page_135">135</a>, <a href="#Page_136">136</a>, <a href="#Page_173">173</a></span><br /> +<br /> +Weight of Pterodactyle, <a href="#Page_106">106</a><br /> +<br /> +Whinchat, <a href="#Page_47">47</a><br /> +<br /> +Whitby, <a href="#Page_33">33</a>, <a href="#Page_135">135</a><br /> +<br /> +Williston, Professor W. S., <a href="#Page_75">75</a>, <a href="#Page_82">82</a>, <a href="#Page_92">92</a>, <a href="#Page_98">98</a>, <a href="#Page_105">105</a>, <a href="#Page_110">110</a><br /> +<br /> +Willow-wren, <a href="#Page_47">47</a><br /> +<br /> +Wing finger, <a href="#Page_116">116</a>, <a href="#Page_130">130</a>, <a href="#Page_133">133</a>, <a href="#Page_151">151</a>, <a href="#Page_178">178</a>, <a href="#Page_197">197</a><br /> +<br /> +— membrane, <a href="#Page_32">32</a>, <a href="#Page_121">121</a>, <a href="#Page_140">140</a>, and <a href="#Fig_47">frontispiece</a><br /> +<br /> +— metacarpal, <a href="#Page_123">123</a>;<br /> +<span style="margin-left: 1em;">in Dimorphodon, <a href="#Page_151">151</a>;</span><br /> +<span style="margin-left: 1em;">in Ornithostoma, <a href="#Page_184">184</a>;</span><br /> +<span style="margin-left: 1em;">in bats, <a href="#Page_131">131</a></span><br /> +<br /> +Wings of Dragons, <a href="#Page_16">16</a><br /> +<br /> +Winkler, T. C., <a href="#Page_231">231</a><br /> +<br /> +Woodwardian Museum, <a href="#Page_34">34</a><br /> +<br /> +Wood-wren, <a href="#Page_47">47</a><br /> +<br /> +Wrist bones, <a href="#Page_122">122</a><br /> +<br /> +Würtemberg, <a href="#Page_33">33</a><br /> +<br /> +<br /> +<b>Y</b><br /> +<br /> +Yale College Museum, <a href="#Page_32">32</a><br /> +<br /> +York Museum, <a href="#Page_34">34</a>, <a href="#Page_176">176</a><br /> +<br /> +<br /> +<b>Z</b><br /> +<br /> +Zittel, Karl von, <a href="#Page_31">31</a>, <a href="#Page_157">157</a>, <a href="#Page_165">165</a>, <a href="#Page_231">231</a><br /> +<br /> +Zygomatic arch, <a href="#Page_67">67</a><br /> +</p></div> + +<p><span class='pagenum'><a name="Page_240" id="Page_240">[Pg 240]</a></span></p> + +<h5> +PRINTED BY<br /> +<br /> +WILLIAM BRENDON AND SON<br /> +<br /> +PLYMOUTH<br /> +</h5> + + + + + + + + +<pre> + + + + + +End of the Project Gutenberg EBook of Dragons of the Air, by H. G. 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