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diff --git a/24800-h/24800-h.htm b/24800-h/24800-h.htm new file mode 100644 index 0000000..6f5e4d1 --- /dev/null +++ b/24800-h/24800-h.htm @@ -0,0 +1,13311 @@ +<!DOCTYPE html PUBLIC "-//W3C//DTD XHTML 1.0 Transitional//EN" + "http://www.w3.org/TR/xhtml1/DTD/xhtml1-transitional.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 Darwin, and After Darwin, Volume I, by George John Romanes. + </title> + <style type="text/css"> +/*<![CDATA[ XML blockout */ +<!-- + a {text-decoration:none; color:blue;} + a:visited {color:gray;} + body {margin-left: 15%; margin-right: 15%;} + h1,h2,h3 {text-align: center; clear: both;} + h3 {margin:0 auto 0 auto;} + hr {width:65%; margin:2em auto 2em auto; clear:both; text-align:center;} + hr.full {width: 100%;} + hr.minor {width: 45%; margin:1em auto 1em auto; clear:both;} + ins.corr {text-decoration:none; border-bottom: thin dotted gray;} + p {margin-top:.75em; text-align:justify; margin-bottom:.75em; text-indent:1.5em;} + p.noin {text-indent:0;} + p.hi {margin-left: 2em; text-indent:-2em;} + table {margin-left:auto; margin-right:auto; empty-cells:show;} + td.c {text-align:center;} + td.r {text-align:right} + td.l {text-align:left} + td.tdc {margin-left: 1.5em; text-indent:-1.5em;} + ul.off {list-style-type:none;} + ul.hi li {margin-left: 2em; text-indent:-2em;} + .b {font-weight:bold;} + .bbox {border:solid 1px; padding:1em; margin:2em 10% 2em 10%;} + .blockquot{margin-left:5%; margin-right:10%;} + .c {text-align:center;} + .caption {font-weight:bold; font-size:75%; text-indent:0;} + .captionh {font-weight:bold; font-size:75%; text-align:left; margin-left: 2em; text-indent:-2em;} + .figcenter {margin:2em auto auto auto; text-align:center;} + .figleft {float:left; clear:left; margin:1em 1em 1em 0; padding:0; text-align:center;} + .figright {float:right; clear:right; margin:1em 0 1em 1em; padding:0; text-align:center;} + .fnanchor {vertical-align:super; font-size:.8em; text-decoration:none;} + .footnote {margin-left:10%; margin-right:10%; font-size:0.9em;} + .footnote p {margin-left: 2em; text-indent:0;} + .footnote .label {position:absolute; right:80%; text-align:right;} + .footnotes {border:dashed 1px;} + .i {font-style:italic;} + .lc {text-transform:lowercase;} + .linenum {position:absolute; top:auto; left:4%;} + .mt2 {margin-top:2em;} + .pagenum {position:absolute; left:95%; font-style:normal; font-size:smaller; text-align:right; text-indent:0;} + .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;} + .ralign {position:absolute; right:20%; text-align:right;} + .sc {font-variant: small-caps;} + .sf30 {font-size:30%;} + .sf50 {font-size:50%;} + .sf75 {font-size:75%;} + .sn {font-style: italic;} /* scientific name */ + .u {text-decoration:underline;} + // --> + /* XML end ]]>*/ + </style> + </head> +<body> + + +<pre> + +The Project Gutenberg EBook of Darwin, and After Darwin (Vol. 1 and 3, of +3), by George John Romanes + +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: Darwin, and After Darwin (Vol. 1 and 3, of 3) + An Exposition of the Darwinian Theory and a Discussion of + Post-Darwinian Questions + +Author: George John Romanes + +Release Date: March 11, 2008 [EBook #24800] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK DARWIN *** + + + + +Produced by Marilynda Fraser-Cunliffe, LN Yaddanapudi and +the Online Distributed Proofreading Team at +http://www.pgdp.net + + + + + + +</pre> + + +<div class="figcenter" style="width: 411px;"> +<img src="images/i_001_frontispiece.jpg" width="369" height="498" alt="Charles Darwin" title="" /></div> + +<p class="pagenum"><a name="page_i" id="page_i">[i]</a></p> +<h1>DARWIN, AND AFTER DARWIN<br /><br /> + +<span class='i sf75'>AN EXPOSITION OF THE DARWINIAN THEORY +AND A DISCUSSION OF +POST-DARWINIAN QUESTIONS</span><br /><br /> + +<span class='sf50'>BY</span><br /><br /> + +<span class='sf75'>GEORGE JOHN ROMANES, M.A., LL.D., F.R.S.</span><br /> +<span class='i sf50'>Honorary Fellow of Gonville and Caius College, Cambridge</span><br /><br /> + +<span class='sf75'>I</span><br /> +<span class='sf75'>THE DARWINIAN THEORY</span><br /> +<span class='sf50'>FOURTH EDITION</span></h1> + +<p class='noin c'>Chicago<br /> +THE OPEN COURT PUBLISHING COMPANY.<br /> +1910</p> + +<hr /><p class="pagenum"><a name="page_ii" id="page_ii">[ii]</a></p> + +<p class='mt2'>The Illustrations of this book (with the exception of the Frontispiece +and the colored plate facing page 332) are copyrighted under the title +“Darwinism Illustrated.”</p> + +<p class='noin'><span class="ralign sc">The Open Court Publishing Co.</span><br /></p> + +<p class='mt2 noin c sf75'>PRESS OF THE<br /> +BLAKELY-OSWALD PRINTING CO.<br /> +CHICAGO</p> + +<hr /><p class="pagenum"><a name="page_iii" id="page_iii">[iii]</a></p> + +<div class="figcenter" style="width: 329px;"> +<img src="images/i_004_letter.jpg" width="329" height="500" alt="Letter from George Romanes to Edward Hageler" title="" /> +<div class='caption' style='text-align:left; font-size:100%;'><pre> + Ch. Ch. Oxford: + March 15th 1892. + + My dear Sir, + + As we have now agreed that +the Open Court Publishing Company is to +undertake the American edition of my +work entitled “Darwin and after +Darwin,” I have much pleasure +in transferring to you the copyright +thereof, with all that this +includes. + +Thanking you very much for +the kindness and liberality which +have marked your conduct of these +negotiations, + + I remain, + Yours very faithfully, + George J. Romanes + +To +Edward C. Hegeler Esq. + La Salle, Ill. U. S.</pre></div></div> + +<hr /><p class="pagenum"><a name="page_v" id="page_v">[v]</a></p> +<h2><a name="PREFACE" id="PREFACE"></a>PREFACE</h2> + +<p>Several years ago Lord Rosebery founded, in the University of Edinburgh, +a lectureship on “The Philosophy of Natural History,” and I +was invited by the Senatus to deliver the lectures. This invitation I +accepted, and subsequently constituted the material of my lectures the +foundation of another course, which was given in the Royal Institution, +under the title “Before and after Darwin.” Here the course +extended over three years—namely from 1888 to 1890. The lectures for +1888 were devoted to the history of biology from the earliest recorded +times till the publication of the “Origin of Species” in +1859; the lectures for 1889 dealt with the theory of organic evolution +up to the date of Mr. Darwin’s death, in 1882; while those of the +third year discussed the further developments of this theory from that +date till the close of the course in 1890.</p> + +<p>It is from these two courses—which resembled each other in +comprising between thirty and forty lectures, but differed largely in +other respects—that the present treatise has grown. Seeing, +however, that it has <span class="pagenum"><a name="page_vi" id="page_vi">[vi]</a></span> +Grown much beyond the bulk of the original lectures, I have thought it +desirable to publish the whole in the form of three separate works. Of +these the first—or that which deals with the purely historical +side of biological science—may be allowed to stand over for an +indefinite time. The second is the one which is now brought out and +which, as its sub-title signifies, is devoted to the general theory of +organic evolution as this was left by the stupendous labours of darwin. +as soon as the translations shall have been completed, the third portion +will follow (probably in the autumn season), under the sub-title, +“post-darwinian questions.”</p> + +<p>As the present volume is thus intended to be merely a systematic +exposition of what may be termed the Darwinism of Darwin, and as on this +account it is likely to prove of more service to general readers than to +professed naturalists, I have been everywhere careful to avoid assuming +even the most elementary knowledge of natural science on the part of +those to whom the exposition is addressed. The case, however, will be +different as regards the next volume, where I shall have to deal with +the important questions touching Heredity, Utility, Isolation, &c., +which have been raised since the death of Mr. Darwin, and which are now +being debated with such salutary vehemence by the best naturalists of +our time.</p> + +<p>My obligations to the Senatus of the University of Edinburgh, and to the +Board of Management of the Royal Institution, have already been +virtually <span class="pagenum"><a name="page_vii" id="page_vii">[vii]</a></span> +expressed; but I should like to take this opportunity of also +expressing my obligations to the students who attended the lectures in +the University of Edinburgh. For alike in respect of their large +numbers, their keen intelligence, and their generous sympathy, the +members of that voluntary class yielded a degree of stimulating +encouragement, without which the labour of preparing the original +lectures could not have been attended with the interest and the +satisfaction that I found in it. My thanks are also due to Mr. R. E. +Holding for the painstaking manner in which he has assisted me in +executing most of the original drawings with which this volume is +illustrated; and likewise to Messrs. Macmillan and Co. for kindly +allowing me to reprint—without special acknowledgment in every +case—certain passages from an essay which they published for me +many years ago, under the title “Scientific Evidences of Organic +Evolution.” Lastly, I must mention that I am indebted to the same +firm for permission to reproduce an excellent portrait of Mr. Darwin, +which constitutes the frontispiece.</p> + +<p class='noin'><span class='ralign'>G. J. R.<br /></span><br /></p> + +<p><span class="sc">Christ Church, Oxford,<br /></span> + <i>April 19th, 1892.</i></p> + +<hr /><p class="pagenum"><a name="page_ix" id="page_ix">[ix]</a></p> + +<h2><a name="CONTENTS" id="CONTENTS"></a>CONTENTS</h2> + +<div class='center'> +<table border="0" cellpadding="4" cellspacing="0" summary=""> +<tr><td class='l'></td><td class='r'>PAGE</td></tr> +<tr><td class='c' colspan='2'><a href="#CHAPTER_I">CHAPTER I.</a></td></tr> +<tr><td class='l'><span class="sc">Introductory</span></td><td class='r'><a href="#page_1">1</a></td></tr> +<tr><td class='c' colspan='2'><a href="#CHAPTER_II">CHAPTER II.</a></td></tr> +<tr><td class='l'><span class="sc">Classification</span></td><td class='r'><a href="#page_23">23</a></td></tr> +<tr><td class='c' colspan='2'><a href="#CHAPTER_III">CHAPTER III.</a></td></tr> +<tr><td class='l'><span class="sc">Morphology</span></td><td class='r'><a href="#page_50">50</a></td></tr> +<tr><td class='c' colspan='2'><a href="#CHAPTER_IV">CHAPTER IV.</a></td></tr> +<tr><td class='l'><span class="sc">Embryology</span></td><td class='r'><a href="#page_98">98</a></td></tr> +<tr><td class='c' colspan='2'><a href="#CHAPTER_V">CHAPTER V.</a></td></tr> +<tr><td class='l'><span class="sc">Palæontology</span></td><td class='r'><a href="#page_156">156</a></td></tr> +<tr><td class='c' colspan='2'><a href="#CHAPTER_VI">CHAPTER VI.</a></td></tr> +<tr><td class='l'><span class="sc">Geographical Distribution</span></td><td class='r'><a href="#page_204">204</a></td></tr> +<tr><td class='c' colspan='2'><a href="#CHAPTER_VII">CHAPTER VII.</a></td></tr> +<tr><td class='l'><span class="sc">The Theory of Natural Selection</span></td><td class='r'><a href="#page_251">251</a></td></tr> +<tr><td class='c' colspan='2'><a href="#CHAPTER_VIII">CHAPTER VIII.</a></td></tr> +<tr><td class='l'><span class="sc">Evidences of the Theory of Natural Selection</span></td><td class='r'><a href="#page_285">285</a></td></tr> +<tr><td class='c' colspan='2'><a href="#CHAPTER_IX">CHAPTER IX.</a><span class="pagenum"><a name="page_x" id="page_x">[x]</a></span></td></tr> +<tr><td class='l'><span class="sc">Criticisms of the Theory of Natural Selection</span></td><td class='r'><a href="#page_333">333</a></td></tr> +<tr><td class='c' colspan='2'><a href="#CHAPTER_X">CHAPTER X.</a></td></tr> +<tr><td class='l'><span class="sc">The Theory of Sexual Selection, and Concluding Remarks</span></td><td class='r'><a href="#page_379">379</a></td></tr> +<tr><td class='c' colspan='2'><hr class='minor' /></td></tr> +<tr><td class='l'><span class="sc"><a href="#APPENDIX_TO_CHAPTER_V">Appendix to Chapter V.</a></span></td><td class='r'><a href="#page_421">421</a></td></tr> +<tr><td class='l'><span class="sc"><a href="#Note_A_to_page_257">Note A to Page 257</a></span></td><td class='r'><a href="#page_443">443</a></td></tr> +<tr><td class='l'><span class="sc"><a href="#Note_B_to_page_295">Note B to Page 295</a></span></td><td class='r'><a href="#page_445">445</a></td></tr> +<tr><td class='l'><span class="sc"><a href="#Note_C_to_page_394">Note C to Page 394</a></span></td><td class='r'><a href="#page_448">448</a></td></tr> +<tr><td class='l'><span class="sc"><a href="#INDEX">Index</a></span></td><td class='r'><a href="#page_451">451</a></td></tr> +</table></div> + +<hr /><p class="pagenum"><a name="page_xi" id="page_xi">[xi]</a></p> +<h2><a name="LIST_OF_ILLUSTRATIONS" id="LIST_OF_ILLUSTRATIONS"></a>LIST OF ILLUSTRATIONS</h2> + +<div class='center'> +<table border="0" cellpadding="4" cellspacing="0" summary=""> +<tr><td class='tdc l'><span class="sc">Fig.</span></td><td class='r'><span class="sc lc">PAGE</span></td></tr> +<tr><td class='tdc l'>1. Successive forms of Paludina, from the Tertiary deposits of Slavonia</td><td class='r'><a href="#page_19">19</a></td></tr> +<tr><td class='tdc l'>2. Skeleton of Seal</td><td class='r'><a href="#page_52">52</a></td></tr> +<tr><td class='tdc l'>3. Skeleton of Greenland Whale</td><td class='r'><a href="#page_53">53</a></td></tr> +<tr><td class='tdc l'>4. Paddle of Whale compared with Hand of Man</td><td class='r'><a href="#page_54">54</a></td></tr> +<tr><td class='tdc l'>5. Wing of Reptile, Mammal, and Bird</td><td class='r'><a href="#page_56">56</a></td></tr> +<tr><td class='tdc l'>6. Skeleton of <span class='sn'>Dinornis gravis</span></td><td class='r'><a href="#page_61">61</a></td></tr> +<tr><td class='tdc l'>7. Hermit crabs compared with the cocoa-nut crab</td><td class='r'><a href="#page_64">64</a></td></tr> +<tr><td class='tdc l'>8. Rudimentary or vestigial hind-limbs of Python</td><td class='r'><a href="#page_67">67</a></td></tr> +<tr><td class='tdc l'>9. <span class='sn'>Apteryx Australis</span></td><td class='r'><a href="#page_69">69</a></td></tr> +<tr><td class='tdc l'>10. Illustrations of the nictitating membrane in various animals named</td><td class='r'><a href="#page_75">75</a></td></tr> +<tr><td class='tdc l'>11. Rudimentary, or vestigial and useless, muscles of the human ear</td><td class='r'><a href="#page_76">76</a></td></tr> +<tr><td class='tdc l'>12. Portrait of a young male gorilla</td><td class='r'><a href="#page_78">78</a></td></tr> +<tr><td class='tdc l'>13. Portrait of a young male child</td><td class='r'><a href="#page_79">79</a></td></tr> +<tr><td class='tdc l'>14. An infant, three weeks old, supporting its own weight</td><td class='r'><a href="#page_81">81</a></td></tr> +<tr><td class='tdc l'>15. Sacrum of Gorilla compared with that of Man, showing the rudimentary tail-bones of each</td><td class='r'><a href="#page_82">82</a></td></tr> +<tr><td class='tdc l'>16. Diagrammatic outline of the human embryo when about seven weeks old</td><td class='r'><a href="#page_83">83</a></td></tr> +<tr><td class='tdc l'>17. Front and back view of adult human sacrum</td><td class='r'><a href="#page_84">84</a></td></tr> +<tr><td class='tdc l'>18. <span class='sn'>Appendix vermiformis</span> in Orang and in Man</td><td class='r'><a href="#page_85">85</a></td></tr> +<tr><td class='tdc l'>19. The same, showing variation in the Orang</td><td class='r'><a href="#page_85">85</a></td></tr> +<tr><td class='tdc l'>20. Human ear</td><td class='r'><a href="#page_86">86</a></td></tr> +<tr><td class='tdc l'>21. Fœtus of an Orang</td><td class='r'><a href="#page_87">87</a></td></tr> +<tr><td class='tdc l'>22. Vestigial characters of human ears</td><td class='r'><a href="#page_88">88</a></td></tr> +<tr><td class='tdc l'>23. Hair-tracts on the arms and hands of Man, as compared with those on the arms and hands of Chimpanzee</td><td class='r'><a href="#page_90">90</a></td></tr> +<tr><td class='tdc l'>24. Molar teeth of lower jaw in Gorilla, Orang, and Man</td><td class='r'><a href="#page_93">93</a></td></tr> +<tr><td class='tdc l'>25. Perforation of the humerus (supra-condyloid foramen) in three species of Quadrumana where it normally occurs, and in Man, where it does not normally occur</td><td class='r'><a href="#page_95">95</a></td></tr> +<tr><td class='tdc l'>26. Antlers of stag, showing successive addition of branches in successive years</td><td class='r'><a href="#page_100">100</a> <span class="pagenum"><a name="page_xii" id="page_xii">[xii]</a></span></td></tr> +<tr><td class='tdc l'>27. Fission of a Protozoön</td><td class='r'><a href="#page_107">107</a></td></tr> +<tr><td class='tdc l'>28. <span class='sn'>Hydra viridis</span>, partly in section</td><td class='r'><a href="#page_111">111</a></td></tr> +<tr><td class='tdc l'>29. Successive stages in the division of the ovum, or egg-cell, of a worm</td><td class='r'><a href="#page_113">113</a></td></tr> +<tr><td class='tdc l'>30. Ovarian ovum of a Mammal</td><td class='r'><a href="#page_121">121</a></td></tr> +<tr><td class='tdc l'>31. Amœboid movements of young egg-cells</td><td class='r'><a href="#page_122">122</a></td></tr> +<tr><td class='tdc l'>32. Human ovum, mature and greatly magnified</td><td class='r'><a href="#page_123">123</a></td></tr> +<tr><td class='tdc l'>33. Stages in the formation of the polar bodies in the ovum of a star-fish</td><td class='r'><a href="#page_125">125</a></td></tr> +<tr><td class='tdc l'>34. Fertilization of the ovum of an echinoderm</td><td class='r'><a href="#page_126">126</a></td></tr> +<tr><td class='tdc l'>35. Fertilization of the ovum of a star-fish</td><td class='r'><a href="#page_127">127</a></td></tr> +<tr><td class='tdc l'>36. Karyokinesis of a typical tissue-cell (epithelium of Salamander)</td><td class='r'><a href="#page_129">129</a></td></tr> +<tr><td class='tdc l'>37. Study of successive changes taking place in the nucleus of an epithelium-cell, preparatory to division of the cell</td><td class='r'><a href="#page_131">131</a></td></tr> +<tr><td class='tdc l'>38. Formation and conjugation of the pronuclei in <span class='sn'>Ascaris megalocephala</span></td><td class='r'><a href="#page_132">132</a>,<br /><a href="#page_133">133</a></td></tr> +<tr><td class='tdc l'>39. Segmentation of ovum</td><td class='r'><a href="#page_135">135</a></td></tr> +<tr><td class='tdc l'>40. The contents of an ovum in an advanced stage of segmentation, drawn in perspective</td><td class='r'><a href="#page_135">135</a></td></tr> +<tr><td class='tdc l'>41. Formation of the gastrula of <span class='sn'>Amphioxus</span></td><td class='r'><a href="#page_137">137</a></td></tr> +<tr><td class='tdc l'>42. Gastrulation</td><td class='r'><a href="#page_138">138</a></td></tr> +<tr><td class='tdc l'>43. Gastrula of a Chalk Sponge</td><td class='r'><a href="#page_139">139</a></td></tr> +<tr><td class='tdc l'>44. <span class='sn'>Prophysema primordiale</span>, an extant gastræa-form</td><td class='r'><a href="#page_140">140</a></td></tr> +<tr><td class='tdc l'>45. Ideal primitive vertebrate, seen from the left side</td><td class='r'><a href="#page_143">143</a></td></tr> +<tr><td class='tdc l'>46. The same in transverse section through the ovaries</td><td class='r'><a href="#page_144">144</a></td></tr> +<tr><td class='tdc l'>47. <span class='sn'>Amphioxus lanceolatus</span></td><td class='r'><a href="#page_145">145</a></td></tr> +<tr><td class='tdc l'>48. <span class='sn'>Balanoglossus</span></td><td class='r'><a href="#page_148">148</a></td></tr> +<tr><td class='tdc l'>49. A large Sea-lamprey (<span class='sn'>Petromyzon marinus</span>)</td><td class='r'><a href="#page_148">148</a></td></tr> +<tr><td class='tdc l'>50. Adult Shark (<span class='sn'>Carcharias melanopterus</span>)</td><td class='r'><a href="#page_149">149</a></td></tr> +<tr><td class='tdc l'>51. Diagram of heart and gill-arches of a fish</td><td class='r'><a href="#page_150">150</a></td></tr> +<tr><td class='tdc l'>52. One gill-arch, with branchial fringe attached</td><td class='r'><a href="#page_150">150</a></td></tr> +<tr><td class='tdc l'>53. Diagram of heart and gill-arches in a lizard</td><td class='r'><a href="#page_150">150</a></td></tr> +<tr><td class='tdc l'>54. Ideal diagram of primitive gill-or aortic-arches</td><td class='r'><a href="#page_151">151</a></td></tr> +<tr><td class='tdc l'>55. The same, modified for a bird</td><td class='r'><a href="#page_151">151</a></td></tr> +<tr><td class='tdc l'>56. The same, modified for a mammal</td><td class='r'><a href="#page_151">151</a></td></tr> +<tr><td class='tdc l'>57. A series of embryos at three comparable and progressive stages of development, representing each of the classes of vertebrated animals below the Mammalia</td><td class='r'><a href="#page_152">152</a></td></tr> +<tr><td class='tdc l'>58. Another series of embryos, also at three comparable and progressive stages of development, representing four different divisions of the class Mammalia</td><td class='r'><a href="#page_153">153</a><span class="pagenum"><a name="page_xiii" id="page_xiii">[xiii]</a></span></td></tr> +<tr><td class='tdc l'>59. Diagram of geological succession of the classes of the Animal Kingdom</td><td class='r'><a href="#page_165">165</a></td></tr> +<tr><td class='tdc l'>60. Skull of <span class='sn'>Oreodon Culbertsoni</span></td><td class='r'><a href="#page_167">167</a></td></tr> +<tr><td class='tdc l'>61,62. Horns of <span class='sn'>Cervus dicrocerus</span></td><td class='r'><a href="#page_168">168</a></td></tr> +<tr><td class='tdc l'>63. Horns of <span class='sn'>C. matheronis</span></td><td class='r'><a href="#page_168">168</a></td></tr> +<tr><td class='tdc l'>64. Horns of <span class='sn'>C. pardinensis</span></td><td class='r'><a href="#page_168">168</a></td></tr> +<tr><td class='tdc l'>65. Horns of <span class='sn'>C. issiodorensis</span></td><td class='r'><a href="#page_168">168</a></td></tr> +<tr><td class='tdc l'>66. Horns of <span class='sn'>C. Sedgwickii</span></td><td class='r'><a href="#page_168">168</a></td></tr> +<tr><td class='tdc l'>67. Successive stages in the development of an existing Deer’s Antlers</td><td class='r'><a href="#page_169">169</a></td></tr> +<tr><td class='tdc l'>68. Homocercal tail</td><td class='r'><a href="#page_169">169</a></td></tr> +<tr><td class='tdc l'>69. Heterocercal tail</td><td class='r'><a href="#page_170">170</a></td></tr> +<tr><td class='tdc l'>70. Vertebrated but symmetrical fin (diphycercal)</td><td class='r'><a href="#page_170">170</a></td></tr> +<tr><td class='tdc l'>71. Tail of <span class='sn'>Archæopteryx</span></td><td class='r'><a href="#page_171">171</a></td></tr> +<tr><td class='tdc l'>72. Tail of modern Bird</td><td class='r'><a href="#page_171">171</a></td></tr> +<tr><td class='tdc l'>73. <span class='sn'>Archæopteryx macura</span>, restored</td><td class='r'><a href="#page_172">172</a></td></tr> +<tr><td class='tdc l'>74. Skeleton of Polar Bear</td><td class='r'><a href="#page_174">174</a></td></tr> +<tr><td class='tdc l'>75. Skeleton of Lion</td><td class='r'><a href="#page_175">175</a></td></tr> +<tr><td class='tdc l'>76. Anterior limb of Man, Dog, Hog, Sheep, and Horse</td><td class='r'><a href="#page_176">176</a></td></tr> +<tr><td class='tdc l'>77. Posterior limb of Man, Monkey, Dog, Sheep, and Horse</td><td class='r'><a href="#page_177">177</a></td></tr> +<tr><td class='tdc l'>78. Posterior limb of <span class='sn'>Baptanodon discus</span>, and anterior limb of <span class='sn'>Chelydra serpentina</span></td><td class='r'><a href="#page_179">179</a></td></tr> +<tr><td class='tdc l'>79. Paddle of a Whale</td><td class='r'><a href="#page_180">180</a></td></tr> +<tr><td class='tdc l'>80. Fossil skeleton of <span class='sn'>Phenacodus primævus</span></td><td class='r'><a href="#page_184">184</a></td></tr> +<tr><td class='tdc l'>81. Bones of the foot of four different forms of the perissodactyl type</td><td class='r'><a href="#page_186">186</a></td></tr> +<tr><td class='tdc l'>82. Bones of the foot of four different forms of the artiodactyl type</td><td class='r'><a href="#page_187">187</a></td></tr> +<tr><td class='tdc l'>83. Feet and teeth In fossil pedigree of the Horse</td><td class='r'><a href="#page_189">189</a></td></tr> +<tr><td class='tdc l'>84. <span class='sn'>Palæotherium</span>. (Lower Tertiary of Paris Basin)</td><td class='r'><a href="#page_190">190</a></td></tr> +<tr><td class='tdc l'>85. <span class='sn'>Hipparion</span>. (New World Pliocene)</td><td class='r'><a href="#page_192">192</a></td></tr> +<tr><td class='tdc l'>86. Comparative series of Brains</td><td class='r'><a href="#page_194">194</a></td></tr> +<tr><td class='tdc l'>87. Ideal section through all the above stages</td><td class='r'><a href="#page_195">195</a></td></tr> +<tr><td class='tdc l'>88. Skulls of Canadian Stag, <span class='sn'>Cervalces Americanus</span>, and Elk</td><td class='r'><a href="#page_198">198</a></td></tr> +<tr><td class='tdc l'>89. Transmutations of <span class='sn'>Planorbis</span></td><td class='r'><a href="#page_200">200</a></td></tr> +<tr><td class='tdc l'>90. Transformation of <span class='sn'>Strombus</span></td><td class='r'><a href="#page_202">202</a></td></tr> +<tr><td class='tdc l'>91. Pigeons. Drawn from life</td><td class='r'><a href="#page_298">298</a><span class="pagenum"><a name="page_xiv" id="page_xiv">[xiv]</a></span></td></tr> +<tr><td class='tdc l'>92. Pigeons (<i>continued</i>). Drawn from life</td><td class='r'><a href="#page_299">299</a></td></tr> +<tr><td class='tdc l'>93. Fowls. Drawn from life</td><td class='r'><a href="#page_300">300</a></td></tr> +<tr><td class='tdc l'>94. Fowls (<i>continued</i>). Drawn from life</td><td class='r'><a href="#page_301">301</a></td></tr> +<tr><td class='tdc l'>95. Pair of Japanese Fowls, long-tailed breed</td><td class='r'><a href="#page_302">302</a></td></tr> +<tr><td class='tdc l'>96. Canaries. Drawn from life</td><td class='r'><a href="#page_303">303</a></td></tr> +<tr><td class='tdc l'>97. Sebastopol, or Frizzled Goose</td><td class='r'><a href="#page_304">304</a></td></tr> +<tr><td class='tdc l'>98. The Dingo, or wild dog of Australia</td><td class='r'><a href="#page_304">304</a></td></tr> +<tr><td class='tdc l'>99. Dogs. Drawn from life</td><td class='r'><a href="#page_305">305</a></td></tr> +<tr><td class='tdc l'>100. Dogs (<i>continued</i>). Drawn from life</td><td class='r'><a href="#page_306">306</a></td></tr> +<tr><td class='tdc l'>101. The Hairless Dog of Japan</td><td class='r'><a href="#page_307">307</a></td></tr> +<tr><td class='tdc l'>102. The skull of a Bull-dog compared with that of a Deer-hound</td><td class='r'><a href="#page_307">307</a></td></tr> +<tr><td class='tdc l'>103. Rabbits. Drawn from life</td><td class='r'><a href="#page_308">308</a></td></tr> +<tr><td class='tdc l'>104. Horses. Drawn from life</td><td class='r'><a href="#page_309">309</a></td></tr> +<tr><td class='tdc l'>105. Sheep. Drawn from life</td><td class='r'><a href="#page_310">310</a></td></tr> +<tr><td class='tdc l'>106. Cattle. Drawn from life</td><td class='r'><a href="#page_311">311</a></td></tr> +<tr><td class='tdc l'>107. Wild Boar contrasted with a modern Domesticated Pig</td><td class='r'><a href="#page_312">312</a></td></tr> +<tr><td class='tdc l'>108. Seasonal changes of colour in Ptarmigan (<span class='sn'>Lagopus mutus</span>)</td><td class='r'><a href="#page_317">317</a></td></tr> +<tr><td class='tdc l'>109.<span class='sn'>Œdicneus crepitans</span>, showing the instinctive attitude of concealment</td><td class='r'><a href="#page_320">320</a></td></tr> +<tr><td class='tdc l'>110. Imitative forms and colours in insects</td><td class='r'><a href="#page_322">322</a></td></tr> +<tr><td class='tdc l'>111. The larva of Puss Moth (<span class='sn'>Cerura vinula</span>)</td><td class='r'><a href="#page_325">325</a></td></tr> +<tr><td class='tdc l'>112. The larva of Puss Moth in disturbed attitude</td><td class='r'><a href="#page_326">326</a></td></tr> +<tr><td class='tdc l'>113. Three cases of mimicry</td><td class='r'><a href="#page_328">328</a></td></tr> +<tr><td class='tdc l'>114. Two further cases of mimicry; flies resembling a wasp in the one and a bee in the other</td><td class='r'><a href="#page_329">329</a></td></tr> +<tr><td class='tdc l'>115. A case of mimicry where a non-venomous species of snake resembles a venomous one</td><td class='r'><a href="#page_330">330</a></td></tr> +<tr><td class='tdc l'>116. A case of mimicry where a homopterous resembles a leaf-cutting ant</td><td class='r'><a href="#page_332">332</a></td></tr> +<tr><td class='tdc l'>117. Feather-footed pigeon</td><td class='r'><a href="#page_359">359</a></td></tr> +<tr><td class='tdc l'>118. <span class='sn'>Raia radiata</span></td><td class='r'><a href="#page_368">368</a></td></tr> +<tr><td class='tdc l'>119. Electric organ of the Skate</td><td class='r'><a href="#page_369">369</a></td></tr> +<tr><td class='tdc l'>120. Electric cells of <span class='sn'>Raia radiata</span></td><td class='r'><a href="#page_370">370</a></td></tr> +<tr><td class='tdc l'>121. The Garden Bower-bird (<span class='sn'>Amblyornis inornata</span>)</td><td class='r'><a href="#page_382">382</a></td></tr> +<tr><td class='tdc l'>122. Courtship of Spiders</td><td class='r'><a href="#page_388">388</a></td></tr> +<tr><td class='tdc l'>123. Courtship of Spiders (<i>continued</i>)</td><td class='r'><a href="#page_389">389</a></td></tr> +<tr><td class='tdc l'>124. The Bell-bird (<span class='sn'>Chasmorhynchus niveus</span>)</td><td class='r'><a href="#page_396">396</a></td></tr> +<tr><td class='tdc l'>125. <span class='sn'>C. tricarunculatus</span></td><td class='r'><a href="#page_397">397</a></td></tr> +</table></div> + +<hr /><p class="pagenum"><a name="page_1" id="page_1">[1]</a></p> +<h2><a name="SECTION_I" id="SECTION_I"></a>SECTION I<br /><br /> +<i>EVOLUTION</i></h2> + +<hr class='minor' /> +<h2><a name="CHAPTER_I" id="CHAPTER_I"></a>CHAPTER I.<br /><br /> +<span class="sc">Introductory.</span></h2> + +<p>Among the many and unprecedented changes that have been wrought by Mr. +Darwin’s work on the <i>Origin of Species</i>, there is one which, +although second in importance to no other, has not received the +attention which it deserves. I allude to the profound modification which +that work has produced on the ideas of naturalists with regard to +method.</p> + +<p>Having had occasion of late years somewhat closely to follow the history +of biological science, I have everywhere observed that progress is not +so much marked by the march of discovery <i>per se</i>, as by the altered +views of method which the march has involved. If we except what +Aristotle called “the first start” in himself, I think one +may fairly say that from the rejuvenescence of biology in the sixteenth +century to the stage of growth which it has now reached in the +nineteenth, there is a direct proportion to be found between the value +of work done and the degree in which the worker has thereby advanced the +true conception of scientific working. Of course, up to a <span class="pagenum"><a name="page_2" id="page_2">[2]</a></span> +certain point, it is notorious that the revolt against the purely +“subjective methods” in the sixteenth century revived the +spirit of <i>inductive</i> research as this had been left by the Greeks; but +even with regard to this revolt there are two things which I should like +to observe.</p> + +<p>In the first place, it seems to me, an altogether disproportionate value +has been assigned to Bacon’s share in the movement. At most, I +think, he deserves to be regarded but as a literary exponent of the +<i>Zeitgeist</i> of his century. Himself a philosopher, as distinguished from +a man of science, whatever influence his preaching may have had upon the +general public, it seems little short of absurd to suppose that it could +have produced any considerable effect upon men who were engaged in the +practical work of research. And those who read the <i>Novum Organon</i> with +a first-hand knowledge of what is required for such research can +scarcely fail to agree with his great contemporary Harvey, that he wrote +upon science like a Lord Chancellor.</p> + +<p>The second thing I should like to observe is, that as the revolt against +the purely subjective methods grew in extent and influence it passed to +the opposite extreme, which eventually became only less deleterious to +the interests of science than was the bondage of authority, and +addiction to <i>a priori</i> methods, from which the revolt had set her free. +For, without here waiting to trace the history of this matter in detail, +I think it ought now to be manifest to everyone who studies it, that up +to the commencement of the present century the progress of science in +general, and of natural history in particular, was seriously retarded by +<span class="pagenum"><a name="page_3" id="page_3">[3]</a></span> +what may be termed the Bugbear of Speculation. Fully awakened to the +dangers of web-spinning from the ever-fertile resources of their own +inner consciousness, naturalists became more and more abandoned to the +idea that their science ought to consist in a mere observation of facts, +or tabulation of phenomena, without attempt at theorizing upon their +philosophical import. If the facts and phenomena presented any such +import, that was an affair for men of letters to deal with; but, as men +of science, it was <i>their</i> duty to avoid the seductive temptations of +the world, the flesh, and the devil, in the form of speculation, +deduction, and generalization.</p> + +<p>I do not allege that this ideal of natural history was either absolute +or universal; but there can be no question that it was both orthodox and +general. Even Linnæus was express in his limitations of true scientific +work in natural history to the collecting and arranging of species of +plants and animals. In accordance with this view, the <i>status</i> of a +botanist or a zoologist was estimated by the number of specific names, +natural habitats, &c., which he could retain in his memory, rather than +by any evidences which he might give of intellectual powers in the way +of constructive thought. At the most these powers might legitimately +exercise themselves only in the direction of taxonomic work; and if a +Hales, a Haller, or a Hunter obtained any brilliant results in the way +of observation and experiment, their merit was taken to consist in the +discovery of facts <i>per se</i>: not in any endeavours they might make in +the way of combining their facts under general principles. Even as late +in the day as Cuvier this ideal was upheld as the strictly <span class="pagenum"><a name="page_4" id="page_4">[4]</a></span> +legitimate one for a naturalist to follow; and although Cuvier himself +was far from being always loyal to it, he leaves no doubt regarding the +estimate in which he held the still greater deviations of his +colleagues, St. Hilaire and Lamarck.</p> + +<p>Now, these traditional notions touching the severance between the facts +of natural history and the philosophy of it, continued more or less to +dominate the minds of naturalists until the publication of the <i>Origin +of Species</i>, in 1859. Then it was that an epoch was marked in this +respect, as in so many other respects where natural history is +concerned. For, looking to the enormous results which followed from a +deliberate disregard of such traditional canons by Darwin, it has long +since become impossible for naturalists, even of the strictest sect, not +to perceive that their previous bondage to the law of a mere ritual has +been for ever superseded by what verily deserves to be regarded as a new +dispensation. Yet it cannot be said, or even so much as suspected, that +Darwin’s method in any way resembled that of pre-scientific days, +the revolt against which led to the straight-laced—and for a long time +most salutary—conceptions of method that we have just been noticing. +Where, then, is the difference? To me it seems that the difference is as +follows; and, if so, that not the least of our many obligations to +Darwin as the great organizer of biological science arises from his +having clearly displayed the true principle which ought to govern +biological research.</p> + +<p>To begin with, he nowhere loses sight of the primary distinction between +fact and theory; so that, thus far, he loyally follows the spirit of +revolt against subjective methods. But, while always holding this +<span class="pagenum"><a name="page_5" id="page_5">[5]</a></span> +distinction clearly in view, his idea of the scientific use of facts is +plainly that of furnishing legitimate material for the construction of +theories. Natural history is not to him an affair of the herbarium or +the cabinet. The collectors and the species-framers are, as it were, his +diggers of clay and makers of bricks: even the skilled observers and the +trained experimentalists are his mechanics. Valuable as the work of all +these men is in itself, its principal value, as he has finally +demonstrated, is that which it acquires in rendering possible the work +of the architect. Therefore, although he has toiled in all the trades +with his own hands, and in each has accomplished some of the best work +that has ever been done, the great difference between him and most of +his predecessors consists in this,—that while to them the discovery or +accumulation of facts was an end, to him it is the means. In their eyes +it was enough that the facts should be discovered and recorded. In his +eyes the value of facts is due to their power of guiding the mind to a +further discovery of principles. And the extraordinary success which +attended his work in this respect of <i>generalization</i> immediately +brought natural history into line with the other inductive sciences, +behind which, in this most important of all respects, she has so +seriously fallen. For it was the <i>Origin of Species</i> which first clearly +revealed to naturalists as a class, that it was the duty of their +science to take as its motto, what is really the motto of natural +science in general,</p> + +<p class='noin c'>Felix qui potuit rerum cognoscere causas.</p> + +<p class='noin'>Not facts, then, or phenomena, but causes or principles, are the +ultimate objects of scientific quest. It remains to ask, How ought this +quest to be prosecuted?</p> + +<p><span class="pagenum"><a name="page_6" id="page_6">[6]</a></span> +Well, in the second place, Darwin has shown that next only to the +importance of clearly distinguishing between facts and theories on the +one hand, and of clearly recognising the relation between them on the +other, is the importance of not being scared by the Bugbear of +Speculation. The spirit of speculation is the same as the spirit of +science, namely, as we have just seen, a desire to know the causes of +things. The <i>hypotheses non fingo</i> of Newton, if taken to mean what it +is often understood as meaning, would express precisely the opposite +spirit from that in which all scientific research must necessarily take +its origin. For if it be causes or principles, as distinguished from +facts or phenomena, that constitute the final aim of scientific +research, obviously the advancement of such research can be attained +only by the framing of hypotheses. And to frame hypotheses is to +speculate.</p> + +<p>Therefore, the difference between science and speculation is not a +difference of spirit; nor, thus far, is it a difference of method. The +only difference between them is in the subsequent process of verifying +hypotheses. For while speculation, in its purest form, is satisfied to +test her explanations only by the degree in which they accord with our +subjective ideas of probability—or with the “Illative +Sense” of Cardinal Newman,—science is not satisfied to rest in +any explanation as final until it shall have been fully verified by an +appeal to objective proof. This distinction is now so well and so +generally appreciated that I need not dwell upon it. Nor need I wait to +go into any details with regard to the so-called canons of verification. +My only object is to make perfectly clear, first, that <span class="pagenum"><a name="page_7" id="page_7">[7]</a></span> +in order to have any question to put to the test of objective +verification, science must already have so far employed the method of +speculation as to have framed a question to be tested; and, secondly, +that the point where science parts company with speculation is the point +where this testing process begins.</p> + +<p>Now, if these things are so, there can be no doubt that Darwin was +following the truest method of inductive research in allowing any amount +of latitude to his speculative thought in the direction of scientific +theorizing. For it follows from the above distinctions that the danger +of speculation does not reside in the width of its range, or even in the +impetuosity of its vehemence. Indeed, the wider its reach, and the +greater its energy, the better will it be for the interests of science. +The only danger of speculation consists in its momentum being apt to +carry away the mind from the more laborious work of adequate +verification; and therefore a true scientific judgment consists in +giving a free rein to speculation on the one hand, while holding ready +the break of verification with the other. Now, it is just because Darwin +did both these things with so admirable a judgment, that he gave the +world of natural history so good a lesson as to the most effectual way +of driving the chariot of science.</p> + +<p>This lesson we have now all more or less learnt to profit by. Yet no +other naturalist has proved himself so proficient in holding the balance +true. For the most part, indeed, they have now all ceased to confound +the process of speculation <i>per se</i> with the danger of inadequate +verification; and therefore the old ideal of natural history as +concerned merely with collecting species, classifying affinities, and, +in general, tabulating <span class="pagenum"><a name="page_8" id="page_8">[8]</a></span> +Facts, has been well-nigh universally superseded. But this great gain +has been attended by some measure of loss. For while not a few +naturalists have since erred on the side of insufficiently +distinguishing between fully verified principles of evolution and merely +speculative deductions therefrom, a still larger number have formed for +themselves a darwinian creed, and regard any further theorizing on the +subject of evolution as <i>ipso facto</i> unorthodox.</p> + +<p>Having occupied the best years of my life in closely studying the +literature of Darwinism, I shall endeavour throughout the following +pages to avoid both these extremes. No one in this generation is able to +imitate Darwin, either as an observer or a generalizer. But this does +not hinder that we should all so far endeavour to follow his <i>method</i>, +as always to draw a clear distinction, not merely between observation +and deduction, but also between degrees of verification. At all events, +my own aim will everywhere be to avoid dogmatism on the one hand, and +undue timidity as regards general reasoning on the other. For everything +that is said justification will be given; and, as far as prolonged +deliberation has enabled me to do so, the exact value of such +justification will be rendered by a statement of at least the main +grounds on which it rests. The somewhat extensive range of the present +treatise, however, will not admit of my rendering more than a small +percentage of the facts which in each case go to corroborate the +conclusion. But although a great deal must thus be necessarily lost on +the one side, I am disposed to think that more will be gained on the +other, by presenting, in a terser form than would otherwise be +<span class="pagenum"><a name="page_9" id="page_9">[9]</a></span> +possible, the whole theory of organic evolution as I believe that it +will eventually stand. My endeavour, therefore, will be to exhibit the +general structure of this theory in what I take to be its strictly +logical form, rather than to encumber any of its parts by a lengthy +citation of facts. Following this method, I shall in each case give only +what I consider the main facts for and against the positions which have +to be argued; and in most cases I shall arrange the facts in two +divisions, namely, first those of largest generality, and next a few of +the most special character that can be found.</p> + +<p>As explained in the Preface, the present instalment of the treatise is +concerned with the theory of evolution, from the appearance of the +<i>Origin of Species</i> in 1859, to the death of its author in 1882; while +the second part will be devoted to the sundry post-Darwinian questions +which have arisen in the subsequent decade. To the possible criticism +that a disproportionate amount of space will thus be allotted to a +consideration of these post-Darwinian questions, I may furnish in +advance the following reply.</p> + +<p>In the first place, besides the works of Darwin himself, there are a +number of others which have already and very admirably expounded the +evidences, both of organic evolution as a fact, and of natural selection +as a cause. Therefore, in the present treatise it seemed needless to go +beyond the ground which was covered by my original lectures, namely, a +condensed and connected, while at the same time a critical statement of +the main evidences, and the main objections, which have thus far been +published with reference to the distinctively Darwinian theory. +<span class="pagenum"><a name="page_10" id="page_10">[10]</a></span> +Indeed while re-casting this portion of my lectures for the present +publication, I have felt that criticism might be more justly urged from +the side of impatience at a reiteration of facts and arguments already +so well known. But while endeavouring, as much as possible, to avoid +overlapping the previous expositions, I have not carried this attempt to +the extent of damaging my own, by omitting any of the more important +heads of evidence; and I have sought to invest the latter with some +measure of novelty by making good what appears to me a deficiency which +has hitherto obtained in the matter of pictorial illustration. In +particular, there will be found a tolerably extensive series of +woodcuts, serving to represent the more important products of artificial +selection. These, like all the other original illustrations, have been +drawn either direct from nature or from a comparative study of the best +authorities. Nevertheless, I desire it to be understood that the first +part of this treatise is intended to retain its original character, as a +merely educational exposition of Darwinian teaching—an exposition, +therefore, which, in its present form, may be regarded as a compendium, +or hand-book, adapted to the requirements of a general reader, or +biological student as distinguished from those of a professed +naturalist.</p> + +<p>The case, however, is different with the second instalment, which will +be published at no very distant date. Here I have not followed with +nearly so much closeness the material of my original lectures. On the +contrary, I have had in view a special class of readers; and, although I +have tried not altogether to sacrifice the more general class, I shall +desire it to be <span class="pagenum"><a name="page_11" id="page_11">[11]</a></span> +understood that I am there appealing to naturalists who are specialists +in Darwinism. One must say advisedly, naturalists who are specialists in +Darwinism, because, while the literature of Darwinism has become a +department of science in itself, there are nowadays many naturalists +who, without having paid any close attention to the subject, deem +themselves entitled to hold authoritative opinions with regard to it. +These men may have done admirable work in other departments of natural +history, and yet their opinions on such matters as we shall hereafter +have to consider may be destitute of value. As there is no necessary +relation between erudition in one department of science and soundness of +judgment in another, the mere fact that a man is distinguished as a +botanist or zoologist does not in itself qualify him as a critic where +specially Darwinian questions are concerned. Thus it happens now, as it +happened thirty years ago, that highly distinguished botanists and +zoologists prove themselves incapable as judges of general reasoning. It +was Darwin’s complaint that for many years nearly all his +scientific critics either could not, or would not, understand what he +had written—and this even as regarded the fundamental principles +of his theory, which with the utmost clearness he had over and over +again repeated. Now the only difference between such naturalists and +their successors of the present day is, that the latter have grown up in +a Darwinian environment, and so, as already remarked, have more or less +thoughtlessly adopted some form of Darwinian creed. But this scientific +creed is not a whit less dogmatic and intolerant than was the more +theological one which it <span class="pagenum"><a name="page_12" id="page_12">[12]</a></span> +has supplanted; and while it usually incorporates the main elements of +Darwin’s teaching, it still more usually comprises gross +perversions of their consequences. All this I shall have occasion more +fully to show in subsequent parts of the present work; and allusion is +made to the matter here merely for the sake of observing that in future +I shall not pay attention to unsupported expressions of opinion from any +quarter: I shall consider only such as are accompanied with some +statement of the grounds upon which the opinion is held. And, even as +thus limited, I do not think it will be found that the following +exposition devotes any disproportional amount of attention to the +contemporary movements of Darwinian thought, seeing, as we shall see, +how active scientific speculation has been in the field of Darwinism +since the death of Mr. Darwin.</p> + +<hr class='minor' /> + +<p>Leaving, then, these post-Darwinian questions to be dealt with +subsequently, I shall now begin a systematic <i>résumé</i> of the evidences +in favour of the Darwinian theory, as this was left to the world by +Darwin himself.</p> + +<p>There is a great distinction to be drawn between the fact of evolution +and the manner of it, or between the evidence of evolution as having +taken place somehow, and the evidence of the causes which have been +concerned in the process. This most important distinction is frequently +disregarded by popular writers on Darwinism; and, therefore, in order to +mark it as strongly as possible, I will effect a complete separation +between the evidence which we have of evolution as a fact, and the +evidence which we have <span class="pagenum"><a name="page_13" id="page_13">[13]</a></span> +as to its method. In other words, not until I shall have fully +considered the evidence of organic evolution as a process which somehow +or another <i>has</i> taken place, will I proceed to consider <i>how</i> it has +taken place, or the causes which Darwin and others have suggested as +having probably been concerned in this process.</p> + +<p>Confining, then, our attention in the first instance to a proof of +evolution considered as a fact, without any reference at all to its +method, let us begin by considering the antecedent standing of the +matter.</p> + +<hr class='minor' /> + +<p>First of all we must clearly recognise that there are only two +hypotheses in the field whereby it is possible so much as to suggest an +explanation of the origin of species. Either all the species of plants +and animals must have been supernaturally created, or else they must +have been naturally evolved. There is no third hypothesis possible; for +no one can rationally suggest that species have been eternal.</p> + +<p>Next, be it observed, that the theory of a continuous transmutation of +species is not logically bound to furnish a full explanation of <i>all</i> +the natural causes which it may suppose to have been at work. The +radical distinction between the two theories consists in the one +assuming an immediate action of some supernatural or inscrutable cause, +while the other assumes the immediate action of natural—and therefore +of possibly discoverable—causes. But in order to sustain this latter +assumption, the theory of descent is under no logical necessity to +furnish a full proof of <i>all</i> the natural causes which may have been +concerned in working out the observed results. We do not <span class="pagenum"><a name="page_14" id="page_14">[14]</a></span> +know the natural causes of many diseases; but yet no one nowadays +thinks of reverting to any hypothesis of a supernatural cause, in order +to explain the occurrence of any disease the natural causation of which +is obscure. The science of medicine being in so many cases able to +explain the occurrence of disease by its hypothesis of natural causes, +medical men now feel that they are entitled to assume, on the basis of a +wide analogy, and therefore on the basis of a strong antecedent +presumption, that all diseases are due to natural causes, whether or not +in particular cases such causes happen to have been discovered. And from +this position it follows that medical men are not logically bound to +entertain any supernatural theory of an obscure disease, merely because +as yet they have failed to find a natural theory. And so it is with +biologists and their theory of descent. Even if it be fully proved to +them that the causes which they have hitherto discovered, or suggested, +are inadequate to account for all the facts of organic nature, this +would in no wise logically compel them to vacate their theory of +evolution, in favour of the theory of creation. All that it would so +compel them to do would be to search with yet greater diligence for the +natural causes still undiscovered, but in the existence of which they +are, by their independent evidence in favour of the theory, bound to +believe.</p> + +<p>In short, the issue is not between the theory of a supernatural cause +and the theory of any one particular natural cause, or set of +causes—such as natural selection, use, disuse, and so forth. The issue +thus far—or where only the <i>fact</i> of evolution is concerned—is between +the theory of a supernatural cause as <span class="pagenum"><a name="page_15" id="page_15">[15]</a></span> +operating immediately in numberless acts of special creation, and the +theory of natural causes as a whole, whether these happen, or do not +happen, to have been hitherto discovered.</p> + +<p>This much by way of preliminaries being understood, we have next to +notice that whichever of the two rival theories we choose to entertain, +we are not here concerned with any question touching the origin of life. +We are concerned only with the origin of particular forms of life—that +is to say, with the origin of species. The theory of descent starts from +life as a <i>datum</i> already granted. How life itself came to be, the +theory of descent, as such, is not concerned to show. Therefore, in the +present discussion, I will take the existence of life as a fact which +does not fall within the range of our present discussion. No doubt the +question as to the origin of life is in itself a deeply interesting +question, and although in the opinion of most biologists it is a +question which we may well hope will some day fall within the range of +science to answer, at present, it must be confessed, science is not in a +position to furnish so much as any suggestion upon the subject; and +therefore our wisdom as men of science is frankly to acknowledge that +such is the case.</p> + +<hr class='minor' /> + +<p>We are now in a position to observe that the theory of organic evolution +is strongly recommended to our acceptance on merely antecedent grounds, +by the fact that it is in full accordance with what is known as the +principle of continuity. By the principle of continuity is meant the +uniformity of nature, in virtue of which the many and varied processes +going on in nature are due to the same kind of method, i. e. the method +of <span class="pagenum"><a name="page_16" id="page_16">[16]</a></span> +natural causation. This conception of the uniformity of nature is one +that has only been arrived at step by step through a long and arduous +course of human experience in the explanation of natural phenomena. The +explanations of such phenomena which are first given are always of the +supernatural kind; it is not until investigation has revealed the +natural causes which are concerned that the hypotheses of superstition +give way to those of science. Thus it follows that the hypotheses of +superstition which are the latest in yielding to the explanations of +science, are those which refer to the more recondite cases of natural +causation; for here it is that methodical investigation is longest in +discovering the natural causes. Thus it is only by degrees that +fetishism is superseded by what now appears a common-sense +interpretation of physical phenomena; that exorcism gives place to +medicine; alchemy to chemistry; astrology to astronomy; and so forth. +Everywhere the miraculous is progressively banished from the field of +explanation by the advance of scientific discovery; and the places where +it is left longest in occupation are those where the natural causes are +most intricate or obscure, and thus present the greatest difficulty to +the advancing explanations of science. Now, in our own day there are but +very few of these strongholds of the miraculous left. Nearly the whole +field of explanation is occupied by naturalism, so that no one ever +thinks of resorting to supernaturalism except in the comparatively few +cases where science has not yet been able to explore the most obscure +regions of causation. One of these cases is the origin of life; and, +until quite recently, another of these cases was the origin of +<span class="pagenum"><a name="page_17" id="page_17">[17]</a></span> +Species. But now that a very reasonable explanation of the origin of +species has been offered by science, it is but in accordance with all +previous historical analogies that many minds should prove themselves +unable all at once to adjust themselves to the new ideas, and thus still +linger about the more venerable ideas of supernaturalism. But we are now +in possession of so many of these historical analogies, that all minds +with any instincts of science in their composition have grown to +distrust, on merely antecedent grounds, any explanation which embodies a +miraculous element. Such minds have grown to regard all these +explanations as mere expressions of our own ignorance of natural +causation; or, in other words, they have come to regard it as an <i>a +priori</i> truth that nature is everywhere uniform in respect of method or +causation; that the reign of law universal; the principle of continuity +ubiquitous.</p> + +<p>Now, it must be obvious to any mind which has adopted this attitude of +thought, that the scientific theory of natural descent is recommended by +an overwhelming weight of antecedent presumption, as against the +dogmatic theory of supernatural design.</p> + +<p>To begin with, we must remember that the fact of evolution—or, which is +the same thing, the fact of continuity in natural causation—has now +been unquestionably proved in so many other and analogous departments of +nature, that to suppose any interruption of this method as between +species and species becomes, on grounds of such analogy alone, well-nigh +incredible. For example, it is now a matter of demonstrated fact that +throughout the range of <i>inorganic</i> nature the principles of evolution +have obtained. It is no longer <span class="pagenum"><a name="page_18" id="page_18">[18]</a></span> +possible for any one to believe with our forefathers that the +earth’s surface has always existed as it now exists. For the +science of geology has proved to demonstration that seas and lands are +perpetually undergoing gradual changes of relative +positions—continents and oceans supplanting each other in the +course of ages, mountain-chains being slowly uplifted, again as slowly +denuded, and so forth. Moreover, and as a closer analogy, within the +limits of animate nature we know it is the universal law that every +individual life undergoes a process of gradual development; and that +breeds, races, or strains, may be brought into existence by the +intentional use of natural processes—the results bearing an +unmistakeable resemblance to what we know as natural species. Again, +even in the case of natural species themselves, there are two +considerations which present enormous force from an antecedent point of +view. The first is that organic forms are only then recognised as +species when intermediate forms are absent. If the intermediate forms +are actually living, or admit of being found in the fossil state, +naturalists forthwith regard the whole series as varieties, and name all +the members of it as belonging to the same species. Consequently it +becomes obvious that naturalists, in their work of naming species, may +only have been marking out the cases where intermediate or connecting +forms have been lost to observation. For example, here we have a diagram +representing a very unusually complete series of fossil shells, which +within the last few years has been unearthed from the Tertiary lake +basins of Slavonia. Before the series was completed, some six or eight +of the then <span class="pagenum"><a name="page_19" id="page_19">[19]</a></span> +disconnected forms were described as distinct species; but as soon as +the connecting forms were found—showing a progressive modification +from the older to the newer beds,—the whole were included as +varieties of one species.</p> + +<div class="figcenter" style="width: 481px;"> +<img src="images/i_036_001.jpg" width="481" height="500" alt="Successive forms of Paludina." title="" /> +<div class="caption"><span class='sc'>Fig.</span> 1.—Successive forms of Paludina, from the Tertiary +deposits of Slavonia (after Neumayr).</div></div> + +<p>Of course, other cases of the same kind might be adduced, and therefore, +as just remarked, in their work of naming species naturalists may only +have been marking out the cases where intermediate forms <span class="pagenum"><a name="page_20" id="page_20">[20]</a></span> +have been lost to observation. And this possibility becomes little less +than a certainty when we note the next consideration which I have to +adduce, namely, that in all their systematic divisions of plants and +animals in groups higher than species—such as genera, families, +orders, and the rest—naturalists have at all times recognised the +fact that the one shades off into the other by such imperceptible +gradations, that it is impossible to regard such divisions as other than +conventional. It is important to remember that this fact was fully +recognised before the days of Darwin. In those days the scientifically +orthodox doctrine was, that although species were to be regarded as +fixed units, bearing the stamp of a special creation, all the higher +taxonomic divisions were to be considered as what may be termed the +artificial creation of naturalists themselves. In other words, it was +believed, and in many cases known, that if we could go far enough back +in the history of the earth, we should everywhere find a tendency to +mutual approximation between allied <i>groups of species</i>; so that, for +instance, birds and reptiles would be found to be drawing nearer and +nearer together, until eventually they would seem to become fused in a +single type; that the existing distinctions between herbivorous and +carnivorous mammals would be found to do likewise; and so on with all +the larger group-distinctions, at any rate within the limits of the same +sub-kingdoms. But although naturalists recognised this even in the +pre-Darwinian days, they stoutly believed that a great exception was to +be made in the case of species. These, the lowest or initial members of +their taxonomic series, they supposed to be permanent—the <span class="pagenum"><a name="page_21" id="page_21">[21]</a></span> +miraculously created units of organic nature. Now, all that I have at +present to remark is, that this pre-Darwinian exception which was made +in favour of species to the otherwise recognised principle of gradual +change, was an exception which can at no time have been recommended by +any antecedent considerations. At all times it stood out of analogy with +the principle of continuity; and, as we shall fully find in subsequent +chapters, it is now directly contradicted by all the facts of biological +science.</p> + +<p>There remains one other fact of high generality to which prominent +attention should be drawn from the present, or merely antecedent, point +of view. On the theory of special creation no reason can be assigned why +distinct specific types should present any correlation, either in time +or in space, with their nearest allies; for there is evidently no +conceivable reason why any given species, A, should have been specially +created on the same area and at about the same time as its nearest +representative, B,—still less, of course, that such should be a general +rule throughout all the thousands and millions of species which have +ever inhabited the earth. But, equally of course, on the theory of a +natural evolution this is so necessary a consequence, that if no +correlation of such a two-fold kind were observable, the theory would be +negatived. Thus the question whether there be any indication of such a +two-fold correlation may be regarded as a test-question as between the +two theories; for although the vast majority of extinct species have +been lost to science, there are a countless number of existing species +which furnish ample material for answering the question. And the answer +is so unequivocal <span class="pagenum"><a name="page_22" id="page_22">[22]</a></span> +that Mr. Wallace, who is one of our greatest authorities on +geographical distribution, has laid it down as a general law, applicable +to all the departments of organic nature, that, so far as observation +can extend, “every species has come into existence coincident both +in space and time with a pre-existing and closely allied species.” +As it appears to me that the significance of these words cannot be +increased by any comment upon them, I will here bring this introductory +chapter to a close.</p> + +<hr /><p class="pagenum"><a name="page_23" id="page_23">[23]</a></p> +<h2><a name="CHAPTER_II" id="CHAPTER_II"></a>CHAPTER II.<br /><br /> +<span class="sc">Classification.</span></h2> + +<p>The first line of direct evidence in favour of organic evolution which I +shall open is that which may be termed the argument from Classification.</p> + +<p>It is a matter of observable fact that different forms of plants and +animals present among themselves more or less pronounced resemblances. +From the earliest times, therefore, it has been the aim of philosophical +naturalists to classify plants and animals in accordance with these +resemblances. Of course the earliest attempts at such classification +were extremely crude. The oldest of these attempts with which we are +acquainted—namely, that which is presented in the books of Genesis and +Leviticus—arranges the whole vegetable kingdom in three simple +divisions of Grass, Herbs, and Trees; while the animal kingdom is +arranged with almost equal simplicity with reference, first to habitats +in water, earth, or air, and next as to modes of progression. These, of +course, were what may be termed common-sense classifications, having +reference merely to external appearances and habits of life. But when +Aristotle laboriously investigated the comparative anatomy of animals, +he could not fail to perceive that their entire structures had to be +taken into account in <span class="pagenum"><a name="page_24" id="page_24">[24]</a></span> +Order to classify them scientifically; and, also, that for this purpose +the internal parts were of quite as much importance as the external. +indeed, he perceived that they were of greatly more importance in this +respect, inasmuch as they presented so many more points for comparison; +and, in the result, he furnished an astonishingly comprehensive, as well +as an astonishingly accurate classification of the larger groups of the +animal kingdom. On the other hand, classification of the vegetable +kingdom continued pretty much as it had been left by the book of +genesis—all plants being divided into three groups, herbs, shrubs, +and trees. Nor was this primitive state of matters improved upon till +the sixteenth century, when gesner (1516-1565), and still more +cæsalpino (1519-1603), laid the foundations of systematic botany.</p> + +<p>But the more that naturalists prosecuted their studies on the anatomy of +plants and animals, the more enormously complex did they find the +problem of classification become. Therefore they began by forming what +are called artificial systems, in contradistinction to natural systems. +An artificial system of classification is a system based on the more or +less arbitrary selection of some one part, or set of parts; while a +natural classification is one that is based upon a complete knowledge of +all the structures of all the organisms which are classified.</p> + +<p>Thus, the object of classification has been that of arranging organisms +in accordance with their natural affinities, by comparing organism with +organism, for the purpose of ascertaining which of the constituent +organs are of the most invariable occurrence, and therefore of the most +typical signification. A porpoise, <span class="pagenum"><a name="page_25" id="page_25">[25]</a></span> +for instance, has a large number of teeth, and in this feature +resembles most fish, while it differs from all mammals. But it also +gives suck to its young. Now, looking to these two features alone, +should we say that a porpoise ought to be classed as a fish or as a +mammal? Assuredly as a mammal; because the number of teeth is a very +variable feature both in fish and mammals, whereas the giving of suck is +an invariable feature among mammals, and occurs nowhere else in the +animal kingdom. This, of course, is chosen as a very simple +illustration. Were all cases as obvious, there would be but little +distinction between natural and artificial systems of classification. +But it is because the lines of natural affinity are, as it were, so +interwoven throughout the organic world, and because there is, in +consequence, so much difficulty in following them, that artificial +systems have to be made in the first instance as feelers towards +eventual discovery of the natural system. In other words, while forming +their artificial systems of classification, it has always been the aim +of naturalists—whether consciously or unconsciously—to admit +as the bases of their systems those characters which, in the then state +of their knowledge, seemed most calculated to play an important part in +the eventual construction of the natural system. If we were dealing with +the history of classification, it would here be interesting to note how +the course of it has been marked by gradual change in the principles +which naturalists adopted as guides to the selection of characters on +which to found their attempts at a natural classification. Some of these +changes, indeed, I shall have to mention later on; but at present what +has to be specially noted is, <span class="pagenum"><a name="page_26" id="page_26">[26]</a></span> +that through all these changes of theory or principle, and through all +the ever-advancing construction of their taxonomic science, naturalists +themselves were unable to give any intelligible reason for the faith +that was in them—or the faith that over and above the artificial +classifications which were made for the mere purpose of cataloguing the +living library of organic nature, there was deeply hidden in nature +itself a truly natural classification, for the eventual discovery of +which artificial systems might prove to be of more or less assistance.</p> + +<p>Linnæus, for example, expressly says—“You ask me for the +characters of the natural orders; I confess that I cannot give +them.” Yet he maintains that, although he cannot define the +characters, he knows, by a sort of naturalist’s instinct, what in +a general way will subsequently be found to be the organs of most +importance in the eventual grouping of plants under a natural system. +“I will not give my reasons for the distribution of the natural +orders which I have published,” he said: “you, or some other +person, after twenty or after fifty years, will discover them, and see +that I was right.”</p> + +<p>Thus we perceive that in forming their provisional or artificial +classifications, naturalists have been guided by an instinctive belief +in some general principle of natural affinity, the character of which +they have not been able to define; and that the structures which they +selected as the bases of their classifications when these were +consciously artificial, were selected because it seemed that they were +the structures most likely to prove of use in subsequent attempts at +working out the natural system.</p> + +<p><span class="pagenum"><a name="page_27" id="page_27">[27]</a></span> +This general principle of natural affinity, of which all naturalists +have seen more or less well-marked evidence in organic nature, and after +which they have all been feeling, has sometimes been regarded as +natural, but more often as supernatural. Those who regarded it as +supernatural took it to consist in a divine ideal of creation according +to types, so that the structural affinities of organisms were to them +expressions of an archetypal plan, which might be revealed in its +entirety when all organisms on the face of the earth should have been +examined. Those, on the other hand, who regarded the general principle +of affinity as depending on some natural causes, for the most part +concluded that these must have been utilitarian causes; or, in other +words, that the fundamental affinities of structure must have depended +upon fundamental requirements of function. According to this view, the +natural classification would eventually be found to stand upon a basis +of physiology. Therefore all the systems of classification up to the +earlier part of the present century went upon the apparent axiom, that +characters which are of most importance to the organisms presenting them +must be characters most indicative of natural affinities. But the truth +of the matter was eventually found to be otherwise. For it was +eventually found that there is absolutely no correlation between these +two things; that, therefore, it is a mere chance whether or not organs +which are of importance to organisms are likewise of importance as +guides to classification; and, in point of fact, that the general +tendency in this matter is towards an inverse instead of a direct +proportion. More often than not, the greater the value of a structure +for the purpose of <span class="pagenum"><a name="page_28" id="page_28">[28]</a></span> +indicating natural affinities, the less is its value to the creatures +presenting it.</p> + +<p>Enough has now been said to show three things. First, that long before +the theory of descent was entertained by naturalists, naturalists +perceived the fact of natural affinities, and did their best to +construct a natural system of classification for the purpose of +expressing such affinities. Second, that naturalists had a kind of +instinctive belief in some one principle running through the whole +organic world, which thus served to bind together organisms in groups +subordinate to groups—that is, into species, genera, orders, families, +classes, sub-kingdoms, and kingdoms. Third, that they were not able to +give any very intelligible reason for this faith that was in them; +sometimes supposing the principle in question to be that of a +supernatural plan of organization, sometimes regarding it as dependent +on conditions of physiology, and sometimes not attempting to account for +it at all.</p> + +<p>Of course it is obvious that the theory of descent furnishes the +explanation which is required. For it is now evident to evolutionists, +that although these older naturalists did not know what they were doing +when they were tracing these lines of natural affinity, and thus helping +to construct a natural classification—I say it is now evident to +evolutionists that these naturalists were simply tracing the lines of +genetic relationship. The great principle pervading organic nature, +which was seen so mysteriously to bind the whole creation together as in +a nexus of organic affinity, is now easily understood as nothing more or +less than the principle of Heredity. Let us, therefore, look a little +more closely at the character of this network, in <span class="pagenum"><a name="page_29" id="page_29">[29]</a></span> +order to see how far it lends itself to this new interpretation.</p> + +<p>The first thing that we have to observe about the nexus is, that it is a +nexus—not a single line, or even a series of parallel lines. In other +words, some time before the theory of descent was seriously entertained, +naturalists for the most part had fully recognised that it was +impossible to arrange either plants or animals, with respect to their +mutual affinities, in a ladder-like series (as was supposed to be the +type of classification by the earlier systematists), or even in map-like +groups (as was supposed to be the type by Linnæus). And similarly, also, +with respect to grades of organization. In the case of the larger +groups, indeed, it is usually possible to say that the members of this +group as a whole are more highly organized than the members of that +group as a whole; so that, for instance, we have no hesitation in +regarding the Vertebrata as more highly organized than the Invertebrata, +Birds than Reptiles, and so on. But when we proceed to smaller +subdivisions, such as genera and species, it is usually impossible to +say that the one type is more highly organized than another type. A +horse, for instance, cannot be said to be more highly organized than a +zebra or an ass; although the entire horse-genus is clearly a more +highly organized type than any genus of animal which is not a mammal.</p> + +<p>In view of these facts, therefore, the system of classification which +was eventually arrived at before the days of Darwin, was the system +which naturalists likened to a tree; and this is the system which all +naturalists now agreed upon as the true one. According to this system, a +short trunk may be taken <span class="pagenum"><a name="page_30" id="page_30">[30]</a></span> +to represent the lowest organisms which cannot properly be termed +either plants or animals. This short trunk soon separates into two large +trunks, one of which represents the vegetable and the other the animal +kingdom. Each of these trunks then gives off large branches signifying +classes, and these give off smaller, but more numerous branches, +signifying families, which ramify again into orders, genera, and finally +into the leaves, which may be taken to represent species. Now, in such a +representative tree of life, the height of any branch from the ground +may be taken to indicate the grade of organization which the leaves, or +species, present; so that, if we picture to ourselves such a tree, we +may understand that while there is a general advance of organization +from below upwards, there are many deviations in this respect. Sometimes +leaves growing on the same branch are growing at a different +level—especially, of course, if the branch be a large one, +corresponding to a class or sub-kingdom. And sometimes leaves growing on +different branches are growing at the same level: that is to say, +although they represent species belonging to widely divergent families, +orders, or even classes, it cannot be said that the one species is more +highly organized than the other.</p> + +<p>Now, this tree-like arrangement of species in nature is an arrangement +for which Darwin is not responsible. For, as we have seen, the detecting +of it has been due to the progressive work of naturalists for centuries +past; and even when it was detected, at about the commencement of the +present century, naturalists were confessedly unable to explain the +reason of it, or what was the underlying principle that they were +<span class="pagenum"><a name="page_31" id="page_31">[31]</a></span> +engaged in tracing when they proceeded ever more and more accurately to +define these ramifications of natural affinity. But now, as just +remarked, we can clearly perceive that this underlying principle was +none other than Heredity as expressed in family +likeness,—likeness, therefore, growing progressively more unlike +with remoteness of ancestral relationship. For thus only can we obtain +any explanation of the sundry puzzles and apparent paradoxes, which a +working out of their natural classifications revealed to botanists and +zoologists during the first half of the present century. It will now be +my endeavour to show how these puzzles and paradoxes are all explained +by the theory that natural affinities are merely the expression of +genetic affinities.</p> + +<p>First of all, and from the most general point of view, it is obvious +that the tree-like system of classification, which Darwin found already +and empirically worked out by the labours of his predecessors, is as +suggestive as anything could well be of the fact of genetic +relationship. For this is the form that every tabulation of family +pedigree must assume; and therefore the mere fact that a scientific +tabulation of natural affinities was eventually found to take the form +of a tree, is in itself highly suggestive of the inference that such a +tabulation represents a <i>family</i> tree. If all species were separately +created, there can be no assignable reason why the ideas of earlier +naturalists touching the form which a natural classification would +eventually assume should not have represented the truth—why, for +example, it should not have assumed the form of a ladder (as was +anticipated in the seventeenth century), or of a map (as was anticipated +in <span class="pagenum"><a name="page_32" id="page_32">[32]</a></span> +the eighteenth), or, again, of a number of wholly unrelated lines, +circles, &c. (as certain speculative writers of the present century +have imagined). But, on the other hand, if all species were separately +and independently created, it becomes virtually incredible that we +should everywhere observe this progressive arborescence of characters +common to larger groups into more and more numerous, and more and more +delicate, ramifications of characters distinctive only of smaller and +smaller groups. A man would be deemed insane if he were to attribute the +origin of every branch and every twig of a real tree to a separate act +of special creation; and although we have not been able to witness the +growth of what we may term in a new sense the Tree of Life, the +structural relations which are now apparent between its innumerable +ramifications bear quite as strong a testimony to the fact of their +having been due to an organic growth, as is the testimony furnished by +the branches of an actual tree.</p> + +<p>Or, to take another illustration. Classification of organic forms, as +Darwin, Lyell, and Häckel have pointed out, strongly resembles the +classification of languages. In the case of languages, as in the case of +species, we have genetic affinities strongly marked; so that it is +possible to some extent to construct a Language-tree, the branches of +which shall indicate, in a diagrammatic form, the progressive divergence +of a large group of languages from a common stock. For instance, Latin +may be regarded as a fossil language, which has given rise to a group of +living languages—Italian, Spanish, French, and, to a large extent, +English. Now what would be thought of a philologist who should maintain +that English, French, Spanish, <span class="pagenum"><a name="page_33" id="page_33">[33]</a></span> +and Italian were all specially created languages—or languages +separately constructed by the Deity, and by as many separate acts of +inspiration communicated to the nations which now speak them—and +that their resemblance to the fossil form, Latin, must be attributed to +special design? Yet the evidence of the natural transmutation of species +is in one respect much stronger than that of the natural transmutation +of languages—in respect, namely, of there being a vastly greater +number of cases all bearing testimony to the fact of genetic +relationship.</p> + +<p>But, quitting now this most general point of view—or the suggestive +fact that what we have before us is a <i>tree</i>—let us next approach this +tree for the purpose of examining its structure more in detail. When we +do this, the fact of next greatest generality which we find is as +follows.</p> + +<p>In cases where a very old form of life has continued to exist +unmodified, so that by investigation of its anatomy we are brought back +to a more primitive type of structure than that of the newer forms +growing higher up <i>upon the same branch</i>, two things are observable. In +the first place, the old form is less differentiated than the newer +ones; and, in the next place, it is seen much more closely to resemble +types of structure belonging to some of the other and larger branches of +the tree. The organization of the older form is not only <i>simpler</i>; but +it is, as naturalists say, more <i>generalized</i>. It comprises within +itself characters belonging to its own branch, and also characters +belonging to neighbouring branches, or to the trunk from which allied +branches spring. Hence it becomes a general rule of classification, that +it is by the lowest, <span class="pagenum"><a name="page_34" id="page_34">[34]</a></span> +or by the oldest, forms of any two natural groups that the affinities +between the two groups admit of being best detected. And it is obvious +that this is just what ought to be the case on the theory of descent +with divergent modification; while, upon the alternative theory of +special creation, no reason can be assigned why the lowest or the oldest +types should thus combine the characters which afterwards become +severally distinctive of higher or newer types.</p> + +<p>Again, I have already alluded to the remarkable fact that there is no +correlation between the value of structures to the organisms which +present them, and their value to the naturalist for the purpose of +tracing natural affinity; and I have remarked that up to the close of +the last century it was regarded as an axiom of taxonomic science, that +structures which are of most importance to the animals or plants +possessing them must likewise prove of most importance in any natural +system of classification. On this account, all attempts to discover the +natural classification went upon the supposition that such a direct +proportion must obtain—with the result that organs of most +physiological importance were chosen as the bases of systematic work. +And when, in the earlier part of the present century, De Candolle found +that instead of a direct there was usually an inverse proportion between +the functional and the taxonomic value of a structure, he was unable to +suggest any reason for this apparently paradoxical fact. For, upon the +theory of special creation, no reason can be assigned why organs of +least importance to organisms should prove of most importance as marks +of natural affinity. But on the theory of descent with progressive +modification <span class="pagenum"><a name="page_35" id="page_35">[35]</a></span> +the apparent paradox is at once explained. For it is evident that +organs of functional importance are, other things equal, the organs +which are most likely to undergo different modifications in different +lines of family descent, and therefore in time to have their genetic +relationships in these different lines obscured. On the other hand, +organs or structures which are of no functional importance are never +called upon to change in response to any change of habit, or to any +change in the conditions of life. They may, therefore, continue to be +inherited through many different lines of family descent, and thus +afford evidence of genetic relationship where such evidence fails to be +given by any of the structures of vital importance, which in the course +of many generations have been required to change in many ways according +to the varied experiences of different branches of the same family. +Here, then, we have an empirically discovered rule in the science of +classification, the <i>raison d’être</i> of which we are at once +able to appreciate upon the theory of evolution, whereas no possible +explanation of why it should ever have become a rule could be furnished +upon the theory of special creation.</p> + +<p>Here, again, is another empirically determined rule. The larger the +<i>number</i>, as distinguished from the <i>importance</i>, of structures which +are found common to different groups, the greater becomes their value as +guides to the determination of natural affinity. Or, as Darwin puts it, +“the value of an aggregate of characters, even when none are +important, alone explains the aphorism enunciated by Linnæus, namely, +that the characters do not give the genus, <span class="pagenum"><a name="page_36" id="page_36">[36]</a></span> +but the genus gives the characters; for this seems founded on the +appreciation of many trifling points of resemblance, too slight to be +defined<a name="FNanchor_1_1" id="FNanchor_1_1"></a><a href="#Footnote_1_1" class="fnanchor">[1]</a>.”</p> + +<p>Now it is evident, without comment, of how much value aggregates of +characters ought to be in classification, if the ultimate meaning of +classification be that of tracing lines of pedigree; whereas, if this +ultimate meaning were that of tracing divine ideals manifested in +special creation, we can see no reason why single characters are not +such sure tokens of a natural arrangement as are aggregates of +characters, even though the latter be in every other respect +unimportant. For, on the special creation theory, we cannot explain why +an assemblage, say of four or five trifling characters, should have been +chosen to mark some unity of plan, rather than some one character of +functional importance, which would have served at least equally well any +such hypothetical purpose. On the other hand, as Darwin remarks, +“we care not how trifling a character may be—let it be the mere +inflection of the angle of the jaw, the manner in which an +insect’s wing is folded, whether the skin be covered with hair or +feathers—if it prevail throughout many and different species, +especially those having very different habits of life, it assumes high +value; for we can account for its presence in so many forms, with such +different habits, only by inheritance from a common parent. We may err +in this respect in regard to single points of structure, but when +several characters, let them be ever so trifling, concur throughout a +large group of beings having different habits, we may feel almost sure, +on the theory of descent, <span class="pagenum"><a name="page_37" id="page_37">[37]</a></span> +that these characters have been inherited from a common ancestor; and +we know that such aggregated characters have especial value in +classification<a name="FNanchor_2_2" id="FNanchor_2_2"></a><a href="#Footnote_2_2" class="fnanchor">[2]</a>.”</p> + +<p>It is true that even a single character, if found common to a large +number of forms, while uniformly absent from others, is also regarded by +naturalists as of importance for purposes of classification, although +they recognise it as of a value subordinate to that of aggregates of +characters. But this also is what we should expect on the theory of +descent. If even any one structure be found to run through a number of +animals presenting different habits of life, the readiest explanation of +the fact is to be found in the theory of descent; but this does not +hinder that if several such characters always occur together, the +inference of genetic relationship is correspondingly confirmed. And the +fact that before this inference was ever drawn, naturalists recognised +the value of single characters in proportion to their constancy, and the +yet higher value of aggregates of characters in proportion to their +number—this fact shows that in their work of classification naturalists +empirically observed the effects of a cause which we have now +discovered, to wit, hereditary transmission of characters through +ever-widening groups of changing species.</p> + +<p>There is another argument which appears to tell strongly in favour of +the theory of descent. We have just seen that non-adaptive structures, +not being required to change in response to change of habits or +conditions of life, are allowed to persist unchanged through many +generations, and thus furnish exceptionally good guides in the science +of classification—or, <span class="pagenum"><a name="page_38" id="page_38">[38]</a></span> +according to our theory, in the work of tracing lines of pedigree. But +now, the converse of this statement holds equally true. For it often +happens that adaptive structures are required to change in different +lines of descent in analogous ways, in order to meet analogous needs; +and, when such is the case, the structures concerned have to assume more +or less close resemblances to one another, even though they have +severally descended from quite different ancestors. The paddles of a +whale, for instance, most strikingly resemble the fins of a fish as to +their outward form and movements; yet, on the theory of descent, they +must be held to have had a widely different parentage. Now, in all such +cases where there is thus what is called an analogous (or adaptive) +resemblance, as distinguished from what is called an homologous (or +anatomical) resemblance—in all such cases it is observable that +the similarities do not extend further into the structure of the parts +than it is necessary that they should extend, in order that the +structures should both perform the same functions. The whole anatomy of +the paddles of a whale is quite unlike that of the fins of a +fish—being, in fact, that of the fore-limb of a mammal. The +change, therefore, which the fore-limb has here undergone to suit it to +the aquatic habits of this mammal, is no greater than was required for +that purpose: the change has not extended to any one feature of +<i>anatomical</i> significance. This, of course, is what we should expect on +the theory of descent with modification of ancestral characters; but on +the theory of special creation it is not intelligible why there should +always be so marked a distinction between resemblances as analogical or +<span class="pagenum"><a name="page_39" id="page_39">[39]</a></span> +adaptive, and resemblances as homological or of meaning in reference to +a natural classification. To take another and more detailed instance, +the Tasmanian wolf is an animal separated from true wolves in a natural +system of classification. Yet its jaws and teeth bear a strong general +resemblance to those of all the dog tribe, although there are +differences of anatomical detail. In particular, while the dogs all have +on each side of the upper jaw four pre-molars and two molars, the +Tasmanian wolf has three pre-molars and four molars. Now there is no +reason, so far as their common function of dealing with flesh is +concerned, why the teeth of the Tasmanian wolf should not have resembled +homologically as well as analogically the teeth of a true wolf; and +therefore we cannot assign any intelligible reason why, if all the +species of the dog genus were separately created with one pattern of +teeth, the unallied Tasmanian wolf should have been furnished with what +is practically the same pattern from a functional point of view, while +differing from a structural point of view. But, of course, on the theory +of descent with modification, we can well understand why similarities of +habit should have led to similarities of structural appearance of an +adaptive kind in different lines of descent, without there being any +trace of such real or anatomical similarities as could possibly point to +genetic relationship.</p> + +<p>Lastly, to adduce the only remaining argument from classification which +I regard as of any considerable weight, naturalists have found it +necessary, while constructing their natural classifications, to set +great store on what Mr. Darwin calls “chains of +affinities.<span class="pagenum"><a name="page_40" id="page_40">[40]</a></span>” +Thus, for instance, “nothing can be easier than to define +a number of characters common to all birds; but with crustaceans any +such definition has hitherto been found impossible. There are +crustaceans at the opposite ends of the series, which have hardly a +character in common; yet the species at both ends, from being plainly +allied to others, and these to others, and so onwards, can be recognised +as unequivocally belonging to this, and to no other class of the +articulata<a name="FNanchor_3_3" id="FNanchor_3_3"></a><a href="#Footnote_3_3" class="fnanchor">[3]</a>.” +Now it is evident that this progressive modification of specific +types—where it cannot be said that the continuity of resemblance +is anywhere broken, and yet terminates in modification so great that but +for the connecting links no one could divine a natural relationship +between the extreme members of the series,—it is evident that such +chains of affinity speak most strongly in favour of a transmutation of +the species concerned, while it is impossible to suggest any explanation +of the fact in terms of the rival theory. For if all the links of such a +chain were separately forged by as many acts of special creation, we can +see no reason why B should resemble A, C resemble B, and so on, but with +ever slight though accumulating differences, until there is no +resemblance at all between A and Z.</p> + +<hr class='minor' /> + +<p>I hope enough has now been said to show that all the general principles +and particular facts appertaining to the natural classification of +plants and animals, are precisely what they ought to be according to the +theory of genetic descent; while no one of them is such as might +be—and, indeed, used to be—expected <span class="pagenum"><a name="page_41" id="page_41">[41]</a></span> +upon the theory of special creation. Therefore, the only possible way +in which all this uniform body of direct evidence can be met by a +supporter of the latter theory, is by falling back upon the argument +from ignorance. We do not know, it may be said, what hidden reasons +there may have been for following all these general principles in the +separate creation of specific types. Now, it is evident that this is a +form of argument which admits of being brought against all the +actual—and even all the possible—lines of evidence in favour +of evolution. Therefore I deem it desirable thus early in our +proceedings to place this argument from ignorance on its proper logical +footing.</p> + +<p>If there were any independent evidence in favour of special creation as +a <i>fact</i>, then indeed the argument from ignorance might be fairly used +against any sceptical cavils regarding the <i>method</i>. In this way, for +example, Bishop Butler made a legitimate use of the argument from +ignorance when he urged that it is no reasonable objection against a +revelation, <i>otherwise accredited</i>, to show that it has been rendered in +a form, or after a method, which we should not have antecedently +expected. But he could not have legitimately employed this argument, +except on the supposition that he had some independent evidence in +favour of the revelation; for, in the absence of any such independent +evidence, appeal to the argument from ignorance would have become a mere +begging of the question, by simply assuming that a revelation had been +made. And thus it is in the present case. A man, of course, may quite +legitimately say, <i>Assuming that the theory of special creation is +true</i>, it is not for us to anticipate the <span class="pagenum"><a name="page_42" id="page_42">[42]</a></span> +form or method of the process. But where the question is as to whether +or not the theory <i>is</i> true, it becomes a mere begging of this question +to take refuge in the argument from ignorance, or to represent in effect +that there is no question to be discussed. And if, when the form or +method is investigated, it be found everywhere charged with evidence in +favour of the theory of descent, the case becomes the same as that of a +supposed revelation, which has been discredited by finding that all +available evidence points to a natural growth. In short, the argument +from ignorance is in any case available only as a negative foil against +destructive criticism: in no case has it any positive value, or value of +a constructive kind. Therefore, if a theory on any subject is destitute +of positive evidence, while some alternative theory is in possession of +such evidence, the argument from ignorance can be of no logical use to +the former, even though it maybe of such use to the latter. For it is +only the possession of positive evidence which can furnish a logical +justification of the argument from ignorance: in the absence of such +evidence, even the negative value of the argument disappears, and it +then implies nothing more than the gratuitous assumption of a theory.</p> + +<hr class='minor' /> + +<p>I will now sum up the various considerations which have occupied us +during the present chapter.</p> + +<p>First of all we must take note that the classification of plants and +animals in groups subordinate to groups is not merely arbitrary, or +undertaken only for a matter of convenience and nomenclature—such, for +instance, as the classification of stars in constellations. On the +contrary, the classification of a naturalist <span class="pagenum"><a name="page_43" id="page_43">[43]</a></span> +differs from that of an astronomer, in that the objects which he has to +classify present structural resemblances and structural differences in +numberless degrees; and it is the object of his classification to +present a tabular statement of these facts. Now, long before the theory +of evolution was entertained, naturalists became fully aware that these +facts of structural resemblances running through groups subordinate to +groups were really facts of nature, and not merely poetic imaginations +of the mind. No one could dissect a number of fishes without perceiving +that they were all constructed on one anatomical pattern, which differed +considerably from the equally uniform pattern on which all mammals were +constructed, even although some mammals bore an extraordinary +resemblance to fish in external form and habits of life. And similarly +with all the smaller divisions of the animal and vegetable kingdoms. +Everywhere investigation revealed the bonds of close structural +resemblances between species of the same genus, resemblance less close +between genera of the same family, resemblance still less close between +families of the same order, resemblance yet more remote between orders +of the same class, and resemblance only in fundamental features between +classes of the same sub-kingdom, beyond which limit all anatomical +resemblance was found to disappear—the different sub-kingdoms +being formed on wholly different patterns. Furthermore, in tracing all +these grades of structural relationship, naturalists were slowly led to +recognise that the form which a natural classification must eventually +assume would be that of a tree, wherein the constituent branches would +<span class="pagenum"><a name="page_44" id="page_44">[44]</a></span> +display a progressive advance of organization from below upwards.</p> + +<p>Now we have seen that although this tree-like arrangement of natural +groups was as suggestive as anything could well be of all the forms o£ +life being bound together by the ties of genetic relationship, such was +not the inference which was drawn from it. Dominated by the theory of +special creation, naturalists either regarded the resemblance of type +subordinate to type as expressive of divine ideals manifested in such +creation, or else contented themselves with investigating the facts +without venturing to speculate upon their philosophical import. But even +those naturalists who abstained from committing themselves to any theory +of archetypal plans, did not doubt that facts so innumerable and so +universal must have been due to some one co-ordinating principle—that, +even though they were not able to suggest what it was, there must have +been some hidden bond of connexion running through the whole of organic +nature. Now, as we have seen, it is manifest to evolutionists that this +hidden bond can be nothing else than heredity; and, therefore, that +these earlier naturalists, although they did not know what they were +doing, were really tracing the lines of genetic descent as revealed by +degrees of structural resemblance,—that the <ins class='corr' +title="Transcriber’s Note: The original showed +‘arboresent’.">arborescent</ins> grouping of organic forms which +their labours led them to begin, and in large measure to execute, was in +fact a family tree of life.</p> + +<p>Here, then, is the substance of the argument from classification. The +mere fact that all organic nature thus incontestably lends itself to a +natural arrangement <span class="pagenum"><a name="page_45" id="page_45">[45]</a></span> +of group subordinate to group, when due regard is paid to degrees of +anatomical resemblance—this mere fact of itself tells so weightily +in favour of descent with progressive modification in different lines, +that even if it stood alone it would be entitled to rank as one of our +strongest pieces of evidence. But, as we have seen, it does not stand +alone. When we look beyond this large and general fact of all the +innumerable forms of life being thus united in a tree-like system by an +unquestionable relationship of some kind, to those smaller details in +the science of classification which have been found most useful as +guides for this kind of research, then we find that all these details, +or empirically discovered rules, are exactly what we should have +expected them to be, supposing the real meaning of classification to +have been that of tracing lines of pedigree.</p> + +<p>In particular, we have seen that the most archaic types are both simpler +in their organization and more generalized in their characters than are +the more recent types—a fact of which no explanation can be given on +the theory of special creation. But, upon the theory of natural +evolution, we can without difficulty understand why the earlier forms +should have been the simpler forms, and also why they should have been +the most generalized. For it is out of the older forms that the newer +must have grown; and, as they multiplied, they must have become more and +more differentiated.</p> + +<p>Again, we have seen that there is no correlation between the importance +of any structure from a classificatory point of view, and the importance +of that structure to the organism which presents it. On the <span class="pagenum"><a name="page_46" id="page_46">[46]</a></span> +contrary, it is a general rule that “the less any part of the +organization is concerned with special habits, the more important it +becomes for classification.” Now, from the point of view of +special creation it is unintelligible why unity of ideal should be most +manifested by least important structures, whereas from the point of view +of evolution it is to be expected that these life-serving structures +should have been most liable to divergent modification in divergent +lines of descent, or in adaptation to different conditions of life, +while the trivial or less important characters should have been allowed +to remain unmodified. Thus we can now understand why all primitive +classifications were wrong in principle when they went upon the +assumption that divine ideals were best exhibited by resemblances +between life-serving (and therefore adaptive) structures, with the +result that whales were classed with fishes, birds with bats, and so on. +Nevertheless, these primitive naturalists were quite logical; for, from +the premises furnished by the theory of special creation, it is much +more reasonable to expect that unity of ideal should be shown in plainly +adaptive characters than in trivial and more or less hidden anatomical +characters. Moreover, long after biological science had ceased +consciously to follow any theological theory, the apparent axiom +continued to be entertained, that structures of most importance to +organisms must also be structures of most importance to systematists. +And when at last, in the present century, this was found not to be the +case, no reason could be suggested why it was not the case. But now we +are able fully to explain this apparent anomaly.</p> + +<p>Once more, we have seen that aggregates of <span class="pagenum"><a name="page_47" id="page_47">[47]</a></span> +characters presenting resemblances to one another have always been +found to be of special importance as guides to classification. This, of +course, is what we should have expected, if the real meaning of +classification be that of tracing lines of pedigree; but on the theory +of special creation no reason can be assigned why single characters are +not such sure tokens of a natural arrangement as are aggregates of +characters, however trivial the latter may be. For it is obvious that +unity of ideal might have been even better displayed by everywhere +maintaining the pattern of some one important structure, than by doing +so in the case of several unimportant structures. Take an analogous +instance from human contrivances. Unity of ideal in the case of +gun-making would be shown by the same principles of mechanism running +through all the different sizes and shapes of gun-locks, rather than by +the ornamental patterns engraved upon the outside. Yet it must be +supposed that in the mechanisms assumed to have been constructed by +special creation, it was the trivial details rather than the fundamental +principles of these mechanisms which were chosen by the Divinity to +display his ideals.</p> + +<p>And this leads us to the next consideration—namely, that when in two +different lines of descent animals happen to adopt similar habits of +life, the modifications which they undergo in order to fit them for +these habits often induces striking resemblances of structure between +the two animals, as in the case of whales and fish. But in all such +instances it is invariably found that the resemblance is only +superficial and apparent: not anatomical or real. In other words, the +resemblance does not extend further than <span class="pagenum"><a name="page_48" id="page_48">[48]</a></span> +it is necessary that it should, if both sets of organs are to be +adapted to perform the same functions. Now this, again, is just what one +would expect to find as the universal rule on the theory of descent, +with modification of ancestral characters. But, on the opposite theory +of special creation, I know not how it is to be explained that among so +many instances of close superficial resemblance between creatures +belonging to different branches of the tree of life, there are no +instances of any real or anatomical resemblance. So far as their +structures are adapted to perform a common function, there is in all +such cases what may be termed a deceptive appearance of some unity of +ideal; but, when carefully examined, it is always found that two +apparently identical structures occurring on different branches of the +classificatory tree are in fact fundamentally different in respect of +their structural plan.</p> + +<p>Lastly, we have seen that one of the guiding principles of +classification has been empirically found to consist in setting a high +value on “chains of affinities.” That is to say, naturalists +not unfrequently meet with a long series of progressive modifications of +type, which, although it cannot be said that the continuity is anywhere +broken, at last leads to so much divergence of character that, but for +the intermediate links, the members at each end of the chain could not +be suspected of being in any way related. Well, such cases of chains of +affinity obviously tell most strongly in favour of descent with +continuous modification; while it is impossible to suggest why, if all +the links were separately forged by as many acts of special creation, +there should have been this gradual transmutation of <span class="pagenum"><a name="page_49" id="page_49">[49]</a></span> +characters carried to the point where the original creative ideal has +been so completely transformed that, but for the accident of the chain +being still complete, no one of nature’s interpreters could +possibly have discovered the connexion. For, as we have seen, this is +not a case in which any appeal can be logically made to the argument +from ignorance of divine method, unless some independent evidence could +be adduced in favour of special creation. And that no such independent +evidence exists, it will be the object of future chapters to show.</p> + +<hr /><p class="pagenum"><a name="page_50" id="page_50">[50]</a></p> +<h2><a name="CHAPTER_III" id="CHAPTER_III"></a>CHAPTER III.<br /><br /> +<span class="sc">Morphology.</span></h2> + +<p>The theory of evolution supposes that hereditary characters admit of +being slowly modified wherever their modification will render an +organism better suited to a change in its conditions of life. Let us, +then, observe the evidence which we have of such adaptive modifications +of structure, in cases where the need of such modification is apparent. +We may begin by again taking the case of the whales and porpoises. The +theory of evolution infers, from the whole structure of these animals, +that their progenitors must have been terrestrial quadrupeds of some +kind, which gradually became more and more aquatic in their habits. Now +the change in the conditions of their life thus brought about would have +rendered desirable great modifications of structure. These changes would +have begun by affecting the least typical—that is, the least strongly +inherited—structures, such as the skin, claws, and teeth. But, as time +went on, the adaptation would have extended to more typical structures, +until the shape of the body would have become affected by the bones and +muscles required for terrestrial locomotion becoming better adapted for +aquatic locomotion, and <span class="pagenum"><a name="page_51" id="page_51">[51]</a></span> +the whole outline of the animal more fish-like in shape. This is the +stage which we actually observe in the seals, where the hind legs, +although retaining all their typical bones, have become shortened up +almost to rudiments, and directed backwards, so as to be of no use for +walking, while serving to complete the fish-like taper of the body. +(Fig. 2.) But in the whales the modification has gone further than this +so that the hind legs have ceased to be apparent externally, and are +only represented internally—and even this only in some +species—by remnants so rudimentary that it is difficult to make +out with certainty the homologies of the bones; moreover, the head and +the whole body have become completely fish-like in shape. (Fig. 3.) But +profound as are these alterations, they affect only those parts of the +organism which it was for the benefit of the organism to have altered, +so that it might be adapted to an aquatic mode of existence. Thus the +arm, which is used as a fin, still retains the bones of the shoulder, +fore-arm, wrist, and fingers, although they are all enclosed in a +fin-shaped sack, so as to render them useless for any purpose other than +swimming (Fig. 4.) Similarly, the head, although it so closely resembles +the head of a fish in shape, still retains the bones of the mammalian +skull in their proper anatomical relations to one another; but modified +in form so as to offer the least possible resistance to the water. In +short, it may be said that all the modifications have been effected with +the least possible divergence from the typical mammalian type, which is +compatible with securing so perfect an adaptation to a purely aquatic +mode of life.</p> + +<p class="pagenum"><a name="page_52" id="page_52">[52]</a></p> +<div class="figcenter" style="width: 600px;"> +<img src="images/i_069_002.jpg" width="600" height="360" alt="Skeleton of Seal." title="" /> +<div class="caption"><span class='sc'>Fig.</span> 2.—Skeleton of Seal, 1/8 nat. size. Drawn from +nature (R. Coll. Surg. Mus.).</div></div> + +<p class="pagenum"><a name="page_53" id="page_53">[53]</a></p> +<div class="figcenter" style="width: 600px;"> +<img src="images/i_070_003.jpg" width="600" height="277" alt="Skeleton of Greenland Whale" title="" /> +<div class="caption"><span class='sc'>Fig.</span> 3.—Skeleton of Greenland Whale, 1/100 nat. size. +The rudimentary bones of the pelvis are shown on a larger scale in the +upper drawing. (From Prof. Flower.)</div></div> + +<p><span class="pagenum"><a name="page_54" id="page_54">[54]</a></span> +Now I have chosen the case of the whale and porpoise group, because they +offer so extreme an example of profound modification of structure in +adaptation to changed conditions of life. But the same thing may be seen +in hundreds and hundreds of other cases. For instance, to confine our +attention to the arm, not only is the limb modified in the whale for +swimming, but in another mammal—the bat—it is modified for flying, by +having the fingers enormously elongated and overspread with a membranous +web.</p> + +<div class="figcenter" style="width: 485px;"> +<img src="images/i_071_004.jpg" width="485" height="389" alt="Paddle of Whale compared with Hand of Man." title="" /> +<div class="caption"><span class='sc'>Fig.</span> 4.—Paddle of Whale compared with Hand of Man. Drawn +from nature (R. Coll. Surg. Mus.).</div></div> + +<p>In birds, again, the arm is modified for flight in a wholly different +way—the fingers here being very <span class="pagenum"><a name="page_55" id="page_55">[55]</a></span> +Short and all run together, while the chief expanse of the wing is +composed of the shoulder and fore-arm. In frogs and lizards, again, we +find hands more like our own; but in an extinct species of flying +reptile the modification was extreme, the wing having been formed by a +prodigious elongation of the fifth finger, and a membrane spread over it +and the rest of the hand. (Fig. 5.) lastly, in serpents the hand and arm +have disappeared altogether.</p> + +<p class="pagenum"><a name="page_56" id="page_56">[56]</a></p> +<div class="figcenter" style="width: 318px;"> +<img src="images/i_073_005.jpg" width="318" height="500" alt="Wing of Reptile, Mammal, and Bird." title="" /> +<div class="caption"><span class='sc'>Fig.</span> 5.—Wing of Reptile, Mammal, and Bird. Drawn from +nature (Brit. Mus.).</div></div> + +<p><span class="pagenum"><a name="page_55c" id="page_55c">[55c.]</a></span> +Thus, even if we confine our attention to a single organ, how wonderful +are the modifications which it is seen to undergo, although never losing +its typical character. Everywhere we find the distinction between +homology and analogy which was explained in the last chapter—the +distinction, that is, between correspondence of structure and +correspondence of function. On the one hand, we meet with structures +which are perfectly homologous and yet in no way analogous: the +structural elements remain, but are profoundly modified so as to perform +wholly different functions. On the other hand, we meet with structures +which are perfectly analogous, and yet in no way homologous: totally +different structures are modified to perform the same functions. How, +then, are we to explain these things? By design manifested in special +creation, or by descent with adaptive modification? If it is said by +design manifested in special creation, we must suppose that the Deity +formed an archetypal plan of certain structures, and that he determined +to adhere to this plan through all the modifications which those +structures exhibit. But, if so, why is it that some structures are +selected<span class="pagenum"><a name="page_57" id="page_57">[57]</a></span> +as typical and not others? Why should the vertebral skeleton, for +instance, be tortured into every conceivable variety of modification in +order to subserve as great a variety of functions; while another +structure, such as the eye, is made in different sub-kingdoms on +fundamentally different plans, notwithstanding that it has throughout to +perform the same function? Will any one have the hardihood to assert +that in the case of the skeleton the Deity has endeavoured to show his +<i>ingenuity</i>, by the manifold functions to which he has made the same +structure subservient; while in the case of the eye he has endeavoured +to show his <i>resources</i>, by the manifold structures which he has adapted +to serve the same function? If so, it becomes a most unfortunate +circumstance that, throughout both the vegetable and animal kingdoms, +all cases which can be pointed to as showing ingenious adaptation of the +same typical structure to the performance of widely different +functions—or cases of homology without analogy,—are cases +which come within the limits of the same natural group of plants and +animals, and therefore admit of being equally well explained by descent +from a common ancestry; while all cases of widely different structures +performing the same function—or cases of analogy without +homology,—are to be found in different groups of plants or +animals, and are therefore suggestive of independent variations arising +in the different lines of hereditary descent.</p> + +<p>To take a specific illustration. The octopus, or devil-fish, belongs to +a widely different class of animals from a true fish; and yet its eye, +in general appearance, looks wonderfully like the eye of a true fish. +<span class="pagenum"><a name="page_58" id="page_58">[58]</a></span> +Now, Mr. Mivart pointed to this fact as a great difficulty in the way of +the theory of evolution by natural selection, because it must clearly be +a most improbable thing that so complicated a structure as the eye of a +fish should happen to be arrived at through each of two totally +different lines of descent. And this difficulty would, indeed, be a +formidable one to the theory of evolution, if the similarity were not +only analogical but homological. Unfortunately for the objection, +however, Darwin clearly showed in his reply that in no one anatomical or +homologous feature do the two structures resemble one another; so that, +in point of fact, the two organs do not resemble one another in any +particular further than it is necessary that they should, if both are to +be analogous, or to serve the same function as organs of sight. But now, +suppose that this had not been the case, and that the two structures, +besides presenting the necessary superficial or analogical resemblance, +had also presented an anatomical or homologous resemblance, with what +force might it have then been urged,—Your hypothesis of hereditary +descent with progressive modification being here excluded by the fact +that the animals compared belong to two widely different branches of the +tree of life, how are we to explain the identity of type manifested by +these two complicated organs of vision? <ins class='corr' title="Transcriber’s Note: The original showed ‘the’.">The</ins> +only hypothesis open to us is intelligent adherence to an ideal plan or +mechanism. But as this cannot now be urged in any comparable case +throughout the whole organic world, we may on the other hand present it +as a most significant fact, that while within the limits of the same +large branch of the tree of life we constantly find the same <span class="pagenum"><a name="page_59" id="page_59">[59]</a></span> +typical structures modified so as to perform very different functions, +we never find any of these particular types of structure in other large +branches of the tree. That is to say, we never find typical structures +appearing except in cases where their presence may be explained by the +hypothesis of hereditary descent; while in thousands of such cases we +find these structures undergoing every conceivable variety of adaptive +modification.</p> + +<p>Consequently, special creationists must fall back upon another position +and say,—Well, but it may have pleased the Deity to form a certain +number of ideal types, and never to have allowed the structures +occurring in one type to appear in any of the others. We +answer,—Undoubtedly such may have been the case; but, if so, it is a +most unfortunate thing for your theory, because the fact implies that +the Deity has planned his types in such a way as to suggest the +counter-theory of descent. For instance, it would seem most capricious +on the part of the Deity to have made the eyes of an innumerable number +of fish on exactly the same ideal type, and then to have made the eye of +the octopus so exactly like these other eyes in superficial appearance +as to deceive so accomplished a naturalist as Mr. Mivart, and yet to +have taken scrupulous care that in no one ideal particular should the +one type resemble the other. However, adopting for the sake of argument +this great assumption, let us suppose that God did lay down these +arbitrary rules for his own guidance in creation, and then let us see to +what the assumption leads. If the Deity formed a certain number of ideal +types, and determined that on no account should he allow any part of one +type <span class="pagenum"><a name="page_60" id="page_60">[60]</a></span> +to appear in any part of another, surely we should expect that within +the limits of the same type the same typical structures should always be +present. Thus, remember what efforts, so to speak, have been made to +maintain the uniformity of type in the case of the fore-limb as +previously explained, and should we not expect that in other and similar +cases a similar method should have been followed? Yet we repeatedly find +that this is not the case. Even in the whale, as we have seen, the +hind-limbs are either altogether absent or dwindled almost to nothing; +and it is impossible to see in what respect the hind-limbs are of any +less ideal value than the fore-limbs—which are carefully preserved +in all vertebrated animals except the snakes, and the extinct <span class='sn'>Dinornis</span>, +where again we meet in this particular with a sudden and sublime +indifference to the maintenance of a typical structure. (Fig. 6.)<a name="FNanchor_4_4" id="FNanchor_4_4"></a><a href="#Footnote_4_4" class="fnanchor">[4]</a> +Now I say that if the theory of ideal types is true, we have in these +facts evidence of a most unreasonable inconsistency. But the theory of +descent with continued adaptive modification fully explains all the +known cases; for in every case the degree of divergence from the typical +structure which an organism presents corresponds, in a general way, with +the length of time during which the divergence has been going on. Thus +we scarcely ever meet with any great departure from the typical form +with respect to one of the organs, without some of the other organs +being so far modified as of themselves to indicate, on the supposition +<span class="pagenum"><a name="page_62" id="page_62">[62]</a></span> +of descent with modification, that the animal or plant must have been +subject to the modifying influences for an enormously long series of +generations. And this combined testimony of a number of organs in the +same organism is what the theory of descent would lead us to expect, +while the rival theory of design can offer no explanation of the fact, +that when one organ shows a conspicuous departure from the supposed +ideal type, some of the other organs in the same organism should tend to +keep it company by doing likewise.</p> + +<p class="pagenum"><a name="page_61" id="page_61">[61]</a></p> +<div class="figcenter" style="width: 372px;"> +<img src="images/i_078_006.jpg" width="372" height="500" alt="Skeleton of Dinornis gravis." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 6.—Skeleton of Dinornis gravis, 1/16 nat. size. +Drawn from nature (Brit. Mus.). As separate cuts on a larger scale are +shown, 1st, the sternum, as this appears in mounted skeletons, and, 2nd, +the same in profile, with its (hypothetical) scapulo-coracoid attached.</div></div> + +<p><span class="pagenum"><a name="page_62c" id="page_62c">[62c.]</a></span> +As an illustration both of this and of other points which have been +mentioned, I may draw attention to what seems to me a particularly +suggestive case. So-called soldier-or hermit-crabs, are crabs which have +adopted the habit of appropriating the empty shells of mollusks. In +association with this peculiar habit, the structure of these animals +differs very greatly from that of all other crabs. In particular, the +hinder part of the body, which occupies the mollusk-shell, and which +therefore has ceased to require any hard covering of its own, has been +suffered to lose its calcareous integument, and presents a soft fleshy +character, quite unlike that of the more exposed parts of the animal. +Moreover, this soft fleshy part of the creature is specially adapted to +the particular requirements of the creature by having its lateral +appendages—i. e. appendages which in other crustacea perform the +function of legs—modified so as to act as claspers to the inside of the +mollusk-shell; while the tail-end of the part in question is twisted +into the form of a spiral, which fits into the spiral of the +mollusk-shell. Now, in Keeling Island there is a large kind of crab +called<span class="pagenum"><a name="page_63" id="page_63">[63]</a></span> +<span class='sn'>Birgus latro</span>, which lives upon land and there feeds upon cocoa-nuts. +The whole structure of this crab, it seems to me, unmistakeably +resembles the structure of a hermit-crab (see drawings on the next page, +Fig. 7). Yet this crab neither lives in the shell of a mollusk, nor is +the hinder part of its body in the soft and fleshy condition just +described: on the contrary, it is covered with a hard integument like +all the other parts of the animal. Consequently, I think we may infer +that the ancestors of <span class='sn'>Birgus</span> were hermit-crabs living in mollusk-shells; +but that their descendants gradually relinquished this habit as they +gradually became more and more terrestrial, while, concurrently with +these changes in habit, the originally soft posterior parts acquired a +hard protective covering to take the place of that which was formerly +supplied by the mollusk-shell. So that, if so, we now have, within the +limits of a single organism, evidence of a whole series of morphological +changes in the past history of its species. First, there must have been +the great change from an ordinary crab to a hermit-crab in all the +respects previously pointed out. Next, there must have been the change +back again from a hermit-crab to an ordinary crab, so far as living +without the necessity of a mollusk-shell is concerned. From an +evolutionary point of view, therefore, we appear to have in the existing +structure of <span class='sn'>Birgus</span> a morphological record of all these changes, and one +which gives us a reasonable explanation of why the animal presents the +extraordinary appearance which it does. But, on the theory of special +creation, it is inexplicable why this land-crab should have been formed +on the pattern of a hermit-crab, when it never has need to enter the +shell <span class="pagenum"><a name="page_65" id="page_65">[65]</a></span> +of a mollusk. In other words, its peculiar structure is not specially +in keeping with its present habits, although so curiously allied to the +similar structure of certain other crabs of totally different habits, in +relation to which the peculiarities are of plain and obvious +significance.</p> + +<p class="pagenum"><a name="page_64" id="page_64">[64]</a></p> +<div class="figcenter" style="width: 600px;"> +<img src="images/i_081_007.jpg" width="600" height="377" alt="Hermit-crabs compared with the cocoa-nut crab." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 7.—Hermit-crabs compared with the cocoa-nut crab. +On the left of the illustration one hermit-crab is represented as +occupying a mollusk-shell, and another (larger specimen) as it appears +when withdrawn from such a shell. On the right of the illustration the +cocoa-nut crab is represented in its natural habitat on land. When +full-grown, however, it is much larger than our hermit-crabs. The latter +are drawn from life, natural size, the former from a specimen in the +British Museum, 1/6 natural size.</div></div> + +<hr class='minor' /><span class="pagenum"><a name="page_65c" id="page_65c">[65c.]</a></span> + +<p>I will devote the remainder of this chapter to considering another +branch of the argument from morphology, to which the case of <span class='sn'>Birgus</span> +serves as a suitable introduction: I mean the argument from rudimentary +structures.</p> + +<p>Throughout both the animal and vegetable kingdoms we constantly meet +with dwarfed and useless representatives of organs, which in other and +allied kinds of animals and plants are of large size and functional +utility. Thus, for instance, the unborn whale has rudimentary teeth, +which are never destined to cut the gums; and throughout its life this +animal retains, in a similarly rudimentary condition, a number of organs +which never could have been of use to any kind of creature save a +terrestrial quadruped. The whole anatomy of its internal ear, for +example, has reference to hearing in air—or, as Hunter long ago +remarked, “is constructed upon the same principle as in the +quadruped"; yet, as Owen says, “the outer opening and passage +leading therefrom to the tympanum can rarely be affected by sonorous +vibrations of the atmosphere, and indeed they are reduced, or have +degenerated, to a degree which makes it difficult to conceive how such +vibrations can be propagated to the ear-drum during the brief moments in +which the opening may be raised above the water.”</p> + +<p class="pagenum"><a name="page_67" id="page_67">[67]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_084_008.jpg" width="500" height="341" alt="Rudimentary or vestigial hind-limbs of Python." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 8.—Rudimentary or vestigial hind-limbs of Python, +as exhibited in the skeleton and on the external surface of the animal. +Drawn from nature, ¼ nat. size (Zoological Gardens).</div></div> + +<p><span class="pagenum"><a name="page_66" id="page_66">[66]</a></span> +Now, rudimentary organs of this kind are of such frequent occurrence, +that almost every species presents one or more of them—usually, indeed, +a considerable number. How, then, are they to be accounted for? Of +course the theory of descent with adaptive modification has a simple +answer to supply—namely, that when, from changed conditions of life, an +organ which was previously useful becomes useless, it will be suffered +to dwindle away in successive generations, under the influence of +certain natural causes which we shall have to consider in future +chapters. On the other hand, the theory of special creation can only +maintain that these rudiments are formed for the sake of adhering to an +ideal type. Now, here again the former theory appears to be triumphant +over the latter; for, without waiting to dispute the wisdom of making +dwarfed and useless structures merely for the whimsical motive assigned, +surely if such a method were adopted in so many cases, we should expect +that in consistency it would be adopted in all cases. This reasonable +expectation, however, is far from being realized. We have already seen +that in numberless cases, such as that of the fore-limbs of serpents, no +vestige of a rudiment is present. But the vacillating policy in the +matter of rudiments does not end here; for it is shown in a still more +aggravated form where within the limits of the same natural group of +organisms a rudiment is sometimes present and sometimes absent. For +instance, although in nearly all the numerous species of snakes there +are no vestiges of limbs, in the Python we find very tiny rudiments of +the hind-limbs. (Fig. 8.) Now, is it a worthy conception of Deity that, +while neglecting to maintain his unity of <span class="pagenum"><a name="page_67c" id="page_67c">[67c.]</a></span> +ideal in the case of nearly all the numerous species of snakes, he +should have added a tiny rudiment in the case of the Python—and +even in that case should have maintained his ideal very inefficiently, +inasmuch as only two limbs, instead of four, are represented? How much +more reasonable is the naturalistic interpretation; for here the very +irregularity of their appearance in different species, which constitutes +rudimentary structures one of the crowning difficulties to the theory of +special design, furnishes the best possible evidence in favour of +hereditary descent; seeing that this irregularity then becomes what may +be termed the anticipated expression of progressive <span class="pagenum"><a name="page_68" id="page_68">[68]</a></span> +dwindling due to inutility. Thus, for example, to return to the case of +wings, we have already seen that in an extinct genus of bird, <span class='sn'>Dinornis</span>, +these organs were reduced to such an extent as to leave it still +doubtful whether so much as the tiny rudiment hypothetically supplied to +Fig. 6 (p. <a href="#page_61">61</a>) was present in all the species. And here is another +well-known case of another genus of still existing bird, which, as was +the case with <span class='sn'>Dinornis</span>, occurs only in New Zealand. (Fig. 9.) Upon this +island there are no four-footed enemies—either existing or +extinct—to escape from which the wings of birds would be of any +service. Consequently we can understand why on this island we should +meet with such a remarkable dwindling away of wings.</p> + +<p class="pagenum"><a name="page_69" id="page_69">[69]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_086_009.jpg" width="500" height="403" alt="Apteryx Australis." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 9.—Apteryx Australis. Drawn from life in the +Zoological Gardens, 1/8 nat. size. The external wing is drawn to a scale +in the upper part of the cut. The surroundings are supplied from the +most recent descriptions.</div></div> + +<p><span class="pagenum"><a name="page_68c" id="page_68c">[68c.]</a></span> +Similarly, the logger-headed duck of South America can only flap along +the surface of the water, having its wings considerably reduced though +less so than the <span class='sn'>Apteryx</span> of New Zealand. But here the interesting fact +is that the young birds are able to fly perfectly well. Now, in +accordance with a general law to be considered in a future chapter, the +life-history of an individual organism is a kind of condensed +recapitulation of the life-history of its species. Consequently, we can +understand why the little chickens of the logger-headed duck are able to +fly like all other ducks, while their parents are only able to flap +along the surface of the water.</p> + +<p>Facts analogous to this reduction of wings in birds which have no +further use for them, are to be met with also in insects under similar +circumstances. Thus, there are on the island of Madeira somewhere +between 500 and 600 species of beetles, which are in <span class="pagenum"><a name="page_69c" id="page_69c">[69c.]</a></span> +large part peculiar to that island, though related to other—and +therefore presumably parent—species on the neighbouring continent. +Now, no less than 200 species—or nearly half the whole +number—are so far deficient in wings that they cannot fly. And, if +we disregard the species which are not peculiar to the island—that +is to say, all the species which likewise occur on the neighbouring +continent, and therefore, as evolutionists conclude, have but <i>recently</i> +migrated to the island,—we find this very remarkable proportion. +There are altogether 29 peculiar genera, and out of <span class="pagenum"><a name="page_70" id="page_70">[70]</a></span> +these no less than 23 have <i>all</i> their species in this condition.</p> + +<p>Similar facts have been recently observed by the Rev. A. E. Eaton with +respect to insects inhabiting Kerguelen Island. All the species which he +found on the island—viz. a moth, several flies, and numerous +beetles—he found to be incapable of flight; and therefore, as Wallace +observes, “as these insects could hardly have reached the islands +in a wingless state, even if there were any other known land inhabited +by them, which there is not, we must assume that, like the Madeiran +insects, they were originally winged, and lost their power of flight +because its possession was injurious to them"—Kerguelen Island being +“one of the stormiest places on the globe,” and therefore a +place where insects could rarely afford to fly without incurring the +danger of being blown out to sea.</p> + +<p>Here is another and perhaps an even more suggestive class of facts.</p> + +<p>It is now many years ago since the editors of <i>Silliman’s Journal</i> +requested the late Professor Agassiz to give them his opinion on the +following question. In a certain dark subterranean cave, called the +Mammoth cave, there are found some peculiar species of blind fishes. Now +the editors of <i>Silliman’s Journal</i> wished to know whether Prof. +Agassiz would hold that these fish had been specially created in these +caves, and purposely devoided of eyes which could never be of any use to +them; or whether he would allow that these fish had probably descended +from other species, but, having got into the dark cave, gradually lost +their eyes through disuse. Prof. Agassiz, who was a believer in special +creation, <span class="pagenum"><a name="page_71" id="page_71">[71]</a></span> +allowed that this ought to constitute a crucial test as between the two +theories of special design and hereditary descent. “If physical +circumstances,” he said, “ever modified organized beings, it +should be easily ascertained here.” And eventually he gave it as +his opinion, that these fish “were created under the circumstances +in which they now live, within the limits over which they now range, and +with the structural peculiarities which now characterise them.”</p> + +<p>Since then a great deal of attention has been paid to the fauna of this +Mammoth cave, and also to the faunas of other dark caverns, not only in +the New, but also in the Old World. In the result, the following general +facts have been fully established.</p> + +<p>(1) Not only fish, but many representatives of other classes, have been +found in dark caves.</p> + +<p>(2) Wherever the caves are totally dark, all the animals are blind.</p> + +<p>(3) If the animals live near enough to the entrance to receive some +degree of light, they may have large and lustrous eyes.</p> + +<p>(4) In all cases the species of blind animals are closely allied to +species inhabiting the district where the caves occur; so that the blind +species inhabiting American caves are closely allied to American +species, while those inhabiting European caves are closely allied to +European species.</p> + +<p>(5) In nearly all cases structural remnants of eyes admit of being +detected, in various degrees of obsolescence. In the case of some of the +crustaceans of the Mammoth cave the foot-stalks of the eyes are present, +although the eyes themselves are entirely absent.</p> + +<p>Now, it is evident that all these general facts are in <span class="pagenum"><a name="page_72" id="page_72">[72]</a></span> +full agreement with the theory of evolution, while they offer serious +difficulties to the theory of special creation. As Darwin remarks, it is +hard to imagine conditions of life more similar than those furnished by +deep limestone caverns under nearly the same climate in the two +continents of America and Europe; so that, in accordance with the theory +of special creation, very close similarity in the organizations of the +two sets of faunas might have been expected. But, instead of this, the +affinities of these two sets of faunas are with those of their +respective continents—as of course they ought to be on the theory +of evolution. Again, what would have been the sense of creating useless +foot-stalks for the imaginary support of absent eyes, not to mention all +the other various grades of degeneration in other cases? So that, upon +the whole, if we agree with the late Prof. Agassiz in regarding these +cave animals as furnishing a crucial test between the rival theories of +creation and evolution, we must further conclude that the whole body of +evidence which they now furnish is weighing on the side of evolution.</p> + +<p>So much, then, for a few special instances of what Darwin called +rudimentary structures, but what may be more descriptively +designated—in accordance with the theory of descent—obsolescent or +vestigial structures. It is, however, of great importance to add that +these structures are of such general occurrence throughout both the +vegetable and animal kingdoms, that, as Darwin has observed, it is +almost impossible to point to a single species which does not present +one or more of them. In other words, it is almost impossible to find a +single species which does not in this <span class="pagenum"><a name="page_73" id="page_73">[73]</a></span> +way bear some record of its own descent from other species; and the +more closely the structure of any species is examined anatomically, the +more numerous are such records found to be. Thus, for example, of all +organisms that of man has been most minutely investigated by anatomists; +and therefore I think it will be instructive to conclude this chapter by +giving a list of the more noteworthy vestigial structures which are +known to occur in the human body. I will take only those which are found +in adult man, reserving for the next chapter those which occur in a +transitory manner during earlier periods of his life. But, even as thus +restricted, the number of obsolescent structures which we all present in +our own persons is so remarkable, that their combined testimony to our +descent from a quadrumanous ancestry appears to me in itself conclusive. +I mean, that even if these structures stood alone, or apart from any +more general evidences of our family relationships, they would be +sufficient to prove our parentage. Nevertheless, it is desirable to +remark that of course these special evidences which I am about to detail +do not stand alone. Not only is there the general analogy furnished by +the general proof of evolution elsewhere, but there is likewise the more +special correspondence between the whole of our anatomy and that of our +nearest zoological allies. Now the force of this latter consideration is +so enormous, that no one who has not studied human anatomy can be in a +position to appreciate it. For without special study it is impossible to +form any adequate idea of the intricacy of structure which is presented +by the human form. Yet it is found that this enormously intricate +organization is repeated in all its <span class="pagenum"><a name="page_74" id="page_74">[74]</a></span> +details in the bodies of the higher apes. There is no bone, muscle, +nerve, or vessel of any importance in the one which is not answered to +by the other. Hence there are hundreds of thousands of instances of the +most detailed correspondence, without there being any instances to the +contrary, if we pay due regard to vestigial characters. The entire +corporeal structure of man is an exact anatomical copy of that which we +find in the ape.</p> + +<p>My object, then, here is to limit attention to those features of our +corporeal structure which, having become useless on account of our +change in attitude and habits, are in process of becoming obsolete, and +therefore occur as mere vestigial records of a former state of things. +For example, throughout the vertebrated series, from fish to mammals, +there occurs in the inner corner of the eye a semi-transparent eye-lid, +which is called the nictitating membrane. The object of this structure +is to sweep rapidly, every now and then, over the external surface of +the eye, apparently in order to keep the surface clean. But although the +membrane occurs in all classes of the sub-kingdom, it is more prevalent +in some than in others—e.g. in birds than in mammals. Even, however, +where it does not occur of a size and mobility to be of any use, it is +usually represented, in animals above fishes, by a functionless +rudiment, as here depicted in the case of man. (Fig. 10.)</p> + +<p class="pagenum"><a name="page_75" id="page_75">[75]</a></p> +<div class="figcenter" style="width: 282px;"> +<img src="images/i_092_010.jpg" width="282" height="500" alt="Nictitating membrane in various animals." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 10.—Illustrations of the nictitating membrane in +the various animals named drawn from nature. The letter N indicates the +membrane in each case. In man it is called the plica semilunaris, and is +represented in the two lower drawings under this name. In the case of +the shark (Galeus) the muscular mechanism is shown as dissected.</div></div> + +<p><span class="pagenum"><a name="page_74c" id="page_74c">[74c.]</a></span> +Now the organization of man presents so many vestigial structures thus +referring to various stages of his long ancestral history, that it would +be tedious so much as to enumerate them. Therefore I will yet further +limit the list of vestigial structures to be given <span class="pagenum"><a name="page_76" id="page_76">[76]</a></span> +as examples, by not only restricting these to cases which occur in our +own organization; but of them I shall mention only such as refer us to +the very last stage of our ancestral history—viz. structures which +have become obsolescent since the time when our distinctively human +branch of the family tree diverged from that of our immediate +forefathers, the Quadrumana.</p> + +<div class="figcenter" style="width: 363px;"> +<img src="images/i_093_011.jpg" width="363" height="500" alt="Rudimentary muscles +of the human ear." title="" /> +<div class="caption"><span class='sc'>Fig.</span> 11.—Rudimentary, or vestigial and useless, muscles +of the human ear. (From Gray’s Anatomy.)</div></div> + +<p>(1) <i>Muscles of the external ear.</i>—These, which are of large size and +functional use in quadrupeds, we <span class="pagenum"><a name="page_77" id="page_77">[77]</a></span> +Retain in a dwindled and useless condition (Fig. 11). this is likewise +the case in anthropoid apes; but in not a few other quadrumana (e.g. +baboons, macacus, magots, &c.) degeneration has not proceeded so +far, and the ears are voluntarily moveable.</p> + +<p>(2) <span class='sn'>Panniculus carnosis.</span>—A large number of the mammalia are able to +move their skin by means of sub-cutaneous muscle—as we see, for +instance, in a horse, when thus protecting himself against the sucking +of flies. We, in common with the Quadrumana, possess an active remnant +of such a muscle in the skin of the forehead, whereby we draw up the +eyebrows; but we are no longer able to use other considerable remnants +of it, in the scalp and elsewhere,—or, more correctly, it is rarely +that we meet with persons who can. But most of the Quadrumana (including +the anthropoids) are still able to do so. There are also many other +vestigial muscles, which occur only in a small percentage of human +beings, but which, when they do occur, present unmistakeable homologies +with normal muscles in some of the Quadrumana and still lower +animals<a name="FNanchor_5_5" id="FNanchor_5_5"></a><a href="#Footnote_5_5" class="fnanchor">[5]</a>.</p> + +<p>(3) <i>Feet.</i>—It is observable that in the infant the feet have a strong +deflection inwards, so that the soles in considerable measure face one +another. This peculiarity, which is even more marked in the embryo than +in the infant (see p. <a href="#page_153">153</a>), and which becomes gradually less and less +conspicuous even before the child begins to walk, appears to me a highly +suggestive peculiarity. For it plainly refers to the condition <span class="pagenum"><a name="page_78" id="page_78">[78]</a></span> +of things in the Quadrumana, seeing that in all these animals the feet +are similarly curved inwards, to facilitate the grasping of branches. +And even when walking on the ground apes and monkeys employ to a great +extent the outside edges of their feet, as does also a child when +learning to walk. The feet of a young child are also extraordinarily +mobile in all directions, as are those of apes. In order to show these +points, I here introduce comparative drawings of a young ape and the +portrait of a young male child. These drawings, moreover, serve at the +<span class="pagenum"><a name="page_79" id="page_79">[79]</a></span> +same time to illustrate two other vestigial characters, which have +often been previously noticed with regard to the infant’s foot. I +allude to the incurved form of the legs, and the lateral extension of +the great toe, whereby it approaches the thumb-like character of this +organ in the Quadrumana. As in the case of the incurved position of the +legs and feet, so in this case of the lateral extensibility of the great +toe, the peculiarity is even more marked in embryonic than in infant +life. For, as Prof. Wyman has remarked with regard to the fœtus +when about an inch in length,<span class="pagenum"><a name="page_80" id="page_80">[80]</a></span> +“The great toe is shorter than the others; and, instead of being +parallel to them, is projected at an angle from the side of the foot, +thus corresponding with the permanent condition of this part in the +Quadrumana<a name="FNanchor_6_6" id="FNanchor_6_6"></a><a href="#Footnote_6_6" class="fnanchor">[6]</a>.” +So that this organ, which, according to Owen, “is perhaps the most +characteristic peculiarity in the human structure,” when traced +back to the early stages of its development, is found to present a +notably less degree of peculiarity.</p> + +<p class="pagenum"><a name="page_78i" id="page_78i">[78i.]</a></p> +<div class="figcenter" style="width: 400px;"> +<img src="images/i_095_012.jpg" width="400" height="477" alt="Young male gorilla." title="" /> +<div class="caption"><span class='sc'>Fig.</span> 12.—Portrait of a young male gorilla (after +Hartmann).</div></div> + +<p class="pagenum"><a name="page_79i" id="page_79i">[79i.]</a></p> +<div class="figcenter" style="width: 400px;"> +<img src="images/i_096_013.jpg" width="400" height="378" alt="Young male child." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 13.—Portrait of a young male child. Photographed +from life, when the mobile feet were for a short time at rest in a +position of extreme inflection.</div></div> + +<p><span class="pagenum"><a name="page_80c" id="page_80c">[80c.]</a></span> +(4) <i>Hands.</i>—Dr. Louis Robinson has recently observed that the grasping +power of the whole human hand is so surprisingly great at birth, and +during the first few weeks of infancy, as to be far in excess of present +requirements on the part of a young child. Hence he concludes that it +refers us to our quadrumanous ancestry—the young of anthropoid apes +being endowed with similar powers of grasping, in order to hold on to +the hair of the mother when she is using her arms for the purposes of +locomotion. This inference appears to me justifiable, inasmuch as no +other explanation can be given of the comparatively inordinate muscular +force of an infant’s grip. For experiments showed that very young +babies are able to support their own weight, by holding on to a +horizontal bar, for a period varying from one half to more than two +minutes<a name="FNanchor_7_7" id="FNanchor_7_7"></a><a href="#Footnote_7_7" class="fnanchor">[7]</a>. +With his kind permission I here reproduce one of Dr. Robinson’s +instantaneous, and hitherto unpublished, photographs of a very young +infant. This photograph was taken after the above paragraph (3) was +written, and I introduce it here because it serves to show +incidentally—and perhaps <span class="pagenum"><a name="page_81" id="page_81">[81]</a></span> +even better than the preceding figure—the points there mentioned +with regard to the feet and great toes. Again, as Dr. Robinson observes, +the attitude, and the disproportionately large development of the arms +as compared with the legs, give all the photographs a striking +resemblance to a picture of the chimpanzee “Sally” at the +Zoological Gardens. For “invariably the thighs are bent nearly at +right angles to the body, and in no case did the lower limbs hang down +and take the attitude of the erect position.” He adds, “In +many cases no sign of distress is evinced, <span class="pagenum"><a name="page_82" id="page_82">[82]</a></span> +and no cry uttered, until the grasp begins to give way.”</p> + +<p class="pagenum"><a name="page_81i" id="page_81i">[81i.]</a></p> +<div class="figcenter" style="width: 332px;"> +<img src="images/i_098_014.jpg" width="332" height="400" alt="3 weeks old infant supporting its own weight." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 14.—An infant, three weeks old, supporting its own +weight for over two minutes. The attitude of the lower limbs, feet, and +toes, is strikingly simian. Reproduced from an instantaneous photograph, +kindly given for the purpose by Dr. L. Robinson.</div></div> + +<p class="pagenum"><a name="page_82c" id="page_82c">[82c.]</a></p> + +<div class="figcenter" style="width: 450px;"> +<img src="images/i_099_015.jpg" width="450" height="413" alt="Sacrum of Gorilla and Man." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 15.—Sacrum of Gorilla compared with that of Man, +showing the rudimentary tail-bones of each. Drawn from nature (R. Coll. +Surg. Mus.).</div></div> + +<p>(5) <i>Tail.</i>—The absence of a tail in man is popularly supposed to +constitute a difficulty against the doctrine of his quadrumanous +descent. As a matter of fact, however, the absence of an external tail +in man is precisely what this doctrine would expect, seeing that the +nearest allies of man in the quadrumanous series are likewise destitute +of an external tail. Far, then, from this deficiency in man constituting +any difficulty to be accounted for, if the case were not so—i. e. if +man <i>did</i> possess an external tail,—the difficulty would be +<span class="pagenum"><a name="page_83" id="page_83">[83]</a></span> +to understand how he had managed to retain an organ which had been +renounced by his most recent ancestors. Nevertheless, as the anthropoid +apes continue to present the rudimentary vestiges of a tail in a few +caudal vertebræ below the integuments, we might well expect to +find a similar state of matters in the case of man. And this is just +what we do find, as a glance at these two comparative illustrations will +show. (Fig. 15.) Moreover, during embryonic life, both of the anthropoid +apes and of man, the tail much more closely resembles that of the lower +kinds of quadrumanous animals from which these higher representatives of +the group have descended. For at a certain stage of embryonic life the +tail, both of apes and of human beings, is actually longer than the legs +(see Fig. 16). And at this stage of development, also, the tail admits +of being moved by muscles which later on dwindle away. Occasionally, +however, <span class="pagenum"><a name="page_84" id="page_84">[84]</a></span> +these muscles persist, and are then described by anatomists as +abnormalities. The following illustrations serve to show the muscles in +question, when thus found in adult man.</p> + +<p class="pagenum"><a name="page_83i" id="page_83i">[83i.]</a></p> +<div class="figcenter" style="width: 350px;"> +<img src="images/i_100_016.jpg" width="350" height="204" alt="Human embryo, +about seven weeks old." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 16.—Diagrammatic outline of the human embryo when +about seven weeks old, showing the relations of the limbs and tail to +the trunk (after Allen Thomson), <i>r</i>, the radial, and <i>u</i>, the ulnar, +border of the hand and fore-arm; <i>t</i>, the tibial, and <i>f</i>, the fibular, +border of the foot and lower leg; <i>au</i>, ear; <i>s</i>, spinal cord; <i>v</i>, +umbilical cord; <i>b</i>, branchial gill-slits; <i>c</i>, tail.</div></div> + +<p class="pagenum"><a name="page_84c" id="page_84c">[84c.]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_101_017.jpg" width="500" height="379" alt="Front and back of adult human sacrum." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 17.—Front and back view of adult human sacrum, +showing abnormal persistence of vestigial tail-muscles. (The first +drawing is copied from Prof. Watson’s paper in Journl. Anat. and +Physiol., vol. 79: the second is compiled from different specimens.)</div></div> + +<p>(6) <i>Vermiform Appendix of the Cæcum.</i>—This is of large size and +functional use in the process of digestion among many herbivorous +animals; while in man it is not only too small to serve any such +purpose, but is even a source of danger to life—many persons dying +every year from inflammation set up by the lodgement in this blind tube +of fruit-stones, &c.</p> + +<p>In the orang it is longer than in man (Fig. 18), as <span class="pagenum"><a name="page_86" id="page_86">[86]</a></span> +it is also in the human fœtus proportionally compared with the +adult. (Fig. 19.) In some of the lower herbivorous animals it is longer +than the entire body.</p> + +<p class="pagenum"><a name="page_85" id="page_85">[85]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_102_018.jpg" width="500" height="193" alt="Appendix vermiformis in Orang and in Man." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 18.—Appendix vermiformis in Orang and in Man. Drawn +from dried inflated specimens in the Cambridge Museum by Mr. J. J. +Lister. <i>Il</i>, ilium; <i>Co</i>, colon; <i>C</i>, cæcum; W, a window cut in the +wall of the cæcum; X X X, the appendix.</div></div> + +<div class="figcenter" style="width: 500px;"> +<img src="images/i_102_019.jpg" width="500" height="398" alt="The same, showing variation in the Orang." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 19.—The same, showing variation in the Orang. Drawn +from a specimen in the Museum of the Royal College of Surgeons.</div></div> + +<p><span class="pagenum"><a name="page_86c" id="page_86c">[86c.]</a></span> +Like vestigial structures in general, however, this one is highly +variable. Thus the above cut (Fig. 19) serves to show that it may +sometimes be almost as short in the orang as it normally is in man—both +the human subjects of this illustration having been normal.</p> + +<p>(7) <i>Ear.</i>—Mr. Darwin writes:—</p> + +<div class='blockquot'><p>The celebrated sculptor, Mr. Woolner, informs me of one little +peculiarity in the external ear, which he has often observed both +<span class="figleft" style="width: 184px;"> +<img src="images/i_103_020.jpg" width="184" height="250" alt="Human ear." title="" /> +<span class="caption"><span class='sc'>Fig.</span> 20.—Human ear, modelled and drawn by Mr. Woolner. +<i>a</i>, the projecting point.</span></span> +in men and women.... The peculiarity consists in a little blunt +point, projecting from the inwardly folded margin, or helix. When +present, it is developed at birth, and, according to Prof. Ludwig +Meyer, more frequently in man than in woman. Mr. Woolner made an +exact model of one such case, and sent me the accompanying +drawing.... The helix obviously consists of the extreme margin of +the ear folded inwards; and the folding appears to be in some +manner connected with the whole external ear being permanently +pressed backwards. In many monkeys, which do not stand high in the +order, as baboons and some species of macacus, the upper portion of +the ear is slightly pointed, and the margin is not at all folded +inwards; but if the margin were to be thus folded, a slight point +would necessarily project towards the centre.... The following +wood-cut is an accurate copy of a photograph of the fœtus of an +orang (kindly sent me by Dr. Nitsche), in which it may be seen how +different the pointed outline of the ear is at this period from its +adult condition, when it bears a close <span class="pagenum"><a name="page_87" id="page_87">[87]</a></span> +general resemblance to that of man [including even the occasional +appearance of the projecting point shown in the preceding woodcut]. It +is evident that the folding over of the tip of such an ear, unless it +changed greatly during its further development, would give rise to a +point projecting inwards<a name="FNanchor_8_8" id="FNanchor_8_8"></a><a href="#Footnote_8_8" class="fnanchor">[8]</a>.</p></div> + +<div class="figcenter" style="width: 300px;"> +<img src="images/i_104_021.jpg" width="300" height="292" alt="Fœtus of an Orang." title="" /> +<div class="caption"><span class='sc'>Fig.</span> 21.—Fœtus of an Orang. Exact copy of a +photograph, showing the form of the ear at this early stage.</div></div> + +<p>The following woodcut serves still further to show vestigial +resemblances between the human ear and that of apes. The last two +figures illustrate the general resemblance between the normal ear of +fœtal man and the ear of an adult orang-outang. The other two figures +on the lower line are intended to exhibit occasional modifications of +the adult human ear, which approximate simian characters somewhat more +closely than does the normal type. It will be observed that in their +comparatively small lobes these ears resemble those of all the apes; and +that while the outer margin of one is not unlike that of the Barbary +<span class="pagenum"><a name="page_89" id="page_89">[89]</a></span> +ape, the outer margin of the other follows those of the chimpanzee and +orang. Of course it would be easy to select individual human ears which +present either of these characters in a more pronounced degree; but +these ears have been chosen as models because they present both +characters in conjunction. The upper row of figures likewise shows the +close similarity of hair-tracts, and the direction of growth on the part +of the hair itself, in cases where the human ear happens to be of an +abnormally hirsute character. But this particular instance (which I do +not think has been previously noticed) introduces us to the subject of +hair, and hair-growth, in general.</p> + +<p class="pagenum"><a name="page_88" id="page_88">[88]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_105_022.jpg" width="500" height="305" alt="Vestigial characters of human ears." title="" /> +<div class="caption"><span class='sc'>Fig.</span> 22.—Vestigial characters of human ears. Drawn from +nature.</div></div> + +<p><span class="pagenum"><a name="page_89c" id="page_89c">[89c.]</a></span> +(8) <i>Hair.</i>—Adult man presents rudimentary hair over most parts of the +body. Wallace has sought to draw a refined distinction between this +vestigial coating and the useful coating of quadrumanous animals, in the +absence of the former from the human back. But even this refined +distinction does not hold. On the one hand, the comparatively hairless +chimpanzee which died last year in the Zoological Gardens (<span class='sn'>T. calvus</span>) +was remarkably denuded over the back; and, on the other hand, men who +present a considerable development of hair over the rest of their bodies +present it also on their backs and shoulders. Again, in all men the +rudimentary hair on the upper and lower arm is directed towards the +elbow—a peculiarity which occurs nowhere else in the animal kingdom, +with the exception of the anthropoid apes and a few American monkeys, +where it presumably has to do with arboreal habits. For, when sitting in +trees, the orang, as observed by Mr. Wallace, places its hands above its +head with its elbows pointing downwards:<span class="pagenum"><a name="page_91" id="page_91">[91]</a></span> +the disposition of hair on the arms and fore-arms then has the effect +of thatch in turning the rain. Again, I find that in all species of +apes, monkeys, and baboons which I have examined (and they have been +numerous), the hair on the backs of the hands and feet is continued as +far as the first row of phalanges; but becomes scanty, or disappears +altogether, on the second row; while it is invariably absent on the +terminal row. I also find that the same peculiarity occurs in man. We +all have rudimentary hair on the first row of phalanges, both of hands +and feet: when present at all, it is more scanty on the second row; and +in no case have I been able to find any on the terminal row. In all +cases these peculiarities are congenital, and the total absence or +partial presence of hair on the second phalanges is constant in +different species of Quadrumana. For instance, it is entirely absent in +all the chimpanzees, which I have examined, while scantily present in +all the orangs. As in man, it occurs in a patch midway between the +joints.</p> + +<p class="pagenum"><a name="page_90" id="page_90">[90]</a></p> +<div class="figcenter" style="width: 296px;"> +<img src="images/i_107_023.jpg" width="296" height="500" alt="Hair-tracts on the arms and hands of Man and Chimpanzee." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 23.—Hair-tracts on the arms and hands of Man, as +compared with those on the arms and hands of Chimpanzee. Drawn from +life.</div></div> + +<p><span class="pagenum"><a name="page_91c" id="page_91c">[91c.]</a></span> +Besides showing these two features with regard to the disposition of +hair on the human arm and hand, the above woodcut illustrates a third. +By looking closely at the arm of the very hairy man from whom the +drawing was taken, it could be seen that there was a strong tendency +towards a whorled arrangement of the hairs on the backs of the wrists. +This is likewise, as a general rule, a marked feature in the arrangement +of hair on the same places in the gorilla, orang, and chimpanzee. In the +specimen of the latter, however, from which the drawing was taken, this +characteristic was not well marked. The downward direction of the hair +on the backs of the hands <span class="pagenum"><a name="page_92" id="page_92">[92]</a></span> +is exactly the same in man as it is in all the anthropoid apes. Again, +with regard to hair, Darwin notices that occasionally there appears in +man a few hairs in the eyebrows much longer than the others; and that +they seem to be representative of similarly long and scattered hairs +which occur in the chimpanzee, macacus, and baboons.</p> + +<p>Lastly, it may be here more conveniently observed than in the next +chapter on Embryology, that at about the sixth month the human fœtus +is often thickly coated with somewhat long dark hair over the entire +body, except the soles of the feet and palms of the hands, which are +likewise bare in all quadrumanous animals. This covering, which is +called the lanugo, and sometimes extends even to the whole forehead, +ears, and face, is shed before birth. So that it appears to be useless +for any purpose other than that of emphatically declaring man a child of +the monkey.</p> + +<p>(9) <i>Teeth.</i>—Darwin writes:—</p> + +<div class='blockquot'><p>It appears as if the posterior molar or wisdom-teeth were tending +to become rudimentary in the more civilized races of man. These +teeth are rather smaller than the other molars, as is likewise the +case with the corresponding teeth in the chimpanzee and orang; and +they have only two separate fangs.... They are also much more +liable to vary, both in structure and in the period of their +development, than the other teeth. In the Melanian races, on the +other hand, the wisdom-teeth are usually furnished with three +separate fangs, and are usually sound [i. e. not specially liable +to decay]; they also differ from the other molars in size, less +than in the Caucasian races.</p></div> + +<p>Now, in addition to these there are other respects in which the +dwindling condition of wisdom-teeth is manifested—particularly with +regard to the pattern of <span class="pagenum"><a name="page_93" id="page_93">[93]</a></span> +their crowns. Indeed, in this respect it would seem that even in the +anthropoid apes there is the beginning of a tendency to degeneration of +the molar teeth from behind forwards. For if we compare the three molars +in the lower jaw of the gorilla, orang, and chimpanzee, we find that the +gorilla has five well-marked cusps on all three of them; but that in the +orang the cusps are not so pronounced, while in the chimpanzee there are +only four of them on the third molar. Now in man it is only the first of +these three teeth which normally presents five cusps, both the others +presenting only four. So that, comparing all <span class="pagenum"><a name="page_94" id="page_94">[94]</a></span> +these genera together, it appears that the number of cusps is being +reduced from behind forwards; the chimpanzee having lost one of them +from the third molar, while man has not only lost this, but also one +from the second molar,—and, it may be added, likewise partially +(or even totally) from the first molar, as a frequent variation among +civilized races. But, on the other hand, variations are often met with +in the opposite direction, where the second or the third molar of man +presents five cusps—in the one case following the chimpanzee, in +the other the gorilla. These latter variations, therefore, may fairly be +regarded as reversionary. For these facts I am indebted to the kindness +of Mr. C. S. Tomes.</p> + +<p class="pagenum"><a name="page_93i" id="page_93i">[93i.]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_110_024.jpg" width="500" height="419" alt="Molar teeth of lower jaw in Gorilla, Orang, and +Man." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 24.—Molar teeth of lower jaw in Gorilla, Orang, and +Man. Drawn from nature, nat. size (<i>R. Mus. Coll. Surg.</i>).</div></div> + +<p><span class="pagenum"><a name="page_94c" id="page_94c">[94c.]</a></span> +(10) <i>Perforations of the humerus.</i>—The peculiarities which we have to +notice under this heading are two in number. First, the supra condyloid +foramen is a normal feature in some of the lower Quadrumana (Fig. 25), +where it gives passage to the great nerve of the fore-arm, and often +also to the great artery. In man, however, it is not a normal feature. +Yet it occurs in a small percentage of cases—viz., according to Sir W. +Turner, in about one per cent., and therefore is regarded by Darwin as a +vestigial character. Secondly, there is inter-condyloid foramen, which +is also situated near the lower end of the humerus, but more in the +middle of the bone. This occurs, but not constantly, in apes, and also +in the human species. From the fact that it does so much more frequently +in the bones of ancient—and also of some savage—races of mankind (viz. +in 20 to 30 per cent. of cases), Darwin is disposed to regard it also as +a vestigial feature. On the other hand, Prof. Flower tells me that in +his opinion it is but an expression <span class="pagenum"><a name="page_95" id="page_95">[95]</a></span> +of impoverished nutrition during the growth of the bone.</p> + +<div class="figcenter" style="width: 500px;"> +<img src="images/i_112_025.jpg" width="500" height="379" alt="Perforation of the humerus +in three species of Quadrumana and in Man." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 25.—Perforation of the humerus (supra-condyloid +foramen) in three species of Quadrumana where it normally occurs, and in +Man, where it does not normally occur. Drawn from nature (<i>R. Coll. +Surg. Mus.</i>).</div></div> + +<p>(11) <i>Flattening of tibia.</i>—In some very ancient human skeletons, there +has also been found a lateral flattening of the tibia, which rarely +occurs in any existing human beings, but which appears to have been +usual among the earliest races of mankind hitherto discovered. According +to Broca, the measurements of these fossil human tibiæ resemble those of +apes. Moreover, the bone is bent and strongly convex forwards, while its +angles are so rounded as to present the nearly oval section seen in +apes. It is in association with these ape-like human tibiæ that +perforated humeri of man are found in greatest abundance.</p> + +<p><span class="pagenum"><a name="page_96" id="page_96">[96]</a></span> +On the other hand, however, there is reason to doubt whether this form +of tibia in man is really a survival from his quadrumanous ancestry. +For, as Boyd-Dawkins and Hartmann have pointed out, the degree of +flattening presented by some of these ancient human bones is <i>greater</i> +than that which occurs in any existing species of anthropoid ape. Of +course the possibility remains that the unknown species of ape from +which man descended may have had its tibia more flattened than is now +observable in any of the existing species. Nevertheless, as some doubt +attaches to this particular case, I do not press it—and, indeed, only +mention it at all in order that the doubt may be expressed.</p> + +<p>Similarly, I will conclude by remarking that several other instances of +the survival of vestigial structures in man have been alleged, which are +of a still more doubtful character. Of such, for example, are the +supposed absence of the genial tubercle in the case of a very ancient +jaw-bone of man, and the disposition of valves in human veins. From the +former it was argued that the possessor of this very ancient jaw-bone +was probably speechless, inasmuch as the tubercle in existing man gives +attachment to muscles of the tongue. From the latter it has been argued +that all the valves in the veins of the human body have reference, in +their disposition, to the incidence of blood-pressure when the attitude +of the body is horizontal, or quadrupedal. Now, the former case has +already broken down, and I find that the latter does not hold. But we +can well afford to lose such doubtful and spurious cases, in view of all +the foregoing unquestionable and genuine cases of vestigial structures +which are <span class="pagenum"><a name="page_97" id="page_97">[97]</a></span> +to be met with even within the limits of our own organization—and +even when these limits are still further limited by selecting only those +instances which refer to the very latest chapter of our long ancestral +history.</p> + +<hr /><p class="pagenum"><a name="page_98" id="page_98">[98]</a></p> +<h2><a name="CHAPTER_IV" id="CHAPTER_IV"></a>CHAPTER IV.<br /><br /> +<span class="sc">Embryology.</span></h2> + +<p>We will next consider what of late years has become the most important +of the lines of evidence, not only in favour of the general fact of +evolution, but also of its history: I mean the evidence which has been +yielded by the newest of the sciences, the science of Embryology. But +here, as in the analogous case of adult morphology, in order to do +justice to the mass of evidence which has now been accumulated, a whole +volume would be necessary. As in that previous case, therefore, I must +restrict myself to giving an outline sketch of the main facts.</p> + +<p>First I will display what in the language of Paley we may call +“the state of the argument.”</p> + +<p>It is an observable fact that there is often a close correspondence +between developmental changes as revealed by any chronological series of +fossils which may happen to have been preserved, and developmental +changes which may be observed during the life-history of now existing +individuals belonging to the same group of animals. For instance, the +successive development of prongs in the horns of deer-like animals, +which is so clearly shown in the geological history of this tribe, is +closely reproduced <span class="pagenum"><a name="page_99" id="page_99">[99]</a></span> +in the life-history of existing deer. Or, in other words, the antlers +of an existing deer furnish in their development a kind of +<i>résumé</i>, or recapitulation, of the successive phases +whereby the primitive horn was gradually superseded by horns presenting +a greater and greater number of prongs in successive species of extinct +deer (Fig. 26). Now it must be obvious that such a recapitulation in the +life-history of an existing animal of developmental changes successively +distinctive of sundry allied, though now extinct species, speaks +strongly in favour of evolution. For as it is of the essence of this +theory that new forms arise from older forms by way of <i>hereditary</i> +descent, we should antecedently expect, if the theory is true, that the +phases of development presented by the individual organism would follow, +in their main outlines, those phases of development through which their +long line of ancestors had passed. The only alternative view is that as +species of deer, for instance, were separately created, additional +prongs were successively added to their antlers; and yet that, in order +to be so added to successive species every individual deer belonging to +later species was required to repeat in his own lifetime the process of +successive additions which had previously taken place in a remote series +of extinct species. Now I do not deny that this view is a possible view; +but I do deny that it is a probable one. According to the evolutionary +interpretation of such facts, we can see a very good <i>reason</i> why the +life-history of the individual is thus a condensed +<i>résumé</i> of the life-history of its ancestral species. But +according to the opposite view no reason can be assigned why such should +be the case. In a previous chapter—the <span class="pagenum"><a name="page_101" id="page_101">[101]</a></span> +chapter on Classification—we have seen that if each species were +created separately, no reason can be assigned why they should all have +been turned out upon structural patterns so strongly suggestive of +hereditary descent with gradual modifications, or slow +divergence—the result being group subordinated to group, with the +most generalized (or least developed) forms at the bottom, and the +highest products of organization at the top. And now we see—or +shall immediately see—that this consideration admits of being +greatly fortified by a study of the developmental history of every +individual organism. If it would be an unaccountable fact that every +separately created species should have been created with close +structural resemblances to a certain limited number of other species, +less close resemblances to certain further species, and so backwards; +assuredly it would be a still more unaccountable fact that every +individual of every species should exhibit in its own person a history +of developmental change, every term of which corresponds with the +structural peculiarities of its now extinct predecessors—and this +in the exact historical order of their succession in geological time. +The more that we think about this antithesis between the naturalistic +and the non-naturalistic interpretations, the greater must we feel the +contrast in respect of rationality to become; and, therefore, I need not +spend time by saying anything further upon the antecedent standing of +the two theories in this respect. The evidence, then, which I am about +to adduce from the study of development in the life-histories of +individual organisms, will be regarded by me as so much unquestionable +evidence in favour of <span class="pagenum"><a name="page_102" id="page_102">[102]</a></span> +similar processes of development in the life-histories of their +respective species—in so far, I mean, as the two sets of changes +admit of being proved parallel.</p> + +<p class="pagenum"><a name="page_100" id="page_100">[100]</a></p> +<div class="figcenter" style="width: 307px;"> +<img src="images/i_117_026.jpg" width="307" height="500" alt="Antlers of Stag." title="" /> +<div class="caption"><span class='sc'>Fig.</span> 26.—Antlers of Stag, showing successive addition of +branches in successive years. Drawn from nature (<i>Brit. Mus.</i>).</div></div> + +<p><span class="pagenum"><a name="page_102c" id="page_102c">[102c.]</a></span> +In the only illustration hitherto adduced—viz. that of deers’ +horns—the series of changes from a one-pronged horn to a fully +developed arborescent antler, is a series which takes place during the +adult life of the animal; for it is only when the breeding age has been +attained that horns are required to appear. But seeing that every animal +passes through most of the phases of its development, not only before +the breeding age has been attained, but even before the time of its own +birth, clearly the largest field for the study of individual development +is furnished by embryology. For instance, there is a salamander which +differs from most other salamanders in being exclusively terrestrial in +its habits. Now, the young of this salamander before their birth are +found to be furnished with gills, which, however, they are never +destined to use. Yet these gills are so perfectly formed, that if the +young salamanders be removed from the body of their mother shortly +before birth, and be then immediately placed in water, the little +animals show themselves quite capable of aquatic respiration, and will +merrily swim about in a medium which would quickly drown their own +parent. Here, then, we have both morphological and physiological +evidence pointing to the possession of gills by the ancestors of the +land salamander.</p> + +<p>It would be easy to devote the whole of the present chapter to an +enumeration of special instances of the kinds thus chosen for purposes +of illustration; but as it is desirable to take a deeper, and therefore +<span class="pagenum"><a name="page_103" id="page_103">[103]</a></span> +a more general view of the whole subject, I will begin at the +foundation, and gradually work up from the earliest stages of +development to the latest. Before starting, however, I ask the reader to +bear in mind one consideration, which must reasonably prevent our +anticipating that in <i>every case</i> the life-history of an individual +organism should present a <i>full</i> recapitulation of the life-history of +its ancestral line of species. Supposing the theory of evolution to be +true, it must follow that in many cases it would have been more or less +disadvantageous to a developing type that it should have been obliged to +reproduce in its individual representatives all the phases of +development previously undergone by its ancestry—even within the +limits of the same family. We can easily understand, for example, that +the waste of material required for building up the useless gills of the +embryonic salamanders is a waste which, sooner or later, is likely to be +done away with; so that the fact of its occurring at all is in itself +enough to show that the change from aquatic to terrestrial habits on the +part of this species must have been one of comparatively recent +occurrence. Now, in as far as it is detrimental to a developing type +that it should pass through any particular ancestral phases of +development, we may be sure that natural selection—or whatever +other adjustive causes we may suppose to have been at work in the +adaptation of organisms to their surroundings—will constantly seek +to get rid of this necessity, with the result, when successful, of +dropping out the detrimental phases. Thus the foreshortening of +developmental history which takes place in the individual lifetime may +be expected often to take place, not only in the way of <span class="pagenum"><a name="page_104" id="page_104">[104]</a></span> +condensation, but also in the way of excision. Many pages of ancestral +history may be recapitulated in the paragraphs of embryonic development, +while others may not be so much as mentioned. And that this is the true +explanation of what embryologists term “direct” +development—or of a more or less sudden leap from one phase to +another, without any appearance of intermediate phases—is proved +by the fact that in some cases both direct and indirect development +occur within the same group of organisms, some genera or families having +dropped out the intermediate phases which other genera or families +retain.</p> + +<hr class='minor' /> + +<p>The argument from embryology must be taken to begin with the first +beginning of individual life in the ovum. And, in order to understand +the bearings of the argument in this its first stage, we must consider +the phenomena of reproduction in the simplest form which these phenomena +are known to present.</p> + +<p>The whole of the animal kingdom is divided into two great groups, which +are called the Protozoa and the Metazoa. Similarly, the whole of the +vegetable kingdom is divided into the Protophyta and the Metaphyta. The +characteristic feature of all the Protozoa and Protophyta is that the +organism consists of a single physiological cell, while the +characteristic of all the Metazoa and Metaphyta is that the organism +consists of a plurality of physiological cells, variously modified to +subserve different functions in the economy of the animal or plant, as +the case may be. For the sake of brevity, I shall hereafter deal only +with the case of animals (Protozoa and Metazoa); but it may throughout +be understood that everything <span class="pagenum"><a name="page_105" id="page_105">[105]</a></span> +which is said applies also to the case of plants (Protophyta and +Metaphyta).</p> + +<p>A Protozoön (like a Protophyton) is a solitary cell, or a +“unicellular organism,” while a Metazoön (like a Metaphyton) +is a society of cells, or a “multicellular organism.” Now, +it is only in the multicellular organisms that there is any observable +distinction of sex. In all the unicellular organisms the phenomena of +reproduction appear to be more or less identical with those of growth. +Nevertheless, as these phenomena are here in some cases suggestively +peculiar, I will consider them more in detail.</p> + +<p>A Protozoön is a single corpuscle of protoplasm which in different +species of Protozoa varies in size from more than one inch to less than +1/1000 of an inch in diameter. In some species there is an enveloping +cortical substance; in other species no such substance can be detected. +Again, in most species there is a nucleus, while in other species no +such differentiation of structure has hitherto been observed. +Nevertheless, from the fact that the nucleus occurs in the majority of +Protozoa, coupled with the fact that the demonstration of this body is +often a matter of extreme difficulty, not only in some of the Protozoa +where it has been but recently detected, but also in the case of certain +physiological cells elsewhere,—from these facts it is not unreasonable +to suppose that all the Protozoa possess a nucleus, whether or not it +admits of being rendered visible by histological methods thus far at our +disposal. If this is the case, we should be justified in saying, as I +have said, that a Protozoön is an isolated physiological cell, and, like +cells in general, multiplies by means of what Spencer and Häckel +<span class="pagenum"><a name="page_106" id="page_106">[106]</a></span> +have aptly called a process of discontinuous growth. That is to say, +when a cell reaches maturity, further growth takes place in the +direction of a severance of its substance—the separated portion +thus starting anew as a distinct physiological unit. But, +notwithstanding the complex changes which have been more recently +observed to take place in the nucleus of some Protozoa prior to their +division, the process of multiplication by division may still be +regarded as a process of growth, which differs from the previous growth +of the individual cell in being attended by a severance of continuity. +If we take a suspended drop of gum, and gradually add to its size by +allowing more and more gum to flow into it, a point will eventually be +reached at which the force of gravity will overcome that of cohesion, +and a portion of the drop will fall away from the remainder. Here we +have a rough physical simile, although of course no true analogy. In +virtue of a continuous assimilation of nutriment, the protoplasm of a +cell increases in mass, until it reaches the size at which the forces of +disruption overcome those of cohesion—or, in other words, the +point at which increase of size is no longer compatible with continuity +of substance. Nevertheless, it must not be supposed that the process is +thus merely a physical one. The phenomena which occur even in the +simplest—or so-called “direct"—cell-division, are of +themselves enough to prove that the process is vital, or physiological; +and this in a high degree of specialization. But so, likewise, are all +processes of growth in organic structures; and therefore the simile of +the drop of gum is not to be regarded as a true analogy: it serves only +to <span class="pagenum"><a name="page_107" id="page_107">[107]</a></span> +indicate the fact that when cell-growth proceeds beyond a certain point +cell-division ensues. The size to which cells may grow before they thus +divide is very variable in different kinds of cells; for while some may +normally attain a length of ten or twelve inches, others divide before +they measure 1/1000 of an inch. This, however, is a matter of detail, +and does not affect the general physiological principles on which we are +at present engaged.</p> + +<p>Now, as we have seen, a Protozoön is a single cell; for even although in +some of the higher forms of protozoal life a colony of cells may be +bound together in organic connexion, each of these cells is in itself an +“individual,” capable of self-nourishment, reproduction, +and, generally, of independent existence. Consequently, when the growth +of a Protozoön ends in a division of its substance, the two parts wander +away from each other as separate organisms. (Fig. 27.)</p> + +<div class="figcenter" style="width: 400px;"> +<img src="images/i_124_027.jpg" width="400" height="226" alt="Fission of a Protozoön." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 27.—Fission of a Protozoön. In the left-hand +drawing the process is represented as having advanced sufficiently far +to have caused a division and segregation both of the nucleus and the +vesicle. In the right-hand drawing the process is represented as +complete. <i>n</i>, N, severed nucleus; <i>vc</i>, severed vesicle; <i>ps</i>, +pseudopodia; <i>f</i>, ingested food.</div></div> + +<p><span class="pagenum"><a name="page_108" id="page_108">[108]</a></span> +The next point we have to observe is, that in all cases where a cell or +a Protozoön multiplies by way of fissiparous division, the process +begins in the nucleus. If the nucleus divides into two parts, the whole +cell will eventually divide into two parts, each of which retains a +portion of the original nucleus, as represented in the above figure. If +the nucleus divides into three, four, or even, as happens in the +development of some embryonic tissues, into as many as six parts, the +cell will subdivide into a corresponding number, each retaining a +portion of the nucleus. Therefore, in all cases of fissiparous division, +the seat or origin of the process is the nucleus.</p> + +<p>Thus far, then, the phenomena of multiplication are identical in all the +lowest or unicellular organisms, and in the constituent cells of all the +higher or multicellular. And this is the first point which I desire to +make apparent. For where the object is to prove a continuity between the +phenomena of growth and reproduction, it is of primary importance to +show—1st, that there is such a continuity in the case of all the +unicellular organisms, and, 2nd, that there are all the above points of +resemblance between the multiplication of cells in the unicellular and +in the multicellular organisms.</p> + +<p>It remains to consider the points of difference, and, if possible, to +show that these do not go to disprove the doctrine of continuity which +the points of resemblance so forcibly indicate.</p> + +<p>The first point of difference obviously is, that in the case of all the +multicellular organisms the two or more “daughter-cells,” +which are produced by division of the “mother-cell,” do not +wander away from one <span class="pagenum"><a name="page_109" id="page_109">[109]</a></span> +another; but, as a rule, they continue to be held in more or less close +apposition by means of other cells and binding membranes,—with the +result of giving rise to those various “tissues,” which in +turn go to constitute the material of “organs.” I cannot +suppose, however, that any advocate of discontinuity will care to take +his stand at this point. But, if any one were so foolish as to do so, it +would be easy to dislodge him by describing the state of matters in some +of the Protozoa where a number of unicellular “individuals” +are organically united so as to form a “colony.” These cases +serve to bridge this distinction between Protozoa and Metazoa, of which +therefore we may now take leave.</p> + +<p>In the second place, there is the no less obvious distinction that the +result of cell-division in the Metazoa is not merely to multiply cells +all of the same kind: on the contrary, the process here gives rise to as +many different kinds of cells as there are different kinds of tissue +composing the adult organism. But no one, I should think, is likely to +oppose the doctrine of continuity on the ground of this distinction. For +the distinction is clearly one which must necessarily arise, if the +doctrine of continuity between unicellular and multicellular organisms +be true. In other words, it is a distinction which the theory of +evolution itself must necessarily pre-suppose, and therefore it is no +objection to the theory that its pre-supposition is realized. Moreover, +as we shall see better presently, there is no difficulty in +understanding why this distinction should have arisen, so soon as it +became necessary (or desirable) that individual cells, when composing a +“colony,” should <span class="pagenum"><a name="page_110" id="page_110">[110]</a></span> +conform to the economic principle of the division of labour—a +principle, indeed, which is already foreshadowed in the constituent +parts of a single cell, since the nucleus has one set of functions and +its surrounding protoplasm another.</p> + +<p>But now, in the third place, we arrive at a more important distinction, +and one which lies at the root of the others still remaining to be +considered. I refer to sexual propagation. For it is a peculiarity of +the multicellular organisms that, although many of them may likewise +propagate themselves by other means (Fig. 28), they all propagate +themselves by means of sexual congress. Now, in its essence, sexual +congress consists in the fusion of two specialized cells (or, as now +seems almost certain, of the nuclei thereof), so that it is out of such +a combination that the new individual arises by means of successive +cell-divisions, which, beginning in the fertilized ovum, eventually +build up all the tissues and organs of the body.</p> + +<p class="pagenum"><a name="page_111" id="page_111">[111]</a></p> +<div class="figcenter" style="width: 302px;"> +<img src="images/i_128_028.jpg" width="302" height="450" alt="Hydra viridis." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 28.—<span class='sn'>Hydra viridis</span>, partly in section. M, mouth; O, +ovary, or bud containing female reproductive cells; T, testis, or bud +containing male reproductive cells. In addition to these buds containing +germinal elements alone, there is another which illustrates the process +of “gemmation"—i. e. the direct out-growth of a fully formed +offspring.</div></div> + +<p><span class="pagenum"><a name="page_110c" id="page_110c">[110c.]</a></span> +This process clearly indicates very high specialization on the part of +germ-cells. For we see by it that although these cells when young +resemble all other cells in being capable of self-multiplication by +binary division (thus reproducing cells exactly like themselves), when +older they lose this power; but, at the same time, they acquire an +entirely new and very remarkable power of giving rise to a vast +succession of many different kinds of cells, all of which are mutually +correlated as to their several functions, so as to constitute a +hierarchy of cells—or, to speak literally, a multicellular +<i>co-organization</i>. Here it is that we touch the really important +distinction between the Protozoa and the Metazoa; for although I have +<span class="pagenum"><a name="page_111c" id="page_111c">[111c.]</a></span> +said that some of the higher Protozoa foreshadow this state of matters +in forming cell-colonies, it must now be noted that the cells composing +such colonies are all of the same kind; and, therefore, that the +principle of producing different kinds of cells which, by mutual +co-adaptation of functions, shall be capable of constructing a +multicellular Metazoön,—this great principle of <i>co-organization</i> is +but dimly nascent in the cell-colonies <span class="pagenum"><a name="page_112" id="page_112">[112]</a></span> +of Protozoa. And its marvellous development in the Metazoa appears +ultimately to depend upon the highly specialized character of +germ-cells. Even in cases where multicellular organisms are capable of +reproducing their kind without the need of any preceding process of +fertilization (parthenogenesis), and even in the still more numerous +cases where complete organisms are budded forth from any part of their +parent organism (gemmation, Fig. 28), there is now very good reason to +conclude that these powers of a-sexual reproduction on the part of +multicellular organisms are all ultimately due to the specialized +character of their germ-cells. For in all these cases the tissues of the +parent, from which the budding takes place, were ultimately derived from +germ-cells—no matter how many generations of budded organisms may +have intervened. And that propagation by budding, &c., in +multicellular organisms is thus ultimately due to their propagation by +sexual methods, seems to be further shown by certain facts which will +have to be discussed at some length in my next volume. Here, therefore, +I will mention only one of them—and this because it furnishes what +appears to be another important distinction between the Protozoa and the +Metazoa.</p> + +<p>In nearly all cases where a Protozoön multiplies itself by fission, the +process begins by a simple division of the nucleus. But when a Metazoön +is developed from a germ-cell, although the process likewise begins by a +division of the nucleus, this division is not a simple or direct one; on +the contrary, it is inaugurated by a series of processes going on within +the nucleus, which are so enormously complex, and withal so <span class="pagenum"><a name="page_113" id="page_113">[113]</a></span> +beautifully ordered, that to my mind they constitute the most +wonderful—if not also the most suggestive—which have ever +been revealed by microscopical research. It is needless to say that I +refer to the phenomena of karyokinesis. A few pages further on they will +be described more fully. For our present purposes it is sufficient to +give merely a pictorial illustration of their successive phases; for a +glance at such a representation serves to reveal the only point to which +attention has now to be drawn—namely, the immense complexity of +the processes in question, and therefore the contrast which they furnish +to the simple (or “direct") division of the nucleus preparatory to +cell-division in the unicellular organisms. Here, then<span class="pagenum"><a name="page_114" id="page_114">[114]</a></span> +(Fig. 29), we see the complex processes of karyokinesis in the first +two stages of egg-cell division. But similar processes continue to +repeat themselves in subsequent stages; and this, there is now good +reason to believe, throughout <i>all</i> the stages of cell-division, whereby +the original egg-cell eventually constructs an entire organism. In other +words, all the cells composing all the tissues of a multicellular +organism, at all stages of its development, are probably originated by +these complex processes, which differ so much from the simple process of +direct division in the unicellular organisms<a name="FNanchor_9_9" id="FNanchor_9_9"></a><a href="#Footnote_9_9" class="fnanchor">[9]</a>. +In this important respect, therefore, it does at first sight appear that +we have a distinction between the Protozoa and the Metazoa of so +pronounced a character, as fairly to raise the question whether +cell-division is fundamentally identical in unicellular and in +multicellular organisms.</p> + +<p class="pagenum"><a name="page_113i" id="page_113i">[113i.]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_130_029.jpg" width="500" height="288" alt="Stages in the division of the ovum +of a worm." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 29.—Successive stages in the division of the ovum, +or egg-cell, of a worm. (After Strasburger.) <i>a</i> to <i>d</i> show the changes +taking place in the nucleus and surrounding cell-contents, which result +in the first segmentation of the ovum at <i>e</i>; <i>f</i> and <i>g</i> show a +repetition of these changes in each of the two resulting cells, leading +to the second segmentation stage at <i>h</i>.</div></div> + +<p><span class="pagenum"><a name="page_114c" id="page_114c">[114c.]</a></span> +Lastly, the only other distinction of a physiologically significant kind +between a single cell when it occurs as a Protozoön and when it does so +as the unfertilized ovum of a Metazoön is, that in the latter case the +nucleus discharges from its own substance two minute protoplasmic masses +("polar bodies"), which are then eliminated from the cell altogether. +This process, which will be more fully described later on, appears to be +of invariable occurrence in the case of all egg-cells, <span class="pagenum"><a name="page_115" id="page_115">[115]</a></span> +while nothing resembling it has ever been observed in any of the +Protozoa.</p> + +<p>We must now consider these several points of difference <i>seriatim</i>.</p> + +<p>First, with regard to sexual propagation, we have already seen that this +is by no means the only method of propagation among the multicellular +organisms; and it now remains to add that, on the other hand, there is, +to say the least, a suggestive foreshadowing of sexual propagation among +the unicellular organisms. For although simple binary fission is here +the more usual mode of multiplication, very frequently two (rarely three +or more) Protozoa of the same species come together, fuse into a single +mass, and thus become very literally “one flesh.” This +process of “conjugation” is usually (though by no means +invariably) followed by a period of quiescent “encystation"; after +which the contents of the cyst escape in the form of a number of minute +particles, or “spores,” and these severally develope into +the parent type. Obviously this process of conjugation, when it is thus +a preliminary to multiplication, appears to be in its essence the same +as fertilization. And if it be objected that encystation and +spore-formation in the Protozoa are not always preceded by conjugation, +the answer would be that neither is oviparous propagation in the Metazoa +invariably preceded by fertilization.</p> + +<p>Nevertheless, that there are great distinctions between true sexual +propagation and this foreshadowing of it in conjugation I do not deny. +The question, however, is whether they be so great as to justify any +argument against an historical continuity between them. What, then, are +these remaining <span class="pagenum"><a name="page_116" id="page_116">[116]</a></span> +distinctions? Briefly, as we have seen, they are the extrusion from +egg-cells of polar bodies, and the occurrence, both in egg-cells and +their products (tissue-cells), of the process of karyokinesis. But, as +regards the polar bodies, it is surely not difficult to suppose that, +whatever their significance may be, it is probably in some way or +another connected with the high specialization of the functions which an +egg-cell has to discharge. Nor is there any difficulty in further +supposing that, whatever purpose is served by getting rid of polar +bodies, the process whereby they are got rid of was originally one of +utilitarian development—i. e. a process which at its commencement +did not betoken any difference of kind, or breach of continuity, between +egg-cells and cells of simpler constitution.</p> + +<p>Lastly, with respect to karyokinesis, although it is true that the +microscope has in comparatively recent years displayed this apparently +important distinction between unicellular and multicellular organisms, +two considerations have here to be supplied. The first is, that in some +of the Protozoa processes very much resembling those of karyokinesis +have already been observed taking place in the nucleus preparatory to +its division. And although such processes do not present quite the same +appearances as are to be met with in egg-cells, neither do the +karyokinetic processes in tissue-cells, which in their sundry kinds +exhibit great variations in this respect. Moreover, even if such were +not the case, the bare fact that nuclear division is not invariably of +the simple or direct character in the case of all Protozoa, is +sufficient to show that the distinction now before us—like the one last +dealt with—is by no means <span class="pagenum"><a name="page_117" id="page_117">[117]</a></span> +absolute. As in the case of sexual propagation, so in that of +karyokinesis, processes which are common to all the Metazoa are not +wholly without their foreshadowings in the Protozoa. And seeing how +greatly exalted is the office of egg-cells—and even of +tissue-cells—as compared with that of their supposed ancestry in +protozoal cells, it seems to me scarcely to be wondered at if their +specializations of function should be associated with corresponding +peculiarities of structure—a general fact which would in no way +militate against the doctrine of evolution. Could we know the whole +truth, we should probably find that in order to endow the most primitive +of egg-cells with its powers of marshalling its products into a living +army of cell-battalions, such an egg-cell must have been passed through +a course of developmental specialization of so elaborate a kind, that +even the complex processes of karyokinesis are but a very inadequate +expression thereof.</p> + +<p>Probably I have now said enough to show that, remarkable and altogether +exceptional as the properties of germ-cells of the multicellular +organisms unquestionably show themselves to be, yet when these +properties are traced back to their simplest beginnings in the +unicellular organisms, they may fairly be regarded as fundamentally +identical with the properties of living cells in general. Thus viewed, +no line of real demarcation can be drawn between growth and +reproduction, even of the sexual kind. The one process is, so to speak, +physiologically continuous with the other; and hence, so far as the +pre-embryonic stage of life-history is concerned, the facts cannot +fairly be regarded as out of keeping with the theory of evolution.</p> + +<p><span class="pagenum"><a name="page_118" id="page_118">[118]</a></span> +I will now pass on to consider the embryogeny of the Metazoa, beginning +at its earliest stage in the fertilization of the ovum. And here it is +that the constructive argument in favour of evolution which is derived +from embryology may be said properly to commence. For it is surely in +itself a most suggestive fact that all the Metazoa begin their life in +the same way, or under the same form and conditions. <i>Omne vivum ex +ovo.</i> This is a formula which has now been found to apply throughout the +whole range of the multicellular organisms. And seeing, as we have just +seen, that the ovum is everywhere a single cell, the formula amounts to +saying that, physiologically speaking, every Metazoön begins its life as +a Protozoön, and every Metaphyton as a Protophyton<a name="FNanchor_10_10" id="FNanchor_10_10"></a><a href="#Footnote_10_10" class="fnanchor">[10]</a>.</p> + +<p>Now, if the theory of evolution is true, what should we expect to happen +when these germ-cells are fertilized, and so enter upon their severally +distinct processes of development? Assuredly we should expect to find +that the higher organisms pass through the same phases of development as +the lower organisms, up to the time when their higher characters begin +to become apparent. If in the life-history of species these higher +characters were gained by gradual improvement upon lower characters, and +if the development of the higher individual is now a general +recapitulation of that of its ancestral species, in studying this +recapitulation we should expect to find the higher organism successively +unfolding its higher characters from the lower ones through which its +ancestral species had previously passed. And this is just what we do +<span class="pagenum"><a name="page_119" id="page_119">[119]</a></span> +find. Take, for example, the case of the highest organism, Man. Like +that of all other organisms, unicellular or multicellular, his +development starts from the nucleus of a single cell. Again, like that +of all the Metazoa and Metaphyta, his development starts from the +specially elaborated nucleus of an egg-cell, or a nucleus which has been +formed by the fusion of a male with a female element<a name="FNanchor_11_11" id="FNanchor_11_11"></a><a href="#Footnote_11_11" class="fnanchor">[11]</a>. +When his animality becomes established, he exhibits the fundamental +anatomical qualities which characterize such lowly animals as polyps and +jelly-fish. And even when he is marked off as a Vertebrate, it cannot be +said whether he is to be a fish, a reptile, a bird, or a beast. Later on +it becomes evident that he is to be a Mammal; but not till later still +can it be said to which order of mammals he belongs.</p> + +<p>Here, however, we must guard against an error which is frequently met +with in popular expositions of this subject. It is not true that the +embryonic phases in the development of a higher form always resemble so +many adult stages of lower forms. This may or may not be the case; but +what always is the case <span class="pagenum"><a name="page_120" id="page_120">[120]</a></span> +is, that the embryonic phases of the higher form resemble the +corresponding phases of the lower forms. Thus, for example, it would be +wrong to suppose that at any stage of his development a man resembles a +jelly-fish. What he does resemble at an early stage of his development +is the essential or groundplan of the jelly-fish, which that animal +presents in <i>its</i> embryonic condition, or before it begins to assume its +more specialized characters fitting it for its own particular sphere of +life. The similarities, therefore, which it is the function of +comparative embryology to reveal are the similarities of type or +morphological plan: not similarities of specific detail. Specific +details may have been added to this, that, and the other species for +their own special requirements, after they had severally branched off +from the common ancestral stem; and so could not be expected to recur in +the life-history of an independent specific branch. The comparison +therefore must be a comparison of embryo with embryo; not of embryos +with adult forms.</p> + +<hr class='minor' /> + +<p>In order to give a general idea of the results thus far yielded by a +study of comparative embryology in the present connexion, I will devote +the rest of this chapter to giving an outline sketch of the most +important and best established of these results.</p> + +<p>Histologically the ovum, or egg-cell, is nearly identical in all +animals, whether vertebrate or invertebrate. Considered as a cell it is +of large size, but actually it is not more than 1/100, and may be less +than 1/200 of an inch in diameter. In man, as in most mammals, it is +about 1/120. It is a more or less spherical body, presenting a thin +transparent envelope, called <span class="pagenum"><a name="page_121" id="page_121">[121]</a></span> +the <span class='sn'>zona pellucida</span>, which contains—first, the protoplasmic +cell-substance or “yolk,” within which lies, second, the +nucleus or germinal vesicle, within which again lies, third, the +nucleolus or germinal spot. This description is true of the egg-cells of +all animals, if we add that in the case of the lowest animals—such +as sponges, &c.—there is no enveloping membrane: the egg-cell +is here a naked cell, and its constituent protoplasm, being thus +unconfined, is free to perform protoplasmic movements, which it does +after the manner, and with all the activity, of an amœba. But even +with respect to this matter of an enveloping membrane, there is no +essential difference between an ovum of the lowest and an ovum of the +highest animals. For in their early stages of development within the +ovary the ova of the highest animals are likewise in the condition of +naked cells, exhibiting amœbiform movements; the enveloping +membrane of an ovum being the product of a later development.<span class="pagenum"><a name="page_123" id="page_123">[123]</a></span> +Moreover this membrane, when present, is usually provided with one or +more minute apertures, through which the spermatozoön passes when +fertilizing the ovum. It is remarkable that the spermatozoa know, so to +speak, of the existence of these gate-ways,—their snake-like +movements being directed towards them, presumably by a stimulus due to +some emanation therefrom<a name="FNanchor_12_12" id="FNanchor_12_12"></a><a href="#Footnote_12_12" class="fnanchor">[12]</a>. +In the mammalian ovum, however, these apertures are exceedingly minute, +and distributed <span class="pagenum"><a name="page_124" id="page_124">[124]</a></span> +all round the circumference of the pellucid envelope, as represented in +this illustration (Fig. 32).</p> + +<p class="pagenum"><a name="page_121i" id="page_121i">[121i.]</a></p> +<div class="figcenter" style="width: 300px;"> +<img src="images/i_138_030.jpg" width="300" height="229" alt="Ovarian ovum of a Mammal." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 30.—Ovarian ovum of a Mammal, (<i>a</i>) magnified and +viewed under pressure, (<i>b</i>) burst by increased pressure, with yolk and +nucleus escaping: (<i>c</i>) the nucleus more freed from yolk-substance. +(From <i>Quain’s Anatomy</i>, after Allen Thomson.)</div></div> + +<p class="pagenum"><a name="page_122" id="page_122">[122]</a></p> +<div class="figcenter" style="width: 328px;"> +<img src="images/i_139_031.jpg" width="328" height="500" alt="Amœboid movements of young egg-cells." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 31.—Amœboid movements of young egg-cells, <i>a</i>, +Amœboid ovum of <span class='sn'>Hydra</span> (from Balfour, after Kleitnenberg); <i>b</i>, early +ovum of <span class='sn'>Toxopneustes variegatus</span>, with pseudopodia-like processes (from +Balfour, after Selenka); <i>c</i>, ovum of <span class='sn'>Toxopneustes lividus</span>, more nearly +ripe (from Balfour, Hertwig). A1 to A4, the primitive egg-cell of a +Chalk-Sponge (<span class='sn'>Leuculmis echinus</span>), in four successive conditions of +motion. B1 to B8, ditto of a Hermit-Crab (<span class='sn'>Chondracanthus cornutus</span>), in +eight successive stages (after E. von Beneden). C1 to C5, ditto of a +Cat, in five successive stages (after Pflüger). D, ditto of Trout; E, of +a Hen; F, of Man. The first series is taken from the <i>Encycl. Brit.</i>; +the second from Häckel’s <i>Evolution of Man</i>.</div></div> + +<p class="pagenum"><a name="page_123i" id="page_123i">[123i.]</a></p> +<div class="figcenter" style="width: 400px;"> +<img src="images/i_140_032.jpg" width="400" height="401" alt="Human ovum, mature and greatly magnified." title="" /> +<div class="caption"><span class='sc'>Fig.</span> 32.—Human ovum, mature and greatly magnified. +(After Häckel.)</div></div> + +<p><span class="pagenum"><a name="page_124c" id="page_124c">[124c.]</a></span> +In thus saying that the ova of all animals are, so far as microscopes +can reveal, <i>substantially</i> similar, I am of course speaking of the +egg-cell proper, and not of what is popularly known as the egg. The egg +of a bird, for example, is the egg-cell, <i>plus</i> an enormous aggregation +of nutritive material, an egg-shell, and sundry other structures suited +to the subsequent development of the egg-cell when separated from the +parent’s body. But all these accessories are, from our present +point of view, accidental or adventitious. What we have now to +understand by the ovum, the egg, or the egg-cell, is the microscopical +germ which I have just described. So far then as this germ is concerned, +we find that all multicellular organisms begin their existence in the +same kind of structure, and that this structure is anatomically +indistinguishable from that of the permanent form presented by the +lowest, or unicellular organisms. But although anatomically +indistinguishable, physiologically they present the sundry peculiarities +already mentioned.</p> + +<p>Now I have endeavoured to show that none of these peculiarities are such +as to exclude—or even so much as to invalidate—the supposition of +developmental continuity between the lowest egg-cells and the highest +protozoal cells. It remains to show in this place, and on the other +hand, that there is no breach of continuity between the lowest and the +highest egg-cells; but, on the contrary, that the remarkable uniformity +of the complex processes whereby their peculiar characters are exhibited +to the histologist, is such as of itself to sustain the doctrine of +continuity <span class="pagenum"><a name="page_125" id="page_125">[125]</a></span> +in a singularly forcible manner. On this account, therefore, and also +because the facts will again have to be considered in another connexion +when we come to deal with Weismann’s theory of heredity, I will +here briefly describe the processes in question.</p> + +<div class="figcenter" style="width: 500px;"> +<img src="images/i_142_033.jpg" width="500" height="293" alt="Polar bodies in +the ovum of a star-fish." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 33.—Stages in the formation of the polar bodies in +the ovum of a star-fish. (After Hertwig.) <i>g.v.</i>, germinal vesicle +transformed into a spindle-shaped system of fibres; <i>p.</i>′, the first +polar body becoming extruded; <i>p.</i>, <i>p.</i>, both polar bodies fully +extruded; <i>f. pn.</i>, female pronucleus, or residue of the germinal +vesicle.</div></div> + +<p>We have already seen that the young egg-cell multiplies itself by simple +binary division, after the manner of unicellular organisms in +general—thereby indicating, as also by its amœbiform movements, its +fundamental identity with such organisms in kind. But, as we have +likewise seen, when the ovum ceases to resemble these organisms, by +taking on its higher degree of functional capacity, it is no longer able +to multiply itself in this manner. On the contrary, its cell-divisions +are now of an endogenous character, <span class="pagenum"><a name="page_126" id="page_126">[126]</a></span> +and result in the formation of many different kinds of cells, in the +order required for constructing the multicellular organism to which the +whole series of processes eventually give rise. We have now to consider +these processes <i>seriatim</i>.</p> + +<div class="figcenter" style="width: 500px;"> +<img src="images/i_143_034.jpg" width="500" height="241" alt="Fertilization of the ovum of an echinoderm." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 34.—Fertilization of the ovum of an echinoderm. +(From <i>Quain’s Anatomy</i>, after Selenka.) S, spermatozoön; <i>m. +pr.</i>, male pronucleus; <i>f. pr.</i>, female pronucleus. 1 to 4 correspond to +D to G in the next figure.</div></div> + +<p>First of all the nucleus discharges its polar bodies, as previously +mentioned, and in the manner here depicted on the previous page. (Fig. +33.) It will be observed that the nucleus of the ovum, or the germinal +vesicle as it is called, gets rid first of one and afterwards of the +other polar body by an “indirect,” or karyokinetic, process +of division. (Fig. 33.) Extrusion of these bodies from the ovum (or it +may be only from the nucleus) having been accomplished, what remains of +the nucleus retires from the circumference of the ovum, and is called +the female pronucleus. (Fig. 33. <i>f. pn.</i>) The ovum is now ready for +fertilization. A similar emission of nuclear substance is said by some +good <span class="pagenum"><a name="page_127" id="page_127">[127]</a></span> +observers to take place also from the male germ-cell, or +spermatozoön, at or about the close of <i>its</i> development. The +theories to which these facts have given rise will be considered in +future chapters on Heredity.</p> + +<p>Turning now to the mechanism of fertilization, the diagrams (Figs. 34, +35) represent what happens in the case of star-fish.</p> + +<div class="figcenter" style="width: 450px;"> +<img src="images/i_144_035.jpg" width="450" height="147" alt="Fertilization of the ovum of a star-fish." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 35.—Fertilization of the ovum of a star-fish. (From +the <i>Encycl. Brit.</i> after Fol.) A, spermatozoa in the mucilaginous coat +of the ovum; a prominence is rising from the surface of the ovum towards +a spermatozoön; B, they have almost met; C, they have met; D, the +spermatozoön enters the ovum through a distinct opening; H, the entire +ovum, showing extruded polar bodies on its upper surface, and the moving +together of the male and female pronuclei; E, F, G, meeting and +coalescence of the pronuclei.</div></div> + +<p>The sperm-cell, or spermatozoön, is seen in the act of penetrating the +ovum. In the first figure it has already pierced the mucilaginous coat +of the ovum, the limit of which is represented by a line through which +the tail of the spermatozoön is passing: the head of the spermatozoön is +just entering the ovum proper. It may be noted that, in the case of many +animals, the general protoplasm of the ovum becomes aware, so to speak, +of the approach of a spermatozoön, and sends up a process to meet it. +(Fig. 35, A, B, C.) Several—or even many—spermatozoa may thus enter +the coat of the ovum; but normally only one proceeds further, or right +into the substance of the ovum, for the <span class="pagenum"><a name="page_128" id="page_128">[128]</a></span> +purpose of effecting fertilization. This spermatozoön, as soon as +it enters the periphery of the yolk, or cell-substance proper, sets up a +series of remarkable phenomena. First, its own head rapidly increases in +size, and takes on the appearance of a cell-nucleus: this is called the +male pronucleus. At the same time its tail begins to disappear, and the +enlarged head proceeds to make its way directly towards the nucleus of +the ovum which, as before stated, is now called the female pronucleus. +The latter in its turn moves towards the former, and when the two meet +they fuse into one mass, forming a new nucleus. Before the two actually +meet, the spermatozoön has lost its tail altogether; and it is +noteworthy that during its passage through the protoplasmic +cell-contents of the ovum, it appears to exercise upon this protoplasm +an attractive influence; for the granules of the latter in its vicinity +dispose themselves around it in radiating lines. All these various +phenomena are depicted in the above wood-cuts. (Figs. 34, 35.)</p> + +<p>Fertilization having been thus effected by fusion of the male and female +pronuclei into a single (or new) nucleus, this latter body proceeds to +exhibit complicated processes of karyokinesis, which, as before shown, +are preliminary to nuclear division in the case of egg-cells. Indeed the +karyokinetic process may begin in both the pronuclei before their +junction is effected; and, even when their junction is effected, it does +not appear that complete fusion of the so-called chromatin elements of +the two pronuclei takes place. For the purpose of explaining what this +means, and still more for the purpose of giving a general idea of the +karyokinetic processes as a whole,<span class="pagenum"><a name="page_129" id="page_129">[129]</a></span> +I will quote the following description of them, because, for terseness +combined with lucidity, it is unsurpassable.</p> + +<div class="figcenter" style="width: 450px;"> +<img src="images/i_146_036.jpg" width="450" height="293" alt="Karyokinesis of a typical tissue-cell +(epithelium of Salamander)." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 36.—Karyokinesis of a typical tissue-cell +(epithelium of Salamander). (After Flemming and Klein.) The series from +A to I represents the successive stages in the movement of the chromatin +fibres during division, excepting G, which represents the +“nucleus-spindle” of an egg-cell. A, resting nucleus; D, +wreath-form; E, single star, the loops of the wreath being broken; F, +separation of the star into two groups of U-shaped fibres; H, diaster or +double star; I, completion of the cell-division and formation of two +resting nuclei. In G the chromatin fibres are marked <i>a</i>, and correspond +to the “equatorial plate"; <i>b</i>, achromatin fibres forming the +nucleus-spindle; <i>c</i>, granules of the cell-protoplasm forming a +“polar star.” Such a polar star is seen at each end of the +nucleus-spindle, and is not to be confused with the diaster H, the two +ends of which are composed of <span class='sn'>chromatin</span>.</div></div> + +<div class='blockquot'><p>Researches, chiefly due to Flemming, have shown that the nucleus in +very many tissues of higher plants and animals consists of a +capsule containing a plasma of “achromatin,” not deeply +stained by re-agents, ramifying in which is a reticulum of +“chromatin” consisting of fibres which readily take a +deep stain. (Fig. 36, A). Further it is demonstrated that, when the +cell is about to divide into two, definite and very remarkable +movements take place in the nucleus, resulting in the disappearance +of the capsule and in the arrangement of its fibres first in the +<span class="pagenum"><a name="page_130" id="page_130">[130]</a></span> +form of a wreath (D), and subsequently (by the breaking of the loops +formed by the fibres) in the form of a star (E). A further movement +within the nucleus leads to an arrangement of the broken loops in two +groups (F), the position of the open ends of the broken loops being +reversed as compared with what previously obtained. Now the two groups +diverge, and in many cases a striated appearance of the achromatin +substance between the two groups of chromatin loops is observable (H). +In some cases (especially egg-cells) this striated arrangement of the +achromatin is then termed a “nucleus-spindle,” and the group +of chromatin loops (G, <i>a</i>) is known as “the equatorial +plate.” At each end of the nucleus-spindle in these cases there is +often seen a star consisting of granules belonging to the general +protoplasm of the cell (G, <i>c</i>). These are known as “polar +stars.” After the separation of the two sets of loops (H) the +protoplasm of the general substance of the cell becomes constricted, and +division occurs, so as to include a group of chromatin loops in each of +the two fission products. Each of these then rearranges itself together +with the associated chromatin into a nucleus such as was present in the +mother cell to commence with (I)<a name="FNanchor_13_13" id="FNanchor_13_13"></a><a href="#Footnote_13_13" class="fnanchor">[13]</a>.</p></div> + +<p>Since the above was published, however, further progress has been made. +In particular it has been found that the chromatin fibres pass from +phase D to phase F by a process of longitudinal splitting (Fig. 37 <i>g</i>, +<i>h</i>; Fig. 38, VI, VII)—which is a point of great importance for +Weismann’s theory of heredity,—and that the protoplasm outside +the nucleus seems to take as important a part in the karyokinetic +process as does the nuclear substance. For the so-called +“attraction-spheres” (Fig. 38 II <i>a</i>, III, III <i>a</i>, VIII to +XII), which were at first supposed to be of subordinate importance in +the process as a whole, are now known to take an exceedingly active part +in it (see especially IX to XI). Lastly, it may be added that there is a +<span class="pagenum"><a name="page_134" id="page_134">[134]</a></span> +growing consensus of authoritative opinion, that the chromatin +fibres are the seats of the material of heredity, or, in other words, +that they contain those essential elements of the cell which endow the +daughter-cells with their distinctive characters. Therefore, where the +parent-cell is an ovum, it follows from this view that all hereditary +qualities of the future organism are potentially present in the +ultra-microscopical structure of the chromatin fibres.</p> + +<p class="pagenum"><a name="page_131" id="page_131">[131]</a></p> +<div class="figcenter" style="width: 469px;"> +<img src="images/i_148_037.jpg" width="469" height="500" alt="Successive changes in the +nucleus of an epithelium cell, preparatory to division of the cell." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 37.—Study of successive changes taking place in the +nucleus of an epithelium cell, preparatory to division of the cell. +(From <i>Quain’s Anatomy</i>, after Flemming.) <i>a</i>, resting cell, +showing the nuclear network; <i>b</i>, first stage of division, the +chromatoplasm transformed into a skein of closely contorted filaments; +<i>c</i> to <i>f</i>, further stages in the growth and looping arrangement of the +filaments; <i>g</i>, stellate phase, or aster; <i>h</i>, completion of the +splitting of the filaments, already begun in <i>f</i> and <i>g</i>; <i>i</i>, <i>j</i>, <i>k</i>, +successive stages in separation of the filaments into two groups; <i>l</i>, +the final result of this (diaster); <i>m</i> to <i>q</i>, stages in the division +of the whole cell into two, showing increasing contortion of the +filaments, until they reach the resting stage at <i>q</i>.</div></div> + +<p class="pagenum"><a name="page_132" id="page_132">[132]</a></p> +<div class="figcenter" style="width: 438px;"> +<img src="images/i_149_038a.jpg" width="438" height="500" alt="Formation and conjugation of the pronuclei in +Ascaris megalocephala." title="" /> +<div class="captionh"><p class='hi'><span class='sc'>Fig.</span> 38.—Formation and conjugation of the pronuclei in +<span class='sn'>Ascaris megalocephala</span>. (From <i>Quain’s Anatomy</i>, after E. von +Beneden.) <i>f</i>, female pronucleus; <i>m</i>, male pronucleus; <i>p</i>, one of the +polar bodies.</p> + +<p class='hi'>I. The second polar body has just been extruded; both male and female +pronuclei contain two chromatin particles; those of the male pronucleus +are becoming transformed into a skein. II. The chromatin in both +pronuclei now forms into a skein.</p> + +<p class='hi'>II <i>a</i>. The skeins are more distinct. Two attraction (or protoplasmic) +spheres, each with a central particle united with a small spindle of +achromatic fibres, have made their appearance in the general substance +of the egg close to the mutually approaching pronuclei. The male +pronucleus has the remains of the body of the spermatozoön adhering to +it.</p> + +<p class='hi'>III. Only the female pronucleus is shown in this figure. The skein is +contracted and thickened. The attraction-spheres are near one side of +the ovum, and are connected with its periphery by a cone of fibres +forming a polar circle, <i>p.c.</i>; <i>e.c.</i>, equatorial circle.</p> + +<p class='hi'>III <i>a</i>. The pronuclei have come into contact, and the spindle-system is +now arranged across their common axis.</p> + +<p class='hi'>IV. Contraction of the skein, and formation of two U-or V-shaped +chromatin fibres in each pronucleus.</p> + +<p class='hi'>V. The V-shaped chromatin filaments are now quite distinct: the male and +female pronuclei are in close contact.</p></div></div> + +<p class="pagenum"><a name="page_133" id="page_133">[133]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_150_038b.jpg" width="500" height="411" alt="Previous figure continued." title="" /> +<div class="captionh"><p class='hi'>VI., VII. The V-shaped filaments are splitting longitudinally; their +structure of fine granules of chromatin is apparent in VII., which is +more highly magnified. The conjugation of the pronuclei is apparently +complete in VII. The attraction-spheres and achromatic spindle, although +present, are not depicted in IV., V., VI., and VII.</p> + +<p class='hi'>VIII. Equatorial arrangement of the four chromatin loops in the middle +of the now segmenting ovum: the achromatic substance forming a +spindle-shaped system of granules with fibres radiating from the poles +of the spindle (attraction-spheres); the chromatin forms an equatorial +plate. (Compare Fig. 36 G.)</p> + +<p class='hi'>IX. Shows diagrammatically the commencing separation of the chromatin +fibres of the conjugated nuclei, and the system of fibres radiating from +the attraction-spheres. (Compare again Fig. 36 G.) <i>p.c.</i>, polar circle; +<i>e.c.</i>, equatorial circle; <i>c.c.</i>, central particle.</p> + +<p class='hi'>X. Further separation of the chromatin filaments. Each of the central +particles of the attraction-spheres has divided into two.</p> + +<p class='hi'>XI. The chromatin fibres are becoming developed into the skeins of the +two daughter-nuclei. These are still united by fibres of achromatin. The +general protoplasm of the ovum is becoming divided.</p> + +<p class='hi'>XII. The two daughter-nuclei exhibit a chromatin network. Each of the +attraction-spheres has divided into two, which are joined by fibres of +achromatin, and connected with the periphery of the cell in the same way +as in the original or parent sphere, III.</p></div></div> + +<p><span class="pagenum"><a name="page_134c" id="page_134c">[134c.]</a></span> +As I shall have more to say about these processes in the next volume, +when we shall see the important part which they bear in Weismann’s +theory of heredity, it is with a double purpose that I here introduce +these yet further illustrations of them upon a somewhat larger scale. +The present purpose is merely that of showing, more clearly than +hitherto, the great complexity of these processes on the one hand, and, +on the other, the general similarity which they display in egg-cells and +in tissue-cells. But as in relation to this purpose the illustrations +speak for themselves, I may now pass on at once to the history of +embryonic development, which follows fertilization of the ovum.</p> + +<hr class='minor' /> + +<p>We have seen that when the new nucleus of the fertilized ovum (which is +formed by a coalescence of the male pronucleus with the female) has +completed its karyokinetic processes, it is divided into two equal +parts; that these are disposed at opposite poles of the ovum; and that +the whole contents of the ovum are thereupon likewise divided into two +equal parts, with the result that there are now two nucleated cells +within the spherical wall of the ovum where before there had only been +one. Moreover, we have also seen that a <span class="pagenum"><a name="page_135" id="page_135">[135]</a></span> +precisely similar series of events repeat themselves in each of these +two cells, thus giving rise to four cells (see Fig. 29). It must now be +added that such duplication is continued time after time, as shown in +the accompanying illustrations (Figs. 39, 40).</p> + +<div class="figcenter" style="width: 500px;"> +<img src="images/i_152_039.jpg" width="500" height="157" alt="Segmentation of ovum." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 39.—Segmentation of ovum. (After Häckel.) +Successive stages are marked by the letters A, B, C. D represents +several stages in advance of C.</div></div> + +<p>All<span class="figright" style="width: 150px;"> +<img src="images/i_152_040.jpg" width="150" height="157" alt="The contents of an ovum in an advanced stage of +segmentation." title="" /> +<span class="caption"><span class='sc'>Fig.</span> 40.—The contents of an ovum in an advanced stage of +segmentation, drawn in perspective. (After Häckel.)</span></span> +this, it will be noticed, is a case of cell-multiplication, which +differs from that which takes place in the unicellular organisms only in +its being <i>invariably</i> preceded (as far as we know) by karyokinesis, and +in the resulting cells being all confined within a common envelope, and +so in not being free to separate. Nevertheless, from what has already +been said, it will also be noticed that this feature makes all the +difference between a Metazoön and a Protozoön; so that already the ovum +presents the distinguishing character of a Metazoön.</p> + +<p><span class="pagenum"><a name="page_136" id="page_136">[136]</a></span> +I have dealt thus at considerable length upon the processes whereby the +originally unicellular ovum and spermatozoön become converted into the +multicellular germ, because I do not know of any other exposition of the +argument from Embryology where this, the first stage of the argument, +has been adequately treated. Yet it is evident that the fact of all the +processes above described being so similar in the case of sexual (or +metazoal) reproduction among the innumerable organisms where it occurs, +constitutes in itself a strong argument in favour of evolution. For the +mechanism of fertilization, and all the processes which even thus far we +have seen to follow therefrom, are hereby shown to be not only highly +complex, but likewise highly specialized. Therefore, the remarkable +similarity which they present throughout the whole animal kingdom—not +to speak of the vegetable—is expressive of organic continuity, rather +than of absolute discontinuity in every case, as the theory of special +creation must necessarily suppose. And it is evident that this argument +is strong in proportion to the uniformity, the specialization, and the +complexity of the processes in question.</p> + +<p>Having occupied so much space with supplying what appear to me the +deficiencies in previous expositions of the argument from Embryology, I +can now afford to take only a very general view of the more important +features of this argument as they are successively furnished by all the +later stages of individual development. But this is of little +consequence, seeing that from the point at which we have now arrived +previous expositions of the argument are both good and numerous. The +following then is to be regarded as a mere sketch <span class="pagenum"><a name="page_137" id="page_137">[137]</a></span> +Of the evidences of phyletic (or ancestral) evolution, which are so +abundantly furnished by all the subsequent phases of ontogenetic (or +individual) evolution.</p> + +<div class="figcenter" style="width: 500px;"> +<img src="images/i_154_041.jpg" width="500" height="313" alt="Formation of the gastrula of Amphioxus." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 41.—Formation of the gastrula of <span class='sn'>Amphioxus</span>. (After +Kowalevsky.) A, wall of the ovum, composed of a single layer of cells; +B, a stage in the process of gastrulation; C, completion of the process; +S, original or segmentation cavity of ovum; <i>al</i>, alimentary cavity of +gastrula; <i>ect</i>, outer layer of cells; <i>ent</i>, inner layer of cells; <i>b</i>, +orifice, constituting the mouth in permanent forms.</div></div> + +<p>The multicellular body which is formed by the series of segmentations +above described is at first a sphere of cells (Fig. 40). Soon, however, +a watery fluid gathers in the centre, and progressively pushes the cells +towards the circumference, until they there constitute a single layer. +The ovum, therefore, is now in the form of a hollow sphere containing +fluid, confined within a continuous wall of cells (Fig. 41 A). The next +thing that happens is a pitting in of one portion of the sphere (B). The +pit becomes deeper and deeper, until there is a complete invagination of +this part of the sphere—the cells which constitute it being +progressively <span class="pagenum"><a name="page_138" id="page_138">[138]</a></span> +pushed inwards until they come into contact with those at the opposite +pole of the ovum. Consequently, instead of a hollow sphere of cells, the +ovum now becomes an open sac, the walls of which are composed of a +double layer of cells (C). The ovum is now what has been called a +gastrula; and it is of importance to observe that probably all the +Metazoa pass through <span class="pagenum"><a name="page_139" id="page_139">[139]</a></span> +this stage. At any rate it has been found to occur in all the main +divisions of the animal kingdom, as a glance at the accompanying figures +will serve to show (Fig. 42)<a name="FNanchor_14_14" id="FNanchor_14_14"></a><a href="#Footnote_14_14" class="fnanchor">[14]</a>. +Moreover many of the lower kinds of Metazoa never pass beyond it; but +are all their lives nothing else than gastrulæ, wherein the +orifice becomes the mouth of the animal, the internal or invaginated +layer of cells the stomach, and the outer layer the skin. So that if we +take a child’s india-rubber ball, of the hollow <span class="pagenum"><a name="page_140" id="page_140">[140]</a></span> +kind with a hole in it, and push in one side with our fingers till +internal contact is established all round, by then holding the indented +side downwards we should get a very fair anatomical model of a +gastræa form, such as is presented by the adult condition of many +of the most primitive Metazoa—especially the lower <span class='sn'>Cœlenterata</span>. +The preceding figures represent two <span class="pagenum"><a name="page_141" id="page_141">[141]</a></span> +other such forms in nature, the first locomotive and transitory, the +second fixed and permanent (Figs. 43, 44).</p> + +<p class="pagenum"><a name="page_138i" id="page_138i">[138i.]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_155_042.jpg" width="500" height="416" alt="Gastrulation." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 42.—Gastrulation. A, Gastrula of a Zoophyte (<span +class='sn'>Gastrophysema</span>). (After Häckel.) B, Gastrula of a Worm (<span +class='sn'>Sagitta</span>). (After Kowalevsky.) C, Gastrula of an Echinoderm +(<span class='sn'>Uraster</span>). (After A. Agassiz.) D, Gastrula of an Arthropod (<span class='sn'>Nauplius</span>). +(After Häckel.) E, Gastrula of a Mollusk (<span class='sn'>Limnæus</span>). (After Rabl.) F, Gastrula of a Vertebrate (<span class='sn'>Amphioxus</span>). (After Kowalevsky.) In all, <i>d</i>, +indicates the intestinal cavity; <i>o</i>, the primitive mouth; <i>s</i>, the +cleavage-cavity; <i>i</i>, the endoderm, or intestinal layer; <i>e</i>, the +ectoderm or skin-layer.</div></div> + +<p class="pagenum"><a name="page_139i" id="page_139i">[139i.]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_156_043.jpg" width="500" height="289" alt="Gastrula of a Chalk Sponge." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 43.—Gastrula of a Chalk Sponge. (After Häckel.) A, +External view. B, Longitudinal section. <i>g</i>, digestive cavities; <i>o</i>, +mouth; <i>i</i>, endoderm; <i>e</i>, ectoderm.</div></div> + +<p class="pagenum"><a name="page_140i" id="page_140i">[140i.]</a></p> +<div class="figcenter" style="width: 411px;"> +<img src="images/i_157_044.jpg" width="411" height="500" alt="Prophysema primordiale, an extant gastræa-form." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 44.—<span class='sn'>Prophysema primordiale</span>, an extant gastræa-form. +(After Häckel.) (A). External view of the whole animal, attached by its +foot to seaweed. (B). Longitudinal section of the same. The digestive +cavity (<i>d</i>) opens at its upper end in the mouth (<i>m</i>). Among the cells +of the endoderm (<i>g</i>) lie amœboid egg-cells of large size (<i>e</i>). The +ectoderm (<i>h</i>) is encrusted with grains of sand, above the sponge +spicules.</div></div> + +<p><span class="pagenum"><a name="page_141c" id="page_141c">[141c.]</a></span> +Here, then, we leave the lower forms of Metazoa in their condition of +permanent gastrulæ. They differ from the transitory stage of other +Metazoa only in being enormously larger (owing to greatly further +<i>growth</i>, without any further <i>development</i> as to matters of fundamental +importance), and in having sundry tentacles and other organs added later +on to meet their special requirements. The point to remember is, that in +all cases a gastrula is an open sac composed of two layers of cells—the +outer layer being called the ectoderm, and the inner the endoderm. They +have also been called the animal layer and the vegetative layer, because +it is the outer layer (ectoderm) that gives rise to all the organs of +sensation and movement—viz. the skin, the nervous system, and the +muscular system; while it is the inner layer (endoderm) that gives rise +to all the organs of nutrition and reproduction. It is desirable only +further to explain that gastrulation does not take place in all the +Metazoa after exactly the same plan. In different lines of descent +various and often considerable modifications of the original and most +simple plan have been introduced; but I will not burden the present +exposition by describing these modifications<a name="FNanchor_15_15" id="FNanchor_15_15"></a><a href="#Footnote_15_15" class="fnanchor">[15]</a>. +It is enough for us that they always end in the formation of the two +primary layers of ectoderm and endoderm.</p> + +<p>The next stage of differentiation is common to all the Metazoa, except +those lowest forms which, as we <span class="pagenum"><a name="page_142" id="page_142">[142]</a></span> +Have just seen, remain permanently as large gastrulæ, with sundry +specialized additions in the way of tentacles, &c. This stage of +differentiation consists in the formation of either a pouch or an +additional layer between the ectoderm and the endoderm, which is called +the mesoderm. It is probably in most cases derived from the endoderm, +but the exact mode of its derivation is still somewhat obscure. +sometimes it has the appearance of itself constituting two layers; but +it is needless to go into these details; for in any case the ultimate +result is the same—viz. that of converting the metazoön into +the form of a tube, the walls of which are composed of concentric layers +of cells. The outermost layer afterwards gives rise to the epidermis +with its various appendages, and also to the central nervous system with +its organs of special sense. The median layer gives rise to the +voluntary muscles, bones, cartilages, &c., the nutritive systems of +the blood, the chyle, the lymph, and the muscular tube of the intestine. +lastly, the innermost layer developes into the epithelium lining of the +intestine, with its various appendages of liver, lungs, intestinal +glands, &c.</p> + +<p>I have just said that this three or four layered stage is shared by all +the Metazoa, except those very lowest forms—such as sponges and +jelly-fish—which do not pass on to it. But from this point the +developmental histories of all the main branches of the Metazoa +diverge—the Vermes, the Echinodermata, the Mollusca, the Articulata, +and the Vertebrata, each taking a different road in their subsequent +evolution. I will therefore confine attention to only one of these +several roads or methods, namely, that which is <span class="pagenum"><a name="page_143" id="page_143">[143]</a></span> +followed by the Vertebrata—observing merely that, if space +permitted, the same principles of progressive though diverging histories +of evolution would equally well admit of being traced in all the other +sub-kingdoms which have just been named.</p> + +<div class="figcenter" style="width: 500px;"> +<img src="images/i_160_045.jpg" width="500" height="170" alt="Ideal primitive vertebrate, seen from the left side." title="" /> +<div class="captionh"><span class='sc'>Fig.</span> 45.—Ideal primitive vertebrate, seen from the left +side. (After Häckel.) <i>na</i>, nose; <i>au</i>, eye; <i>g</i>, ear; <i>md</i>, mouth; <i>ks</i>, +gill-openings; <i>x</i>, notochord; <i>mr</i>, spinal tube; <i>kg</i>, gill-vessels; +<i>k</i>, gill-intestine; <i>hz</i>, heart; <i>ms</i>, muscles; <i>ma</i>, stomach; <i>v</i>, +intestinal vein; <i>c</i>, body-cavity; <i>a</i>, aorta; <i>l</i>, liver; <i>d</i>, small +intestine; <i>e</i>, ovary; <i>h</i>, testes; <i>n</i>, kidney canal; <i>af</i>, anus; <i>lh</i>, +true or leather-skin; <i>oh</i>, outer-skin (epidermis); <i>f</i>, skin-fold, +acting as a fin.</div></div> + +<p>In order to trace these principles in the case of the Vertebrata, it is +desirable first of all to obtain an idea of the anatomical features +which most essentially distinguish the sub-kingdom as a whole. +<span class="figleft" style="width: 169px;"> +<img src="images/i_161_046.jpg" width="169" height="179" alt="Fig. 46.—The same in transverse section through the +ovaries; lettering as in the preceding Fig." title="" /> +<span class="caption"><span class='sc'>Fig.</span> 46.—The same in transverse section through the +ovaries; lettering as in the preceding Fig.</span></span> +The following, then, is what may be termed the ideal plan of vertebrate +organization, as given by Prof. Häckel. First, occupying the major +axis of body we perceive the primitive vertebral column. The parts lying +above this axis are those which have been developed from the ectoderm +and mesoderm—viz. voluntary muscles, central nervous system, and +organs of special sense. The parts lying below this axis are for the +most part those which have been developed from the +endoderm—namely, <span class="pagenum"><a name="page_144" id="page_144">[144]</a></span> +the digestive tract with its glandular appendages, the circulating +system and the respiratory system. In transverse section, therefore, the +ideal vertebrate consists of a solid axis, with a small tube occupied by +the nervous system above, and a large tube, or body-cavity, below. This +body-cavity contains the viscera, breathing organs, and heart, with its +prolongations into the main blood-vessels of the organism. Lastly, on +either side of the central axis are to be found large masses of +muscle—two on the dorsal and two on the ventral. As yet, however, +there are no limbs, nor even any bony skeleton, for the primitive +vertebral column is hitherto unossified cartilage. This ideal animal, +therefore, is to all appearance as much like a worm as a fish, and swims +by means of a lateral undulation of its whole body, assisted, perhaps, +by a dorsal fin formed out of skin.</p> + +<p class="pagenum"><a name="page_145" id="page_145">[145]</a></p> +<div class="figcenter" style="width: 600px;"> +<img src="images/i_162_047.jpg" width="600" height="136" alt="Amphioxus lanceolatus." title="" /> +<div class="captionh"><span class="sc">Fig</span>. 47.—<span class='sn'>Amphioxus lanceolatus</span>. (After Häckel.) <i>a</i>, +anus; <i>au</i>, eye; <i>b</i>, ventral muscles; <i>c</i>, body-cavity; <i>ch</i>, +notochord; <i>d</i>, intestine; <i>do</i> and <i>du</i>, dorsal and ventral walls of +intestine; <i>f</i>, fin-seam; <i>h</i>, skin; <i>k</i>, gills; <i>ka</i>, gill-artery; +<i>lb</i>, liver; <i>lv</i>, liver-vein; <i>m 1</i>, brain-bladder; <i>m 2</i>, spinal +marrow; <i>mg</i>, stomach; <i>o</i>, mouth; <i>p</i>, ventral pore; <i>r</i>, dorsal +muscle; <i>s</i>, tail-fin; <i>t</i>, aorta; <i>v</i>, intestinal vein; <i>x</i>, boundary +between gill-intestine and stomach-intestine; <i>y</i>, hypobranchial +groove.</div></div> + +<p><span class="pagenum"><a name="page_144c" id="page_144c">[144c.]</a></span> +Now I should not have presented this ideal representation of a primitive +vertebrate—for I have very little faith in the “scientific use of +the imagination” where it aspires to discharge the functions of a +Creator in the manufacture of archetypal forms—I say I should not have +presented this ideal representative of a primitive vertebrate, were it +not that the ideal is actually realized in a still existing animal. For +there still survives what must be an immensely archaic form of +vertebrate, whose anatomy is almost identical with that of the imaginary +type which has just been <span class="pagenum"><a name="page_146" id="page_146">[146]</a></span> +described. I allude, of course, to <span class='sn'>Amphioxus</span>, which is by far the most +primitive or generalized type of vertebrated animal hitherto discovered. +Indeed, we may say that this remarkable creature is almost as nearly +allied to a worm as it is to a fish. For it has no specialized head, and +therefore no skull, brain, or jaws: it is destitute alike of limbs, of a +centralized heart, of developed liver, kidneys, and, in short, of most +of the organs which belong to the other Vertebrata. It presents, +however, a rudimentary backbone, in the form of what is called a +notochord. Now a primitive dorsal axis of this kind occurs at a very +early period of embryonic life in all vertebrated animals; but, with the +exception of <span class='sn'>Amphioxus</span>, in all other existing Vertebrata this structure +is not itself destined to become the permanent or bony vertebral column. +On the contrary, it gives way to, or is replaced by, this permanent bony +structure at a later stage of development. Consequently, it is very +suggestive that so distinctively embryonic a structure as this temporary +cartilaginous axis of all the other known Vertebrata should be found +actually persisting to the present day as the permanent axis of <span +class='sn'>Amphioxus</span>. In many other respects, likewise, the early +embryonic history of other Vertebrata refers us to the permanent +condition of <span class='sn'>Amphioxus</span>. In particular, we must notice that the wall of +the neck is always perforated by what in <span class='sn'>Amphioxus</span> are the +gill-openings, and that the blood-vessels as they proceed from the heart +are always distributed in the form of what are called gill-arches, +adapted to convey the blood round or through the gills for the purpose +of aeration. In all existing fish and other gill-breathing Vertebrata, +this <span class="pagenum"><a name="page_147" id="page_147">[147]</a></span> +arrangement is permanent. It is likewise met with in a peculiar kind of +worm, called <span class='sn'>Balanoglossus</span>—a creature so peculiar, indeed, that it +has been constituted by Gegenbaur a class all by itself. We can see by +the wood-cuts that it presents a series of gill-slits, like the +homologous parts of the fishes with which it is compared—i. e. +fishes of a comparatively low type of organization, which dates from a +time before the development of external gills. (Figs. 48, 49, 50.) Now, +as I have already said, these gill-<i>slits</i> are supported internally by +the gill-<i>arches</i>, or the blood-vessels which convey the blood to be +oxygenized in the branchial apparatus (see below, Figs. 51, 52, 53); and +the whole arrangement is developed from the anterior part of the +intestine—as is likewise the respiratory mechanism of all the +gill-breathing Vertebrata. That so close a parallel to this peculiar +mechanism should be met with in a worm, is a strong additional piece of +evidence pointing to the derivation of the Vertebrata from the Vermes.</p> + +<p class="pagenum"><a name="page_148" id="page_148">[148]</a></p> +<div class="figcenter" style="width: 600px;"> +<img src="images/i_165_048.jpg" width="600" height="163" alt="Balanoglossus." title="" /> +<span class="caption"><span class="sc">Fig</span>. 48.—<span class='sn'>Balanoglossus</span>. (After A. Agassiz.) <i>r</i>, +proboscis; <i>h</i>, collar; <i>k</i>, gill-slits; <i>d</i>, digestive posterior +intestine; <i>v</i>, intestinal vessel; <i>a</i>, anus.</span></div> + +<div class="figcenter" style="width: 600px;"> +<img src="images/i_165_049.jpg" width="600" height="80" alt="A large Sea-lamprey (Petromyzon marinus)" title="" /> +<span class="caption"><span class="sc">Fig</span>. 49.—A large Sea-lamprey (<span class='sn'>Petromyzon marinus</span>), much +reduced in size. (After Cuvier and Häckel.) A series of seven gill-slits +are visible.</span></div> + +<p class="pagenum"><a name="page_149" id="page_149">[149]</a></p> +<div class="figcenter" style="width: 600px;"> +<img src="images/i_166_050.jpg" width="600" height="216" alt="Adult Shark (Carcharias melanopterus)." title="" /> +<span class="caption"><span class="sc">Fig</span>. 50—Adult Shark (<span class='sn'>Carcharias melanopterus</span>). (After +Cuvier and Häckel.)</span></div> + +<p class="pagenum"><a name="page_150" id="page_150">[150]</a></p> +<div class='center'><table border="0" cellpadding="4" cellspacing="0" summary=""> +<tr><td class='c'><div class="figcenter" style="width: 200px;"> +<img src="images/i_167_051.jpg" width="200" height="298" alt="Heart and gill-arches of a fish." title="" /> +<div class='captionh'><span class="sc">Fig</span>. 51.—Diagram of heart and gill-arches of a fish. +(After Owen.)</div></div></td> +<td class='c'><div class="figcenter" style="width: 200px;"> +<img src="images/i_167_052.jpg" width="200" height="245" alt="One gill-arch, with branchial fringe attached." title="" /> +<div class="captionh"><span class="sc">Fig</span>. 52.—One gill-arch, with branchial fringe attached. +(After Owen.) H, Heart.</div></div></td> +<td class='c'><div class="figcenter" style="width: 200px;"> +<img src="images/i_167_053.jpg" width="200" height="357" alt="Heart and gill-arches in a lizard." title="" /> +<div class="captionh"><span class="sc">Fig</span>. 53.—Diagram of heart and gill-arches in a lizard. +(After Owen.) The gill-arches, <i>a a' a''</i>, and <i>b b' b''</i>, are called +aortic arches in air-breathing vertebrata.</div></div></td></tr></table></div> + +<p><span class="pagenum"><a name="page_147c" id="page_147c">[147c.]</a></span> +Well, I have just said that in all the gill-breathing Vertebrata, this +mechanism of gill-slits and vascular gill-arches in the front part of +the intestinal tract is permanent. But in the air-breathing Vertebrata +such an arrangement would obviously be of no use. Consequently, the +gill-slits in the sides of the neck (see Figs. 16 and 57, 58), and the +gill-arches of the large blood-vessels (Figs. 54, 55, 56), are here +exhibited only as transitory phases of development. But as such they +occur in all air-breathing Vertebrata. And, as if to make the homologies +as striking as possible, at the time when the gill-slits and the +gill-arches are developed in the embryonic young of air-breathing<span class="pagenum"><a name="page_154" id="page_154">[154]</a></span> +Vertebrata, the heart is constructed upon the fish-like type. That is to +say, it is placed far forwards, and, from having been a simple tube as +in Worms, is now divided into two chambers, as in Fish. Later on it +becomes progressively pushed further back between the developing lungs, +while it progressively acquires the three cavities distinctive of +Amphibia, and finally the four cavities belonging only to the complete +double circulation of Birds and Mammals. Moreover, it has now been +satisfactorily shown that the lungs of air-breathing Vertebrata, which +are thus destined to supersede the function of gills, are themselves the +modified swim-bladder or float, which belongs to Fish. Consequently, all +these progressive modifications in the important organs of circulation +and respiration in the air-breathing Vertebrata, together make up as +complete a history of their aquatic pedigree as it would be possible for +the most exacting critic to require.</p> + +<p class="pagenum"><a name="page_151" id="page_151">[151]</a></p> +<div class='center'><table border="0" cellpadding="4" cellspacing="0" summary=""> +<tr><td class='c'><div class="figcenter" style="width: 200px;"> +<img src="images/i_168_054.jpg" width="200" height="316" alt="Ideal diagram of primitive gill- or +aortic-arches." title="" /> +<div class="captionh"><span class="sc">Fig</span>. 54.—Ideal diagram, of primitive gill- or +aortic-arches. (After Rathke.) H, outline of heart. The arrows show the +course of the blood.</div></div></td> +<td class='c'><div class="figcenter" style="width: 200px;"> +<img src="images/i_168_055.jpg" width="200" height="292" alt="The same, modified for a bird." title="" /> +<div class="captionh"><span class="sc">Fig</span>. 55.—The same, modified for a bird. (After Le +Conte.) The dark lines show the aortic arches which persist. A, aorta; +<span class='sn'>p</span>, pulmonary arches; SC, S'C', sub-clavian; C, C', carotids.</div></div></td> +<td class='c'><div class="figcenter" style="width: 200px;"> +<img src="images/i_168_056.jpg" width="200" height="302" alt="The same, modified for a mammal." title="" /> +<div class="captionh"><span class="sc">Fig</span>. 56.—The same, modified for a mammal. (After Le +Conte.)</div></div></td></tr></table></div> + +<p class="pagenum"><a name="page_152" id="page_152">[152]</a></p> +<div class="figcenter" style="width: 296px;"> +<img src="images/i_169_057.jpg" width="296" height="500" alt="A series of embryos +of the classes of vertebrated animals below the Mammalia" title="" /> +<div class="captionh"><span class="sc">Fig</span>. 57.—A series of embryos at three comparable and +progressive stages of development (marked I, II, III), representing each +of the classes of vertebrated animals below the Mammalia (After +Häckel.)</div></div> + +<p class="pagenum"><a name="page_153" id="page_153">[153]</a></p> +<div class="figcenter" style="width: 307px;"> +<img src="images/i_170_058.jpg" width="307" height="500" alt="Another series of embryos +representing four different divisions of the class Mammalia." title="" /> +<div class="captionh"><span class="sc">Fig</span>. 58.—Another series of embryos, also at three +comparable and progressive stages of development (marked I, II, III), +representing four different divisions of the class Mammalia. (After +Häckel.)</div></div> + +<p>If space permitted, it would be easy to present abundance of additional +evidence to the same effect from the development of the skeleton, the +skull, the brain, the sense-organs, and, in short, of every constituent +part of the vertebrate organization. Even without any anatomical +dissection, the similarity of all vertebrated embryos at comparable +stages of development admits of being strikingly shown, if we merely +place the embryos one beside the other. Here, for instance, are the +embryos of a fish, a salamander, a tortoise, a bird, and four different +mammals. In each case three comparable stages of development are +represented. Now, if we read the series horizontally, we can see that +there is very little difference between <span class="pagenum"><a name="page_155" id="page_155">[155]</a></span> +the eight animals at the earliest of the three stages +represented—all having fish-like tails, gill-slits, and so on. In +the next stage further differentiation has taken place, but it will be +observed that the limbs are still so rudimentary that even in the case +of Man they are considerably shorter than the tail. But in the third +stage the distinctive characters are well marked.</p> + +<hr class='minor' /> + +<p>So much then for an outline sketch of the main features in the embryonic +history of the Vertebrata. But it must be remembered that the science of +comparative embryology extends to each of the other three great branches +of the tree of life, where these take their origin, through the worms, +from the still lower, or gastræa, forms. And in each of these three +great branches—namely, the Echinodermata, the Mollusca, and the +Arthropoda—we have a repetition of just the same kind of evidence in +favour of continuous descent, with adaptive modification in sundry +lines, as that which I have thus briefly sketched in the case of the +Vertebrata. The roads are different, but the method of travelling is the +same. Moreover, when the embryology of the Worms is closely studied, the +origin of these different roads admits of being clearly traced. So that +when all this mass of evidence is taken together, we cannot wonder that +evolutionists should now regard the science of comparative embryology as +the principal witness to their theory.</p> + +<hr /><p class="pagenum"><a name="page_156" id="page_156">[156]</a></p> +<h2><a name="CHAPTER_V" id="CHAPTER_V"></a>CHAPTER V.<br /><br /> +<span class="sc">Palæontology.</span></h2> + +<p>The present Chapter will be devoted to a consideration of the evidence +of organic evolution which has been furnished by the researches of +geologists. On account of its direct or historical nature, this branch +of evidence is popularly regarded as the most important—so much so, +indeed, that in the opinion of most educated persons the whole doctrine +of organic evolution must stand or fall according to the so-called +“testimony of the rocks.” Now, without at all denying the +peculiar importance of this line of evidence, I must begin by remarking +that it does not present the denominating importance which popular +judgment assigns to it. For although popular judgment is right in +regarding the testimony of the rocks as of the nature of a history, this +judgment, as a rule, is very inadequately acquainted with the great +imperfections of that history. Knowing in a general way what magnificent +advances the science of geology has made during the present century, the +public mind is more or less imbued with the notion, that because we now +possess a tolerably complete record of the chronological succession of +geological formations, we must therefore possess a correspondingly +complete <span class="pagenum"><a name="page_157" id="page_157">[157]</a></span> +record of the chronological succession of the forms of life which from +time to time have peopled the globe. Now in one sense this notion is +partly true, but in another sense it is profoundly false. It is partly +true if we have regard only to those larger divisions of the vegetable +or animal kingdoms which naturalists designate by the terms classes and +orders. But the notion becomes progressively more untrue when it is +applied to families and genera, while it is most of all untrue when +applied to species. That this must be so may be rendered apparent by two +considerations.</p> + +<p>In the first place, it does not follow that because we have a tolerably +complete record of the succession of geological formations, we have +therefore any correspondingly complete record of their fossiliferous +contents. The work of determining the relative ages of the rocks does +not require that every cubic mile of the earth’s surface should be +separately examined, in order to find all the different fossils which it +may contain. Were this the case, we should hitherto have made but very +small progress in our reading of the testimony of the rocks. The +relative ages of the rocks are determined by broad comparative surveys +over extensive areas; and although the identification of widely +separated deposits is often greatly assisted by a study of their +fossiliferous contents, the mere pricking of a continent here and there +is all that is required for this purpose. Hence, the accuracy of our +information touching the relative ages of geological strata does not +depend upon—and, therefore, does not betoken—any equivalent accuracy +of knowledge touching the fossiliferous material which these strata may +at the present time actually contain. And, <span class="pagenum"><a name="page_158" id="page_158">[158]</a></span> +as we well know, the opportunities which the geologist has of +discovering fossils are extremely limited, if we consider these +opportunities in relation to the area of geological formations. The +larger portion of the earth’s surface is buried beneath the sea; +and much the larger portion of the fossiliferous deposits on shore are +no less hopelessly buried beneath the land. Therefore it is only upon +the fractional portion of the earth’s surface which at the present +time happens to be actually exposed to his view that the geologist is +able to prosecute his search for fossils. But even here how miserably +inadequate this search has hitherto been! With the exception of a +scratch or two in the continents of Asia and America, together with a +somewhat larger number of similar scratches over the continent of +Europe, even that comparatively small portion of the earth’s +surface which is available for the purpose has been hitherto quite +unexplored by the palæontologist. How enormously rich a store of +material remains to be unearthed by the future scratchings of this +surface, we may dimly surmise from the astonishing world of bygone life +which is now being revealed in the newly discovered fossiliferous +deposits on the continent of America.</p> + +<p>But, besides all this, we must remember, in the second place, that all +the fossiliferous deposits in the world, even if they could be +thoroughly explored, would still prove highly imperfect, considered as a +history of extinct forms of life. In order that many of these forms +should have been preserved as fossils, it is necessary that they should +have died upon a surface neither too hard nor too soft to admit of their +<span class="pagenum"><a name="page_159" id="page_159">[159]</a></span> +leaving an impression; that this surface should afterwards have +hardened sufficiently to retain the impression; that it should then have +been protected from the erosion of water, as well as from the +disintegrating influence of the air; and yet that it should not have +sunk far enough beneath the surface to have come within the no less +disintegrating influence of subterranean heat. Remembering thus, as a +general rule, how many conditions require to have met before a fossil +can have been both formed and preserved, we must conclude that the +geological record is probably as imperfect in itself as are our +opportunities of reading even the little that has been recorded. If we +speak of it as a history of the succession of life upon the planet, we +must allow, on the one hand, that it is a history which merits the name +of a “chapter of accidents"; and, on the other hand, that during +the whole course of its compilation pages were being destroyed as fast +as others were being formed, while even of those that remain it is only +a word, a line, or at most a short paragraph here and there, that we are +permitted to see. With so fragmentary a record as this to study, I do +not think it is too much to say that no conclusions can be fairly based +upon it, merely from the absence of testimony. Only if the testimony +were positively opposed to the theory of descent, could any argument be +fairly raised against that theory on the grounds of this testimony. In +other words, if any of the fossils hitherto discovered prove the order +of succession to have been incompatible with the theory of genetic +descent, then the record may fairly be adduced in argument, because we +should then be in possession of definite information <span class="pagenum"><a name="page_160" id="page_160">[160]</a></span> +of a positive kind, instead of a mere absence of information of any +kind. But if the adverse argument reaches only to the extent of +maintaining that the geological record does not furnish us with so +complete a series of “connecting links” as we might have +expected, then, I think, the argument is futile. Even in the case of +human histories, written with the intentional purpose of conveying +information, it is an unsafe thing to infer the non-occurrence of an +event from a mere silence of the historian—and this especially in +matters of comparatively small detail, such as would correspond (in the +present analogy) to the occurrence of <i>species</i> and <i>genera</i> as +connecting links. And, of course, if the history had only come down to +us in fragments, no one would attach any importance at all to what might +have been only the <i>apparent</i> silence of the historian.</p> + +<p>In view, then, of the unfortunate imperfection of the geological record +<i>per se</i>, as well as of the no less unfortunate limitation of our means +of reading even so much of the record as has come down to us, I conclude +that this record can only be fairly used in two ways. It may fairly be +examined for positive testimony against the theory of descent, or for +proof of the presence of organic remains of a high order of development +in a low level of strata. And it may be fairly examined for negative +testimony, or for the absence of connecting links, if the search be +confined to the larger taxonomic divisions of the fauna and flora of the +world. The more minute these divisions, the more restricted must have +been the areas of their origin, and hence the less likelihood of their +having been preserved in the <span class="pagenum"><a name="page_161" id="page_161">[161]</a></span> +fossil state, or of our finding them even if they have been. Therefore, +if the theory of evolution is true, we ought not to expect from the +geological record a full history of <i>specific</i> changes in any but at +most a comparatively small number of instances, where local +circumstances happen to have been favourable for the writing and +preservation of such a history. But we might reasonably expect to find a +general concurrence of geological testimony to the larger +fact—namely, of there having been throughout all geological time a +uniform progression as regards the larger taxonomic divisions. And, as I +will next proceed to show, this is, in a general way, what we do find, +although not altogether without some important exceptions, with which I +shall deal in an Appendix.</p> + +<p>There is no <i>positive</i> proof <i>against</i> the theory of descent to be drawn +from a study of palæontology, or proof of the presence of any kind of +fossils in strata where the fact of their presence is incompatible with +the theory of evolution. On the other hand, there is an enormous body of +uniform evidence to prove two general facts of the highest importance in +the present connexion. The first of these general facts is, that an +increase in the diversity of types both of plants and animals has been +constant and progressive from the earliest to the latest times, as we +should anticipate that it must have been on the theory of descent in +ever-ramifying lines of pedigree. And the second general fact is, that +through all these branching lines of ever-multiplying types, from the +first appearance of each of them to their latest known conditions, there +is overwhelming evidence of one great law of organic nature—the law of +gradual advance from the general <span class="pagenum"><a name="page_162" id="page_162">[162]</a></span> +to the special, from the low to the high, from the simple to the +complex.</p> + +<p>Now, the importance of these large and general facts in the present +connexion must be at once apparent; but it may perhaps be rendered more +so if we try to imagine how the case would have stood supposing +geological investigation to have yielded in this matter an opposite +result, or even so much as an equivocal result. If it had yielded an +opposite result, if the lower geological formations were found to +contain as many, as diverse, and as highly organized types as the later +geological formations, clearly there would have been no room at all for +any theory of progressive evolution. And, by parity of reasoning, in +whatever degree such a state of matters were found to prevail, in that +degree would the theory in question have been discredited. But seeing +that these opposite principles do not prevail in any (relatively +speaking) considerable degree<a name="FNanchor_16_16" id="FNanchor_16_16"></a><a href="#Footnote_16_16" class="fnanchor">[16]</a>, +we have so far positive testimony of the largest and most massive +character in favour of this theory. For while all these large and +general facts are very much what they ought to be according to this +theory, they cannot be held to lend any support at all to the rival +theory. In other words, it is clearly no essential part of the theory of +special creation that species should everywhere exhibit this gradual +multiplication as to number, coupled with a gradual diversification and +general elevation of types, in all the growing branches of the tree of +life. No one could adopt seriously the jocular lines of Burns, to the +effect that the Creator required to practise his <span class="pagenum"><a name="page_164" id="page_164">[164]</a></span> +prentice hand on lower types before advancing to the formation of +higher. Yet, without some such assumption, it would be impossible to +explain, on the theory of independent creations, why there should have +been this gradual advance from the few to the many, from the general to +the special, from the low to the high.</p> + +<p><span class="pagenum"><a name="page_163" id="page_163">[163]</a></span></p> + +<div class='center'> +<table border="1" cellpadding="4" cellspacing="0" summary=""> +<tr><td class='l'></td><td class='c'><i>Epochs and Formations.</i></td><td class='c'><i>Faunal Characters.</i></td></tr> +<tr><td class='l' rowspan='5'><span class="sc">Cainozoic</span> or <span class="sc">Tertiary</span></td><td class='l'><span class="sc">Post-Pliocene.</span><br /> Glacial Period.</td><td class='l'>Man. Mammalia principally of living species. Mollusca exclusively recent.</td></tr> +<tr><td class='l'><span class="sc">Pliocene</span>, 3,000 feet.</td><td class='l'>Mammalia principally of recent genera—living species rare. Mollusca very modern.</td></tr> +<tr><td class='l'><span class="sc">Miocene</span>, 4,000 ft.</td><td class='l' rowspan='2'>Mammalia principally of living families; extinct genera numerous; species all extinct. Mollusca largely of recent species.</td></tr> +<tr><td class='l'><span class="sc">Oligocene</span>, 8,000 ft.</td></tr> +<tr><td class='l'><span class="sc">Eocene</span>, 10,000 ft.</td><td class='l'>Mammalia with numerous extinct families and orders; all the species and most of the genera extinct. Modern type Shell-Fish.</td></tr> +<tr><td class='l' rowspan='4'><span class="sc">Mesozoic</span> or <span class="sc">Secondary</span></td><td class='l'><span class="sc">Laramie</span>, 4,000 ft.</td><td class='l'>Passage beds.</td></tr> +<tr><td class='l'><span class="sc">Cretaceous</span>, 12,000 ft.</td><td class='l'>Dinosaurian (bird-like) Reptiles; Pterodactyls (flying Reptiles); toothed Birds; earliest Snake; bony Fishes; Crocodiles; Turtles; Ammonites.</td></tr> +<tr><td class='l'><span class="sc">Jurassic</span>, 6,000 ft.<br /> Oolite.<br /> Lias.</td><td class='l'>Earliest Birds; giant Reptiles (Ichthyosaurs, Dinosaurs, Pterodactyls); Ammonites; Clam- and Snail-Shells very abundant; decline of Brachiopods; Butterfly.</td></tr> +<tr><td class='l'><span class="sc">Trias</span>, 5,000 ft.<br /> New Red Sandstone.</td><td class='l'>First Mammalian (Marsupial); 2-gilled Cephalopods (Cuttle-Fishes, Belemnites); reptilian Foot-Prints.</td></tr> +<tr><td class='l' rowspan='5'><span class="sc">Paleozoic</span> or <span class="sc">Primary</span></td><td class='l'><span class="sc">Permian</span>, 5,000 ft.</td><td class='l'>Earliest true Reptiles.</td></tr> +<tr><td class='l'><span class="sc">Carboniferous</span>, 26,000 ft.<br /> Coal.</td><td class='l'>Earliest Amphibian (Labyrinthodont); extinction of Trilobites; first Cray-fish; Beetles; Cockroaches; Centipedes; Spiders.</td></tr> +<tr><td class='l'><span class="sc">Devonian</span>, 18,000 ft.<br /> Old Red Sandstone.</td><td class='l'>Cartilaginous and Ganoid Fishes; earliest and (snail) and freshwater Shells; Shell-Fish abundant; decline of Trilobites; May-flies; Crab.</td></tr> +<tr><td class='l'><span class="sc">Silurian</span>, 33,000 ft.</td><td class='l'>Earliest Fish; the first Air-Breathers (Insect, Scorpion); Brachiopods and 4-gilled Cephalopods very abundant; Trilobites; Corals; Graptolites.</td></tr> +<tr><td class='l'><span class="sc">Cambrian</span>, 24,000 ft.</td><td class='l'>Trilobites; Brachiopod Mollusks.</td></tr> +<tr><td class='l' rowspan='2'><span class="sc">Azoic</span></td><td class='l'><span class="sc">Archæan</span>, 30,000 ft.<br /> Huronian.<br /> Laurentian.</td><td class='l'>Eozoön, (probably not a fossil).</td></tr> +<tr><td class='l'><span class="sc">Primeval</span>.</td><td class='l'>Non-sedimentary.</td></tr> +</table></div> + +<p><span class="pagenum"><a name="page_164c" id="page_164c">[164c.]</a></span> +I submit, then, that so far as the largest and most general principles +in the matter of palæontology are concerned, we have about as strong and +massive a body of evidence as we could reasonably expect this branch of +science to yield; for it is at once enormous in amount and positive in +character. Therefore, if I do not further enlarge upon the evidence +which we here have, as it were <i>en masse</i>, it is only because I do not +feel that any words could add to its obvious significance. It may best +be allowed to speak for itself in the millions of facts which are +condensed in this tabular statement of the order of succession of all +the known forms of animal life, as presented by the eminent +palæontologist, Professor Cope<a name="FNanchor_17_17" id="FNanchor_17_17"></a><a href="#Footnote_17_17" class="fnanchor">[17]</a>.</p> + +<p>Or, taking a still more general survey, this tabular statement may be +still further condensed, and presented in a diagrammatic form, as it has +been by another eminent American palæontologist, Prof. Le Conte, in his +excellent little treatise on <i>Evolution and its Relations to Religious +Thought</i>. The following is his diagrammatic representation, with his +remarks thereon.</p> + +<div class='blockquot'><p>When each ruling class declined in importance, it did not perish, +but continued in a subordinate position. Thus, the whole organic +kingdom became not only higher and higher in its highest forms, but +also more and more complex in its structure <span class="pagenum"><a name="page_165" id="page_165">[165]</a></span> +and in the interaction of its correlated parts. The whole process and +its result is roughly represented in the accompanying diagram, in which +A B represents the course of geological time, and the curve, the rise, +culmination, and decline of successive dominate classes.</p></div> + +<div class="figcenter" style="width: 500px;"> +<img src="images/i_182_059.jpg" width="500" height="82" alt="Geological Succession of the Classes of the Animal Kingdom." title="" /> +<span class="caption"><span class="sc">Fig</span>. 59.—Diagram of Geological Succession of the Classes +of the Animal Kingdom. (After Le Conte.)</span></div> + +<p>I will here leave the evidence which is thus yielded by the most general +principles that have been established by the science of palæontology; +and I will devote the rest of this chapter to a detailed consideration +of a few highly special lines of evidence. By thus suddenly passing from +one extreme to the other, I hope to convey the best idea that can be +conveyed within a brief compass of the minuteness, as well as the +extent, of the testimony which is furnished by the rocks.</p> + +<hr class='minor' /> + +<p>When Darwin first published his <i>Origin of Species</i>, adverse critics +fastened upon the “missing-link” argument as the strongest +that they could bring against the theory of descent. Although Darwin had +himself strongly insisted on the imperfection of the geological record, +and the consequent precariousness of any negative conclusions raised +upon it, these critics maintained that he was making too great a demand +upon the argument from ignorance—that, even allowing for the +imperfection of the record, they would certainly have expected at least +a few cases of testimony to <i>specific</i> transmutation. For, they urged in +effect, looking to <span class="pagenum"><a name="page_166" id="page_166">[166]</a></span> +the enormous profusion of the extinct species on the one hand, and to +the immense number of known fossils on the other, it was incredible that +no satisfactory instances of specific transmutation should ever have +been brought to light, if such transmutation had ever occurred in the +universal manner which the theory was bound to suppose. But since Darwin +first published his great work palæontologists have been very +active in discovering and exploring fossiliferous beds in sundry parts +of the world; and the result of their labours has been to supply so many +of the previously missing links that the voice of competent criticism in +this matter has now been well-nigh silenced. Indeed, the material thus +furnished to an advocate of evolution at the present time is so abundant +that his principal difficulty is to select his samples. I think, +however, that the most satisfactory result will be gained if I restrict +my exposition to a minute account of some few series of connecting +links, rather than if I were to take a more general survey of a larger +number. I will, therefore, confine the survey to the animal kingdom, and +there mention only some of the cases which have yielded well-detailed +proof of continuous differentiation.</p> + +<p>It is obvious that the parts of animals most likely to have been +preserved in such a continuous series of fossils as the present line of +evidence requires, would have been the hard parts. These are horns, +bones, teeth, and shells. Therefore I will consider each of these four +classes of structures separately.</p> + +<hr class='minor' /> + +<p>Horns wherever they occur, are found to be of high importance for +purposes of classification. They are <span class="pagenum"><a name="page_167" id="page_167">[167]</a></span> +restricted to the Ruminants, and appear under three different forms or +types—namely solid, as in antelopes; hollow, as in sheep; and +deciduous, as in deer. Now, in each of these divisions we have a +tolerably complete palæontological history of the evolution of +horns. The early ruminants were altogether hornless (Fig. 60). Then, in +the middle Miocene, the first antelopes appeared with tiny horns, which +progressively increased in size among the ever-multiplying species of +antelopes until the present day. But it is in the deer tribe that we +meet with even better evidence touching the progressive evolution of +horns; because here not only size, but shape, is concerned. For +deer’s horns, or antlers, are arborescent; and hence in their case +we have an opportunity of reading the history, not only of a progressive +growth in size, but also of an increasing development of form. Among the +older members of the tribe, in the lower Miocene, there are no horns at +<span class="pagenum"><a name="page_168" id="page_168">[168]</a></span> +all. In the mid-Miocene we meet with two-pronged horns (<span class='sn'>Cervus +dicrocerus</span>, Figs. 61, 62, 1/5 nat. size). Next, in the upper Miocene +(<span class='sn'>C. matheronis</span>, Fig. 63, 1/8 nat. size), and extending into the Pliocene +(<span class='sn'>C. pardinensis</span>, Fig. 64, 1/18 nat. size), we meet with three-pronged +horns. Then, in the Pliocene we find also four-pronged horns (<span class='sn'>C. +issiodorensis</span>, Fig. 65, 1/16 nat. size), leading us to five-pronged +(<span class='sn'>C. tetraceros</span>). Lastly, in the Forest-bed of Norfolk we meet with +arborescent horns (<span class='sn'>C. Sedgwickii</span>, Fig. 66, 1/35 nat. size). The +life-history of existing stags furnishes a parallel development (Fig. +67), beginning with a single horn (which has not yet been found +palæontologically), going on to two prongs, three prongs, four +prongs, and afterwards branching.</p> + +<p class="pagenum"><a name="page_167i" id="page_167i">[167i.]</a></p> +<div class="figcenter" style="width: 400px;"> +<img src="images/i_184_060.jpg" width="400" height="288" alt="Fig. 60.—Skull of Oreodon Culbertsoni. (After Leidy.)" title="" /> +<span class="caption"><span class='sc'>Fig.</span> 60.—Skull of Oreodon Culbertsoni. (After Leidy.)</span></div> + +<p class="pagenum"><a name="page_168i" id="page_168i">[168i.]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_185_061-6.jpg" width="500" height="275" alt="Evolution of horns." title="" /> +<span class="caption"><span class='sc'>Figs.</span> 61-6. The series is reduced from Gaudry’s illustrations, after Farge, +Croizet, Jobert and Boyd Dawkins.</span></div> + +<p class="pagenum"><a name="page_169i" id="page_169i">[169i.]</a></p> +<div class="figcenter" style="width: 400px;"> +<img src="images/i_186_067.jpg" width="400" height="199" alt="Successive stages in the development of an +existing Deer’s Antlers." title="" /> +<span class="caption"><span class='sc'>Fig.</span> 67. Successive stages in the development of an +existing Deer’s Antlers. (After Gaudry, but a better illustration +has already been given on p. 100.)</span></div> + +<p class="pagenum"><a name="page_168c" id="page_168c">[168c.]</a></p><hr class='minor' /> + +<p>Coming now to bones, we have a singularly complete record of transition +from one type or pattern of <span class="pagenum"><a name="page_169" id="page_169">[169]</a></span> +structure to another in the phylogenetic history of tails. This has +been so clearly and so tersely conveyed by Prof. Le Conte, that I cannot +do better than quote his statement.</p> + +<p class="pagenum"><a name="page_170" id="page_170">[170]</a></p> + +<div class='blockquot'><p>It has long been noticed that there are among fishes two styles of +tail-fins. These are the even-lobed, or homocercal (Fig. 68), and +the uneven-lobed, or heterocercal (Fig. 69). The one is +characteristic of ordinary fishes (teleosts), the other of sharks +and some other orders. In structure the difference is even more +fundamental than in form. In the former style the backbone stops +abruptly in a series of short, enlarged joints, and thence sends +off rays to form the tail-fin (Fig. 68); in the latter the +<span class="pagenum"><a name="page_171" id="page_171">[171]</a></span> +backbone runs through the fin to its very point, growing slenderer by +degrees, and giving off rays above and below from each joint, but the +rays on the lower side are much longer (Fig. 69). This type of fin is, +therefore, <i>vertebrated</i>, the other <i>non-vertebrated</i>. Figs. 68 and 69 +show these two types in form and structure. But there is still another +type found only in the lowest and most generalized forms of fishes. In +these the tail-fin is vertebrated and yet symmetrical. This type is +shown in Fig. 70.</p> + +<div class="figcenter" style="width: 200px;"> +<img src="images/i_186_068.jpg" width="200" height="328" alt="Homocercal Tail." title="" /> +<span class="caption"><span class="sc">Fig</span>. 68.—Homocercal Tail, showing (A) external form +and (B) internal structure.</span></div> + +<p class="pagenum"><a name="page_170i" id="page_170i">[170i.]</a></p> +<div class="figcenter" style="width: 300px;"> +<img src="images/i_187_069.jpg" width="300" height="266" alt="Heterocercal Tail." title="" /> +<span class="caption"><span class="sc">Fig</span>. 69.—Heterocercal Tail, showing (A) external +form and (B) internal structure.</span></div> + +<div class="figcenter" style="width: 300px;"> +<img src="images/i_187_070.jpg" width="300" height="146" alt="Diphycercal fin." title="" /> +<span class="caption"><span class="sc">Fig</span>. 70.—Vertebrated but symmetrical fin +(diphycercal), showing (A) external form and (B) internal +structure.</span></div> + +<p><span class="pagenum"><a name="page_171c" id="page_171c">[171c.]</a></span> +Now, in the development of a teleost fish (Fig. 68), as has been +shown by Alexander Agassiz, the tail-fin is first like Fig. 70; +then becomes heterocercal, like Fig. 69; and, finally, becomes +homocercal like Fig. 68. Why so? Not because there is any special +advantage in this succession of forms; for the changes take place +either in the egg or else in very early embryonic states. The +answer is found in the fact that <i>this is the order of change in +the phylogenetic series</i>. The earliest fish-tails were either like +Fig. 69 or Fig. 70; never like Fig. 68. The <span class="pagenum"><a name="page_173" id="page_173">[173]</a></span> +earliest of all were almost certainly like Fig. 70; then they became +like Fig. 69; and, finally, only much later in geological history +(Jurassic or Cretaceous), they became like Fig. 68. This order of change +is still retained in the embryonic development of the last introduced +and most specialized order of existing fishes. The family history is +repeated in the individual history.</p> + +<p class="pagenum"><a name="page_171i" id="page_171i">[171i.]</a></p> +<div class='center'><table border="0" cellpadding="4" cellspacing="0" summary=""> +<tr><td style='vertical-align: bottom;'><div class="figcenter" style="width: 150px;"> +<img src="images/i_188_071.jpg" width="150" height="374" alt="Fig. 71.—Tail of Archæopteryx." title="" /> +<div class="captionh"><span class="sc">Fig</span>. 71.—Tail of <span class='sn'>Archæopteryx</span>. A indicates origin +of simply-jointed tail.</div></div></td> +<td style='vertical-align: bottom;'><div class="figcenter" style="width: 200px;"> +<img src="images/i_188_072.jpg" width="200" height="301" alt="Fig. 72.—Tail of modern Bird." title="" /> +<div class="captionh"><span class="sc">Fig</span>. 72.—Tail of modern Bird. The numerals indicate +the foreshortened, enlarged, and consolidated joints; <i>f</i>, terminal +segment of the vertebral column; D, shafts of feathers.</div></div></td></tr></table></div> + +<p class="pagenum"><a name="page_172" id="page_172">[172]</a></p> +<div class="figcenter" style="width: 292px;"> +<img src="images/i_189_073.jpg" width="292" height="500" alt="Fig. 73.—Archæopteryx macura, restored." title="" /> +<span class="caption"><span class="sc">Fig</span>. 73.—<span class='sn'>Archæopteryx macura</span>, restored, ½ nat. +size. (After Flower.) The section of the tail is copied from Owen, +nat. size.</span></div> +</div> + +<p><span class="pagenum"><a name="page_173c" id="page_173c">[173c.]</a></span> +Similar changes have taken place in the form and structure of +birds’ tails. The earliest bird known—the Jurassic <span +class='sn'>Archæopteryx</span>—had a long reptilian tail of twenty-one +joints, each joint bearing a feather on each side, right and left +(Fig. 71): [see also Fig. 73]. In the typical modern bird, on the +contrary, the tail-joints are diminished in number, shortened up, +and enlarged, and give out long feathers, fan-like, to form the +so-called tail (Fig. 72). The <span class='sn'>Archæopteryx’</span> tail is +<i>vertebrated</i>, the typical bird’s <i>non-vertebrated</i>. This +shortening up of the tail did not take place at once, but +gradually. The Cretaceous birds, intermediate in time, had tails +intermediate in structure. The <span class='sn'>Hesperornis</span> of Marsh had twelve +joints. At first—in Jurassic strata—the tail is fully a half of +the whole vertebral column. It then gradually shortens up until it +becomes the aborted organ of typical modern birds. Now, in +embryonic development, the tail of the modern typical bird <i>passes +through all these stages</i>. At first the tail is nearly one half the +whole vertebral column; then, as development goes on, while the +rest of the body grows, the growth of the tail stops, and thus +finally becomes the aborted organ we now find. The ontogeny still +passes through the stages of the phylogeny. The same is true of all +tailless animals.</p> + +<p>The extinct <span class='sn'>Archæopteryx</span> above alluded to presents throughout its whole +organization a most interesting assemblage of “generalized +characters.” For example, its teeth, and its still unreduced +digits of the wings (which, like those of the feet, are covered with +scales), refer us, with almost as much force as does the vertebrated +tail, to the Sauropsidian type—or the trunk from which birds and +reptiles have diverged.</p> + +<p>We will next consider the palæontological evidence +<span class="pagenum"><a name="page_175" id="page_175">[175]</a></span> +which we now possess of the evolution of mammalian limbs, with special +reference to the hoofed animals, where this line of evidence happens to +be most complete.</p> + +<p>I may best begin by describing the bones as these <span class="pagenum"><a name="page_176" id="page_176">[176]</a></span> +occur in the sundry branches of the mammalian type now living. As we +shall presently see, the modifications which the limbs have undergone in +these sundry branches chiefly consist in the suppression of some parts +and the exaggerated development of others. But, by comparing all +mammalian limbs together, it is easy to obtain a generalized type of +mammalian limb, <span class="pagenum"><a name="page_177" id="page_177">[177]</a></span> +which in actual life is perhaps most nearly conformed to in the case of +bears. I will therefore choose the bear for the purpose of briefly +expounding the bones of mammalian limbs in general—merely asking +it to be <span class="pagenum"><a name="page_178" id="page_178">[178]</a></span> +understood, that although in the case of many other mammalia some of +these bones may be dwindled or altogether absent, while others may be +greatly exaggerated as to relative size, in no case do any <i>additional</i> +bones appear.</p> + +<p>On looking, then, at the skeleton of a bear (Fig. 74), the first thing +to observe is that there is a perfect serial homology between the bones +of the hind legs and of the fore legs. The thigh-bone, or femur, +corresponds to the shoulder-bone, or humerus; the two shank bones (tibia +and fibula) correspond to the two arm-bones (radius and ulna); the many +little ankle-bones (tarsals) correspond to the many little wrist-bones +(carpals); the foot-bones (meta-tarsals) correspond to the hand-bones +(meta-carpals); and, lastly, the bones of each of the toes correspond to +those of each of the fingers.</p> + +<p class="pagenum"><a name="page_174" id="page_174">[174.]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_191_074.jpg" width="500" height="321" alt="Skeleton of Polar Bear." title="" /> +<span class="caption"><span class="sc">Fig</span>. 74.—Skeleton of Polar Bear, drawn from nature +(<i>Brit. Mus.</i>).</span></div> + +<p>The next thing to observe is, that the disposition of bones in the case +of the bear is such that the animal walks in the way that has been +called plantigrade. That is to say, all the bones of the fingers, as +well as those of the toes, feet, and ankles, rest upon the ground, or +help to constitute the “soles.” Our own feet are constructed +on a closely similar pattern. But in the majority of living mammalian +forms this is not the case. For the majority of mammals are what has +been called digitigrade. That is to say, the bones of the limb are so +disposed that both the foot and hand bones, and therefore also the ankle +and wrist, are removed from the ground altogether, so that the animal +walks exclusively upon its toes and fingers—as in the case of this +skeleton (Fig. 75), which is the skeleton of a lion. The next figures +display a series of limbs, showing the progressive passage of a +completely <span class="pagenum"><a name="page_179" id="page_179">[179]</a></span> +plantigrade into a highly digitigrade type—the curved lines of +connexion serving to indicate the homologous bones (Figs. 76, 77).</p> + +<p class="pagenum"><a name="page_175i" id="page_175i">[175i.]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_192_075.jpg" width="500" height="333" alt="Skeleton of Lion." title="" /> +<span class="caption"><span class='sc'>Fig</span>. 75.—Skeleton of Lion. (After Huxley.)</span></div> + +<p class="pagenum"><a name="page_176i" id="page_176i">[176i.]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_193_076.jpg" width="500" height="344" alt="Anterior limb of Man, Dog, Hog, Sheep, and +Horse." title="" /> +<div class="captionh"><span class="sc">Fig</span>. 76.—Anterior limb of Man, Dog, Hog, Sheep, and +Horse. (After Le Conte.) <i>Sc</i>, shoulder-blade; <i>c</i>, coracoid; <i>a</i>, <i>b</i>, +bones of fore-arm; 5, bones of the wrist; 6, bones of the hand; 7, bones +of the fingers.</div></div> + +<div class="figcenter" style="width: 500px;"> +<img src="images/i_194_077.jpg" width="500" height="323" alt="Posterior limb of Man, Monkey, Dog, Sheep and +Horse." title="" /> +<div class="captionh"><span class="sc">Fig</span>. 77.—Posterior limb of Man, Monkey, Dog, Sheep and +Horse. (After Le Conte.) 1, Hip-joint; 2, thigh-bone; 3, knee-joint; 4, +bones of leg; 5, ankle-joint; 6, bones of foot; 7, bones of toes.</div></div> + +<p><span class="pagenum"><a name="page_179c" id="page_179c">[179c.]</a></span> +I will now proceed to detail the history of mammalian limbs, as this has +been recorded for us in fossil remains.</p> + +<div class="figcenter" style="width: 400px;"> +<img src="images/i_196_078.jpg" width="400" height="387" alt="Posterior limb of Baptanodon discus +and anterior limb of Chelydra serpentina." title="" /> +<div class="captionh"><span class="sc">Fig</span>. 78.—A, posterior limb of <span class='sn'>Baptanodon discus</span>. (After +Marsh.) F, thigh-bone; I to VI, undifferentiated bones of the leg and +foot. B, anterior limb of <span class='sn'>Chelydra serpentina</span>. (After Gegenbaur.) U and +R, bones of the fore-arm; I to V, fully differentiated bones of the +hand, following those of the wrist.</div></div> + +<p>The most generalized or primitive types of limb hitherto discovered in +any vertebrated animal above the class of fishes, are those which are +met with in some of the extinct aquatic reptiles. Here, for instance, is +a diagram of the left hind limb of <span class='sn'>Baptanodon discus</span> (Fig. 78). It has +six rows of little symmetrical bones springing from a leg-like origin. +<span class="pagenum"><a name="page_180" id="page_180">[180]</a></span> +<span class="figleft" style="width: 200px;"> +<img src="images/i_197_079.jpg" width="200" height="357" alt="Paddle of a Whale." title="" /> +<span class="caption"><span class="sc">Fig</span>. 79.—Paddle of a Whale.</span></span> +But the whole structure resembles the fin of a fish about as nearly as +it does the leg of a mammal. For not only are there six rows of bones, +instead of five, suggestive of the numerous rays which characterise the +fin of a fish; but the structure as a whole, having been covered over +with blubber and skin, was throughout flexible and unjointed—thus in +function, even more than in structure, resembling a fin. In this +respect, also, it must have resembled the paddle of a whale (see Fig. +79); but of course the great difference will be noted, that the paddle +of a whale reveals the dwindled though still clearly typical bones of a +true mammalian limb; so that although in outward form and function these +two paddles are alike, their inward structure clearly shows that while +the one testifies to the absence of evolution, the other testifies to +the presence of degeneration. If the paddle of <span class='sn'>Baptanodon</span> had occurred +in a whale, or the paddle of a whale had occurred in <span class='sn'>Baptanodon</span>, either +fact would in itself have been well-nigh destructive of the whole theory +of evolution.</p> + +<p>Such, then, is the most generalized as it is the most ancient type of +vertebrate limb above the class of <span class="pagenum"><a name="page_181" id="page_181">[181]</a></span> +fishes. Obviously it is a type suited only to aquatic life. +Consequently, when aquatic Vertebrata began to become terrestrial, the +type would have needed modification in order to serve for terrestrial +locomotion. In particular, it would have needed to gain in consolidation +and in firmness, which means that it would have needed also to become +jointed. Accordingly, we find that this archaic type gave place in +land-reptiles to the exigencies of these requirements. Here for example +is a diagram, copied from Gegenbaur, of the right fore-foot of <span class='sn'>Chelydra +serpentina</span> (Fig. 78). As compared with the homologous limb of its purely +aquatic predecessor, there is to be noticed the disappearance of one of +the six rows of small bones, a confluence of some of the remainder in +the other five rows, a duplication of the arm-bone into a radius and +ulna, in order to admit of jointed rotation of the hand, and a general +disposition of the small bones below these arm-bones, which clearly +foreshadows the joint of the wrist. Indeed, in this fore-foot of <span +class='sn'>Chelydra</span>, a child could trace all the principal homologies of +the mammalian counterpart, growing, like the next stage in a dissolving +view, out of the primitive paddle of <span class='sn'>Baptanodon</span>—namely, first the +radius and ulna, next the carpals, then the meta-carpals, and, lastly, +the three phalanges in each of the five digits.</p> + +<p>Such a type of foot no doubt admirably meets the requirements of slow +reptilian locomotion over swampy ground. But for anything like rapid +locomotion over hard and uneven ground, greater modifications would be +needed. Such modifications, however, need not be other in kind: it is +enough that they should continue in the same line of advance, so as to +reach a <span class="pagenum"><a name="page_182" id="page_182">[182]</a></span> +higher degree of firmness, combined with better joints. Accordingly we +find that this took place, not indeed among reptiles, whose habits of +cold-blooded life have not changed, but among their warm-blooded +descendants, the mammals. Moreover, when we examine the whole mammalian +series, we find that the required modifications must have taken place in +slightly different ways in three lines of descent simultaneously. We +have first the plantigrade and digitigrade modifications already +mentioned (pp. <a href="#page_178">178</a>, <a href="#page_179">179</a>) +Of these the plantigrade walking entailed least change, because most +resembling the ancestral or lizard-like mode of progression. All that +was here needed was a general improvement as to relative lengths of +bones, with greater consolidation and greater flexibility of joints. +Therefore I need not say anything more about the plantigrade division. +But the digitigrade modification necessitated a change of structural +plan, to the extent of raising the wrist and ankle joints off the +ground, so as to make the quadruped walk on its fingers and toes. We +meet with an interesting case of this transition in the existing hare, +which while at rest supports itself on the whole hind foot after the +manner of a plantigrade animal, but when running does so upon the ends +of its toes, after the manner of a digitigrade animal.</p> + +<p>It is of importance for us to note that this transition from the +original plantigrade to the more recent digitigrade type, has been +carried out on two slightly different plans in two different lines of +mammalian descent. The hoofed mammals—which are all digitigrade—are +sub-classified as artiodactyls and perissodactyls, i. e. even-toed and +odd-toed. Now, <span class="pagenum"><a name="page_183" id="page_183">[183]</a></span> +whether an animal has an even or an odd number of toes may seem a +curiously artificial distinction on which to found so important a +classification of the mammalian group. But if we look at the matter from +a less empirical and more intelligent point of view, we shall see that +the alternative of having an even or an odd number of toes carries with +it alternative consequences of a practically important kind to any +animal of the digitigrade type. For suppose an aboriginal five-toed +animal, walking on the ends of its five toes, to be called upon to +resign some of his toes. If he is left with an even number, it must be +two or four; and in either case the animal would gain the firmest +support by so disposing his toes as to admit of the axis of his foot +passing between an equal number of them—whether it be one or two +toes on each side. On the other hand, if our early mammal were called +upon to retain an odd number of toes, he would gain best support by +adjusting matters so that the axis of his foot should be coincident with +his middle toe, whether this were his only toe, or whether he had one on +either side of it. This consideration shows that the classification into +even-toed and odd-toed is not so artificial as it no doubt at first +sight appears. Let us, then, consider the stages in the evolution of +both these types of feet.</p> + +<p>Going back to the reptile <span class='sn'>Chelydra</span>, it will be observed that the axis of +the foot passes down the middle toe, which is therefore supported by two +toes on either side (Fig. 78). It may also be noticed that the wrist or +ankle bones do not interlock, either with one another or with the bones +of the hand or foot below <span class="pagenum"><a name="page_184" id="page_184">[184]</a></span> +them. This, of course, would give a weak foot, suited to slow +progression over marshy ground—which, as we have seen, was no +doubt the origin of the mammalian plantigrade foot. Here, for instance, +to all intents and purposes, is a similar type of foot, which +<span class="pagenum"><a name="page_185" id="page_185">[185]</a></span> +belonged to a very early mammal, antecedent to the elephant series, the +horse series, the rhinoceros, the hog, and, in short, all the known +hoofed mammalia (Fig. 80). It was presumably an inhabitant of swampy +ground, slow in its movements, and low in its intelligence.</p> + +<p class="pagenum"><a name="page_184i" id="page_184i">[184i.]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_201_080.jpg" width="500" height="296" alt="Fossil skeleton of Phenacodus primavus." title="" /> +<span class="caption"><span class="sc">Fig</span>. 80.—Fossil skeleton of <span class='sn'>Phenacodus primavus</span>. (After +Cope.)</span></div> + +<p><span class="pagenum"><a name="page_185c" id="page_185c">[185c.]</a></span> +But now, as we have seen, for more rapid progression on hard uneven +ground, a stronger and better jointed foot would be needed. Therefore we +find the bones of the wrist and ankle beginning to interlock, both among +themselves and also with those of the foot and hand immediately below +them. Such a stage of evolution is still apparent in the now existing +elephant. (See Fig. 81.)</p> + +<p class="pagenum"><a name="page_186" id="page_186">[186]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_203_081.jpg" width="500" height="236" alt="Bones of the foot of four forms of +the perissodactyl type." title="" /> +<div class="captionh"><span class="sc">Fig</span>. 81.—Bones of the foot of four different forms of +the perissodactyl type, showing gradual reduction in the number of +digits, coupled with a greater consolidation of the bones above the +digits. The series reads from right to left. Drawn from nature (<i>Brit. +Mus.</i>).</div></div> + +<p><span class="pagenum"><a name="page_185c2" id="page_185c2">[185c2.]</a></span> +Next, however, a still stronger foot was made by the still further +interlocking of the wrist and ankle bones, so that both the first and +second rows of them were thus fitted into each other, as well as into +the bones of the hand and foot beneath. This further modification is +clearly traceable in some of the earlier perissodactyls, and occurs in +the majority at the present time. Compare, for example, the greater +interlocking and consolidation of these small bones in the Rhinoceros as +contrasted with the Elephant (Fig. 81). Moreover, simultaneously with +these consolidating improvements in the mechanism of the wrist and ankle +joints, or possibly at a somewhat later period, a reduction in the +number of digits began to take place. This was a continuation of the +policy of consolidating the foot, analogous to the dropping out of the +sixth row of small bones in the paddle of <span class='sn'>Baptanodon</span>. +(Fig. 78.) In the pentadactyl plantigrade foot of the early mammals, the +first digit, being the shortest, was the <span class="pagenum"><a name="page_188" id="page_188">[188]</a></span> +first to leave the ground, to dwindle, and finally to disappear. More +work being thus thrown on the remaining four, they were strengthened by +interlocking with the wrist (or ankle) bones above them, as just +mentioned; and also by being brought closer together.</p> + +<p class="pagenum"><a name="page_187" id="page_187">[187]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_204_082.jpg" width="500" height="297" alt="Bones of the foot of four forms of +the artiodactyl type." title="" /> +<div class="captionh"><span class="sc">Fig</span>. 82.—Bones of the foot of four different forms of +the artiodactyl type, showing gradual reduction of the number of digits, +coupled with a greater consolidation of the bones above the digits. The +series reads from right to left. Drawn from nature (<i>Brit. Mus.</i>).</div></div> + +<p><span class="pagenum"><a name="page_188c" id="page_188c">[188c.]</a></span> +The changes which followed I will render in the words of Professor +Marsh.</p> + +<div class='blockquot'><p>Two kinds of reduction began. One leading to the existing +perissodactyl foot, and the other, apparently later, resulting in +the artiodactyl type. In the former the axis of the foot remained +in the middle of the third digit, as in the pentadactyl foot. [See +Fig. 81.] In the latter, it shifted to the outer side of this +digit, or between the third and fourth toe. [See Fig. 82.]</p> + +<p>In the further reduction of the perissodactyl foot, the fifth +digit, being shorter than the remaining three, next left the +ground, and gradually disappeared. [Fig. 81 B.] Of the three +remaining toes, the middle or axial one was the longest, and +retaining its supremacy as greater strength and speed were +required, finally assumed the chief support of the foot [Fig. 81 +C], while the outer digits left the ground, ceased to be of use, +and were lost, except as splint-bones [Fig. 81 D]. The feet of the +existing horse shows the best example of this reduction in the +Perissodactyls, as it is the most specialized known in the +Ungulates [Fig. 81 D].</p> + +<p>In the artiodactyl foot, the reduction resulted in the gradual +diminution of the two outer of the four remaining toes, the third +and fourth doing all the work, and thus increasing in size and +power. The fifth digit, for the same reasons as in the +perissodactyl foot, first left the ground and became smaller. Next, +the second soon followed, and these two gradually ceased to be +functional, [and eventually disappeared altogether, as shown in the +accompanying drawing of the feet of still existing animals, Fig. 82 +B, C, D].</p> + +<p>The limb of the modern race-horse is a nearly perfect piece of +machinery, especially adapted to great speed on dry, level ground. +The limb of an antelope, or deer, is likewise well fitted <span class="pagenum"><a name="page_190" id="page_190">[190]</a></span>for +rapid motion on a plain, but the foot itself is adapted to rough +mountain work as well, and it is to this advantage, in part, that +the Artiodactyls owe their present supremacy. The plantigrade +pentadactyl foot of the primitive Ungulate—and even the +perissodactyl foot that succeeded it—both belong to the past humid +period of the world’s history. As the surface of the earth +slowly dried up, in the gradual desiccation still in progress, new +types of feet became a necessity, and the horse, antelope, and +camel were gradually developed, to meet the altered conditions.</p></div> + +<p>The best instance of such progressive modifications in the case of +perissodactyl feet is furnished by the fossil pedigree of the existing +horse, because here, within the limits of the same continuous family +line, we have presented the entire series of modifications.</p> + +<p class="pagenum"><a name="page_189" id="page_189">[189]</a></p> +<div class="figcenter" style="width: 327px;"> +<img src="images/i_206_083.jpg" width="327" height="500" alt="Feet and teeth in fossil pedigree of the Horse." title="" /> +<div class="captionh"><span class="sc">Fig</span>. 83.—Feet and teeth in fossil pedigree of the Horse. +(After Marsh.) <i>a</i>, bones of the fore-foot; <i>b</i>, bones of the hind-foot; +<i>c</i>, radius and ulna; <i>d</i>, tibia and fibula; <i>e</i>, roots of a tooth; <i>f</i> +and <i>g</i>, crowns of upper and lower molar teeth.</div></div> + +<p class="pagenum"><a name="page_190c" id="page_190c">[190c.]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_207_084.jpg" width="500" height="279" alt="Palæotherium." title="" /> +<span class="caption"><span class="sc">Fig</span>. 84.—<span class='sn'>Palæotherium</span>. (Lower Tertiary of Paris Basin.)</span></div> + +<p>There are now known over thirty species of horse-like creatures, +beginning from the size of a fox, then progressively increasing in bulk, +and all standing in <span class="pagenum"><a name="page_191" id="page_191">[191]</a></span> +linear series in structure as in time. Confining attention to the teeth +and feet, it will be seen from the wood-cut on page 189 that the former +grow progressively longer in their sockets, and also more complex in the +patterns of their crowns. On the other hand, the latter exhibit a +gradual diminution of their lateral toes, together with a gradual +strengthening of the middle one. (See Fig. 83.) So that in the +particular case of the horse-ancestry we have a practically complete +chain of what only a few years ago were “missing links.” And +this now practically completed chain shows us the entire history of what +happens to be the most peculiar, or highly specialized, limb in the +whole mammalian class—namely, that of the existing horse. Of the +other two wood-cuts, the former (Fig. 84) shows the skeleton of a very +early and highly generalized ancestor, while the other is a partial +restoration of a much more recent and specialized one. (Fig. 85.)</p> + +<p class="pagenum"><a name="page_192i" id="page_192i">[192i.]</a></p> +<div class="figcenter" style="width: 400px;"> +<img src="images/i_209_085.jpg" width="400" height="324" alt="Hipparion." title="" /> +<span class="caption"><span class="sc">Fig</span>. 85.—<span class='sn'>Hipparion</span>. (New World Pliocene.)</span></div> + +<p><span class="pagenum"><a name="page_191c" id="page_191c">[191c.]</a></span> +On the other hand, progressive modifications of the artiodactyl feet may +be traced geologically up to the different stages presented by living +ruminants, in some of which it has proceeded further than in others. For +instance, if we compare the pig, the deer, and the camel (Fig. 82), we +immediately perceive that the dwindling of the two rudimentary digits +has proceeded much further in the case of the deer than in that of the +pig, and yet not so far as in that of the camel, seeing that here they +have wholly disappeared. Moreover, complementary differences are to be +observed in the degree of consolidation presented by the two useful +digits. For while in the pig the two foot-bones are still clearly +distinguishable <span class="pagenum"><a name="page_192" id="page_192">[192]</a></span> +throughout their entire length, in the deer, and still more in the +camel, their union is more complete, so that they go to constitute a +single bone, whose double or compound character is indicated externally +only by a slight bifurcation at the base. Nevertheless, if we examine +the state of matters in the unborn young of these animals, we find that +the two bones in question are still separated throughout their length. +and thus precisely resemble what used to be their permanent condition in +some of the now fossil species of hoofed mammalia.</p> + +<p>Turning next from bones of the limb to other parts of the mammalian +skeleton, let us briefly consider the evidence of evolution that is here +likewise presented by the vertebral column, the skull, and the teeth.</p> + +<p>As regards the vertebral column, if we examine this <span class="pagenum"><a name="page_193" id="page_193">[193]</a></span> +structure in any of the existing hoofed animals, we find that the bony +processes called zygapophyses, which belong to each of the constituent +vertebræ, are so arranged that the anterior pair belonging to each +vertebra interlocks with the posterior pair belonging to the next +vertebra. In this way the whole series of vertebræ are connected +together in the form of a chain, which, while admitting of considerable +movement laterally, is everywhere guarded against dislocation. But if we +examine the skeletons of any ungulates from the lower Eocene deposits, +we find that in no case is there any such arrangement to secure +interlocking. In all the hoofed mammals of this period the zygapophyses +are flat. Now, from this flat condition to the present condition of full +interlocking we obtain a complete series of connecting links. In the +middle Miocene period we find a group of hoofed animals in which the +articulation begins by a slight rounding of the previously flat +surfaces: later on this rounding progressively increases, until +eventually we get the complete interlocking of the present time.</p> + +<p>As regards teeth, and still confining attention to the hoofed mammals, +we find that low down in the geological series the teeth present on +their grinding surfaces only three simple tubercles. Later on a fourth +tubercle is added, and later still there is developed that complicated +system of ridges and furrows which is characteristic of these teeth at +the present time, and which was produced by manifold and various +involutions of the three or four simple tubercles of Eocene and lower +Miocene times. In other words, the principle of gradual improvement in +the <span class="pagenum"><a name="page_195" id="page_195">[195]</a></span> +Construction of teeth, which has already been depicted as regards the +particular case of the horse-family (Fig. 83), is no less apparent in +the pedigree of all the other mammalia, wherever the palæontological +history is sufficiently intact to serve as a record at all.</p> + +<p class="pagenum"><a name="page_194" id="page_194">[194]</a></p> +<div class="figcenter" style="width: 348px;"> +<img src="images/i_211_086.jpg" width="348" height="500" alt="Comparative series of Brains." title="" /> +<div class="captionh"><span class="sc">Fig</span>. 86.—Comparative series of Brains. (After Le Conte.) +The series reads from above downwards, and represents diagrammatically +the brain of a Fish, a Reptile, a Bird, a Mammal, and a Man. In each +case the letter A marks a side view, and the letter B a top view. The +small italics throughout signify the following homologous parts: <i>m</i>, +medulla; <i>cb</i>, cerebellum; <i>op</i>, optic lobes; <i>cr</i>, cerebrum and +thalamus; <i>ol</i>, olfactory lobes. The series shows a progressive +consolidation and enlargement of the brain in general, and of the +cerebrum and cerebellum in particular, which likewise exhibit +continually advancing structure in respect of convolution. In the case +of Man, these two parts of the brain have grown to so great a size that +they conceal all the other parts from the superficial points of view +represented in the diagram.</div></div> + +<p class="pagenum"><a name="page_195c" id="page_195c">[195c.]</a></p> + +<div class="figcenter" style="width: 400px;"> +<img src="images/i_212_087.jpg" width="400" height="294" alt="Ideal section through all the above stages." title="" /> +<span class="caption"><span class="sc">Fig</span>. 87.—Ideal section through all the above stages. +(After Le Conte.)</span></div> + +<p>Lastly, as regards the skull, casts of the interior show that all the +earlier mammals had small brains with comparatively smooth or +unconvoluted surfaces; and that as time went on the mammalian brain +gradually advanced in size and complexity. Indeed so small were the +cerebral hemispheres of the primitive mammals that they did not overlap +the cerebellum, while their smoothness must have been such as in this +respect to have resembled the brain of a bird or reptile. This, of +course, is just as it ought to be, if the brain, which the skull has to +accommodate, has been gradually <span class="pagenum"><a name="page_196" id="page_196">[196]</a></span> +evolved into larger and larger proportions in respect of its cerebral +hemispheres, or the upper masses of it which constitute the seat of +intelligence. Thus, if we look at the above series of wood-cuts, which +represents the comparative structure of the brain in the existing +classes of the Vertebrata, we can immediately understand why the fossil +skulls of Mammalia should present a gradual increase in size and +furrowing, so as to accommodate the general increase of the brain in +both these respects between the level marked “maml” and that +marked “man,” in the last of the diagrams. (Fig. 87.)</p> + +<p>The tabular statement on the following diagram, which I borrow from +Prof. Cope, will serve at a glance to reveal the combined significance +of so many lines of evidence, united within the limits of the same group +of animals.</p> + +<p>To give only one special illustration of the principle of evolution as +regards the skull, here is one of the most recent instances that has +occurred of the discovery of a missing link, or connecting form (see +Fig. 88). The fossil (B), which was found in New Jersey, stands in an +intermediate position between the stag and the elk. In the stag (A) the +skull is high, showing but little of that anterior attenuation which is +such a distinctive feature of the skull of the elk (C). The nasal bones +(N) of the former, again, are remarkably long when compared with the +similar bones of the latter, and the premaxillaries (PMX), instead of +being projected forward along the horizontal plane of the base of the +skull, are deflected sharply downward. In all these points, it will be +seen, the newly discovered form (<span class='sn'>Cervalces</span>) holds an intermediate +<span class="pagenum"><a name="page_198" id="page_198">[198]</a></span> +position (B). “The skull exhibits a partial attenuation +anteriorly, the premaxillaries are directed about equally downward and +forward, and the nasal bones are measurably contracted in size. The +horns likewise <span class="pagenum"><a name="page_199" id="page_199">[199]</a></span> +furnish characters which further serve to establish this dual +relationship<a name="FNanchor_18_18" id="FNanchor_18_18"></a><a href="#Footnote_18_18" class="fnanchor">[18]</a>.”</p> + +<p class="pagenum"><a name="page_198i" id="page_198i">[198i.]</a></p> +<div class="figcenter" style="width: 307px;"> +<img src="images/i_215_088.jpg" width="307" height="500" alt="Skulls of Canadian Stag, Cervalces +Americanus and Elk." title="" /> +<span class="caption"><span class="sc">Fig.</span> 88.—Skulls of—A, Canadian Stag; B, <span class='sn'>Cervalces +Americanus</span>; and C, Elk. (After Heilprin.)</span></div> + +<p class="pagenum"><a name="page_197" id="page_197">[197]</a></p> +<div class='center mt2'> +<table border="1" cellpadding="4" cellspacing="0" summary="" style='font-size:0.8em;'> +<tr><td class='c'>Formation.</td><td class='c'>No. of toes.</td><td class='c'>Feet.</td><td class='c'>Astragalus.</td><td class='c'>Carpus and tarsus.</td><td class='c'>Ulno-radius</td><td class='c'>Superior molars.</td><td class='c'>Zygapophyses.</td><td class='c'>Brain.</td></tr> +<tr><td class='l'> Pliocene.</td><td class='c'>1-1<br />2-2</td><td class='l'>Digitigrade.<br />(Plantigrade.)</td><td class='l'>Grooved.<br />(Flat.)</td><td class='l'>Interlocking.<br />(Opposite.)</td><td class='l'>Faceted.</td><td class='l'>4-tubercles, crested and cemented</td><td class='l'>Doubly involute.<br />Singly involute.</td><td class='l'>Hemispheres larger, convoluted.</td></tr> +<tr><td class='l'>Upper Miocene.<br /> (Loup Fork.)</td><td class='c'>3-3<br />4-4<br />(5-5)</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr> +<tr><td class='l'>Middle.<br />(John Day.)</td><td class='c'>2-2<br />3-3<br />4-4</td><td class='l'>Digitigrade.</td><td class='l'>Grooved.</td><td class='l'>Interlocking.</td><td class='l'>Faceted.<br />Smooth.</td><td class='l'>4-tubercles, and crested.</td><td class='l'>Singly involute.<br />Doubly involute.</td><td class='l'>Hemispheres larger, convoluted.</td></tr> +<tr><td class='l'>Lower<br />(White River.)</td><td class='c'>3-3<br />4-3<br />4-4</td><td class='l'>Digitigrade.<br />Plantigrade.</td><td class='l'>Grooved.</td><td class='l'>Interlocking.</td><td class='l'>Smooth.<br />Faceted.</td><td class='l'>4-tubercles, and crested.</td><td class='l'>? Singly involute.</td><td class='l'>Hemispheres small, and larger.</td></tr> +<tr><td class='l'>Eocene.<br />Upper<br />(bridger.)</td><td class='c'>3-3<br />4-3<br />4-5<br />5-5</td><td class='l'>(Digitigrade.)<br />Plantigrade.</td><td class='l'>Grooved.<br />(Flat.)</td><td class='l'>Opposite. Interlocking.</td><td class='l'>Smooth.</td><td class='l'>4-tubercles.<br />3-tubercles, and crested.</td><td class='l'>Singly involute.<br />Plane.</td><td class='l'>Hemispheres small</td></tr> +<tr><td class='l'>Middle.<br />(Wasatch.)</td><td class='c'>4-3<br />4-5<br />5-5</td><td class='l'>Plantigrade.<br />(Digitigrade.)</td><td class='l'>Flat.<br />(Grooved.)</td><td class='l'>Opposite. Interlocking.</td><td class='l'>Smooth.</td><td class='l'>4-tubercles.<br />3-tubercles, a few crested.</td><td class='l'>Plane.<br />Singly involute.</td><td class='l'>Hemispheres small; mesencehpalon sometimes exposed.</td></tr> +<tr><td class='l'>Lower<br />(Puerco.)</td><td class='c'>5-5</td><td class='l'>Plantigrade.</td><td class='l'>Flat.</td><td class='l'>Opposite.</td><td class='l'>Smooth.</td><td class='l'>3-tubercles.<br />(4-tubercles), none crested.</td><td class='l'>Plane.</td><td class='l'>Mesencephalon exposed; hemisphere small and smoother.</td></tr> +</table></div> + +<p><span class="pagenum"><a name="page_199c" id="page_199c">[199c.]</a></span> +The evidence, then, which is furnished by all parts of the vertebral +skeleton—whether we have regard to Fishes, Reptiles, Birds, or +Mammals—is cumulative and consistent. Nowhere do we meet with any +deviation or ambiguity, while everywhere we encounter similar proofs of +continuous transformation—proofs which vary only with the varying +amount of material which happens to be at our disposal, being most +numerous and detailed in those cases where the greatest number of fossil +forms has been preserved by the geological record. Here, therefore, we +may leave the vertebral skeleton; and, having presented a sample of the +evidence as yielded by horns and bones, I will conclude by glancing with +similar brevity at the case of shells—which, as before remarked, +constitute the only other sufficiently hard or permanent material to +yield unbroken evidence touching the fossil ancestry of animals.</p> + +<p>Of course it will be understood that I am everywhere giving merely +samples of the now superabundant evidence which is yielded by +palæontology; and, as this chapter is already a long one, I must content +myself with citing only the case of mollusk-shells, although shells of +other classes might be made to yield highly important additions to the +testimony. Moreover, even as regards the one division of mollusk-shells, +I can afford to quote only a very few cases. These, however, are in my +opinion the strongest single pieces of evidence in favour of +transmutation which have thus far been brought to light.</p> + +<p class="pagenum"><a name="page_200" id="page_200">[200]</a></p> +<div class="figcenter" style="width: 293px;"> +<img src="images/i_217_089.jpg" width="293" height="500" alt="Transmutations of Planorbis." title="" /> +<span class="caption"><span class="sc">Fig.</span> 89.—Transmutations of <span class='sn'>Planorbis</span>. (After Hyatt.)</span></div> + +<p>Near the village of Steinheim, in Würtemberg, there is an ancient +lake-basin, dating from Tertiary times. The lake has long ago dried up; +but its aqueous deposits are extraordinarily rich in fossil shells, +especially of different species of the genus<span class="pagenum"><a name="page_201" id="page_201">[201]</a></span> +<span class='sn'>Planorbis</span>. The following is an authoritative +<i>résumé</i> of the facts.</p> + +<div class='blockquot'><p>As the deposits seem to have been continuous for ages, and the +fossil shells very abundant, this seemed to be an excellent +opportunity to test the theory of derivation. With this end in +view, they have been made the subject of exhaustive study by +Hilgendorf in 1866, and by Hyatt in 1880. In passing from the +lowest to the highest strata the species change greatly and many +times, the extreme forms being so different that, were it not for +the intermediate forms, they would be called not only different +species, but different genera. And yet the gradations are so +insensible that the whole series is nothing less than a +demonstration, in this case at least, of origin of species by +derivation with modifications. The accompanying plate of successive +forms (Fig. 89), which we take from Prof. Hyatt’s admirable +memoir, will show this better than any mere verbal explanation. It +will be observed that, commencing with four slight +varieties—probably sexually isolated varieties—of one species, +each series shows a gradual transformation as we go upward in the +strata—i. e. onward in time. Series I branches into three +sub-series, in two of which the change of form is extreme. Series +IV is remarkable for great increase in size as well as change in +form. In the plate we give only selected stages, but in the fuller +plates of the memoir, and still more in the shells themselves, the +subtilest gradations are found<a name="FNanchor_19_19" id="FNanchor_19_19"></a><a href="#Footnote_19_19" class="fnanchor">[19]</a>.</p></div> + +<p>Here is another and more recently observed case of transmutation in the +case of mollusks.</p> + +<p>The recent species, <span class='sn'>Strombus accipitrinus</span>, still inhabits the coasts of +Florida. Its extinct prototype, <span class='sn'>S. Leidy</span>, was discovered a few years ago +by Prof. Heilprin in the Pliocene formations of the interior of Florida. +The peculiar shape of the wing, and tuberculation of the whorl, are thus +proved to have grown but of a previously more conical form of shell.</p> + +<p class="pagenum"><a name="page_202" id="page_202">[202]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_219_090.jpg" width="500" height="217" alt="Transformation of Strombus." title="" /> +<span class="caption"><span class="sc">Fig.</span> 90.—Transformation of <span class='sn'>Strombus</span>. (After Heilprin.) +1, 1<i>a</i>, <span class='sn'>Strombus Leidy</span> (1, typical), Pliocene; 2, 2<i>a</i>, <span class='sn'>Strombus +accipitrinus</span> (2<i>a</i> typical) Recent.</span></div> + +<p><span class="pagenum"><a name="page_203" id="page_203">[203]</a></span></p> +<p>Lastly, attention may here again be directed to the very instructive +series of shells which has already been shown in a previous chapter, and +which serves to illustrate the successive geological forms of <span class='sn'>Paludina</span> +from the Tertiary beds of Slavonia, as depicted by Prof. Neumayr of +Vienna. (Fig. 1, p. 19.)</p> + +<hr /><p class="pagenum"><a name="page_204" id="page_204">[204]</a></p> +<h2><a name="CHAPTER_VI" id="CHAPTER_VI"></a>CHAPTER VI.<br /><br /> +<span class="sc">Geographical Distribution.</span></h2> + +<p>The argument from geology is the argument from the distribution of +species in time. I will next take the argument from the distribution of +species in space—that is, the present geographical distribution of +plants and animals.</p> + +<p>Seeing that the theory of descent with adaptive modification implies +slow and gradual change of one species into another, and progressively +still more slow and gradual changes of one genus, family, or order into +another genus, family, or order, we should expect on this theory that +the organic types living on any given geographical area would be found +to resemble or to differ from organic types living elsewhere, according +as the area is connected with or disconnected from other geographical +areas. For instance, the large continental islands of Australia and New +Zealand are widely disconnected from all other lands of the world, and +deep sea soundings show that they have probably been thus disconnected, +either since the time of their origin, or, at the least, through immense +geological epochs. The theory of evolution, therefore, would expect to +find two general facts with regard to the inhabitants of these islands. +First, that <span class="pagenum"><a name="page_205" id="page_205">[205]</a></span> +the inhabitants should form, as it were, little worlds of their own, +more or less unlike the inhabitants of any other parts of the globe. And +next, that some of these inhabitants should present us with independent +information touching archaic forms of life. For it is manifestly most +improbable that the course of evolutionary history should have run +exactly parallel in the case of these isolated oceanic continents and in +continents elsewhere. Australia and New Zealand, therefore, ought to +present a very large number, not only of peculiar species and genera, +but even of families, and possibly of orders. Now this is just what +Australia and New Zealand do present. The case of the dog being +doubtful, there is an absence of all mammalian life, except that of one +of the oldest and least highly developed orders, the Marsupials. There +even occurs a unique order, still lower in the scale of +organization—so low, in fact, that it deserves to be regarded as +but nascent mammalian: I mean, of course, the Monotremata. As regards +Birds, we have the peculiar wingless forms alluded to in a previous +chapter (viz. that on Morphology); and, without waiting to go into +details, it is notorious that the faunas of Australia and New Zealand +are not only highly peculiar, but also suggestively archaic. Therefore, +in both the respects above mentioned, the anticipations of our theory +are fully borne out. But as it would take too long to consider, even +cursorily, the faunas and floras of these immense islands, I here allude +to them only for the sake of illustration. In order to present the +argument from geographical distribution within reasonable limits, I +think it is best to restrict our examination to smaller areas; for these +will better <span class="pagenum"><a name="page_206" id="page_206">[206]</a></span> +admit of brief and yet adequate consideration. But of course it will be +understood that the less isolated the region, and the shorter the time +that it has been isolated, the smaller amount of peculiarity should we +expect to meet with on the part of its present inhabitants. Or, +conversely stated, the longer and the greater the isolation, the more +peculiarity of species would our theory expect to find. The object of +the present chapter will be to show that these, and other cognate +expectations, are fully realized by facts; but, before proceeding to do +this, I must say a few words on the antecedent standing of the argument.</p> + +<p>Where the question is, as at present, between the rival theories of +special creation and gradual transmutation, it may at first sight well +appear that no test can be at once so crucial and so easily applied as +this of comparing the species of one geographical area with those of +another, in order to see whether there is any constant correlation +between differences of type and degrees of separation. But a little +further thought is enough to show that the test is not quite so simple +or so absolute—that it is a test to be applied in a large and general +way over the surface of the whole earth, rather than one to be relied +upon as exclusively rigid in every special case.</p> + +<p>In the first place, there is the obvious consideration that lands or +seas which are discontinuous now may not always have been so, or not for +long enough to admit of the effects of separation having been exerted to +any considerable extent upon their inhabitants. Next, there is the +scarcely less important consideration, that although land areas may long +have been separated from one another by extensive tracts of ocean, +<span class="pagenum"><a name="page_207" id="page_207">[207]</a></span> +birds and insects may more or less easily have been able to fly from +one to the other; while even non-flying animals and plants may often +have been transported by floating ice or timber, wind or water currents, +and sundry other means of dispersal. Again, there is the important +influence of climate to be taken into account. We know from geological +evidence that in the course of geological time the self-same continents +have been submitted to enormous changes of temperature—varying in +fact from polar cold to almost tropical heat; and as it is manifestly +impossible that forms of life suited to one of these climates could have +survived during the other, we can here perceive a further and most +potent cause interfering with the test of geographical distribution as +indiscriminately applied in all cases. When the elephant and +hippopotamus were flourishing in England amid the luxuriant vegetation +which these large animals require, it is evident that scarcely any one +species of either the fauna or the flora of this country can have been +the same as it was when its African climate gave place to that of +Greenland. Therefore, as Mr. Wallace observes, “If glacial epochs +in temperate lands and mild climates near the poles have, as now +believed by men of eminence, occurred several times over in the past +history of the earth, the effects of such great and repeated changes +both on migration, modification, and extinction of species, must have +been of overwhelming importance—of more importance perhaps than +even the geological changes of sea and land.”</p> + +<p>But although for these, and certain other less important reasons which I +need not wait to detail, we must conclude that the evidence from +geographical <span class="pagenum"><a name="page_208" id="page_208">[208]</a></span> +distribution is not to be regarded as a crucial test between the rival +theories of creation and evolution in all cases indiscriminately, I must +next remark that it is undoubtedly one of the strongest lines of +evidence which we possess. When we once remember that, according to the +general theory of evolution itself, the present geographical +distribution of plants and animals is “the visible outcome or +residual product of the whole past history of the earth,” and, +therefore, that of the conditions determining the characters of life +inhabiting this and that particular area continuity or discontinuity +with other areas is but one,—when we remember this, we find that +no further reservation has to be made: all the facts of geographical +distribution speak with one consent in favour of the naturalistic +theory.</p> + +<hr class='minor' /> + +<p>The first of these facts which I shall adduce is, that although the +geographical range of any given species is, as a rule, continuous, such +is far from being always the case. Very many species have more or less +discontinuous ranges—the mountain-hare, for instance, extending from +the Arctic regions over the greater portion of Europe to the Ural +Mountains and the Caucasus, and yet over all this enormous tract +appearing only in isolated or discontinuous patches, where there happen +to be either mountain ranges or climates cold enough to suit its nature. +Now, in all such cases of discontinuity in the range of a species the +theory of evolution has a simple explanation to offer—namely, either +that some representatives of the species have at some former period been +able to migrate from one region to the other, or else that at <span class="pagenum"><a name="page_209" id="page_209">[209]</a></span> +one time the species occupied the whole of the range in question, but +afterwards became broken up as geographical, climatic, or other changes +rendered parts of the area unfit for the species to inhabit. Thus, for +instance, it is easy to understand that during the last cold epoch the +mountain-hare would have had a continuous range; but that as the Arctic +climate gradually receded to polar regions, the species would be able to +survive in southern latitudes only on mountain ranges, and thus would +become broken up into many discontinuous patches, corresponding with +these ranges. In the same way we can explain the occurrence of Arctic +vegetation on the Alps and Pyrenees—namely, as left behind by the +retreat of the Arctic climate at the close of the glacial period.</p> + +<p>But now, on the other hand, the theory of special creation cannot so +well afford to render this obvious explanation of discontinuity. In the +case of the Arctic flora of the Alps, for instance, although it is true +that much of this vegetation is of an Arctic type, it is not true that +the species are all identical with those which occur in the Arctic +regions. Therefore the theory of special creation would here have to +assume that, although the now common species were left behind on the +Alps by the retreat of glaciation northwards, the peculiar Alpine +species were afterwards created separately upon the Alps, and yet +created with such close affinities to the pre-existing species as to be +included with them under the same genera. Looking to the absurdity of +this supposition, as well as of others which I need not wait to mention, +certain advocates of special creation have sought to take refuge in +another hypothesis—namely, that <span class="pagenum"><a name="page_210" id="page_210">[210]</a></span> +species which present a markedly discontinuous range may have had a +corresponding number of different centres of creation, the same specific +type having been turned down, so to speak, on widely separated areas. +But to me it seems that this explanation presents even greater +difficulty than the other. If it is difficult to say why the Divinity +should have chosen to create new species of plants on the Alps on so +precisely the same pattern as the old, much more would it be difficult +to say why, in addition to these new species, he should also have +created again the old species which he had already placed in the Arctic +regions.</p> + +<hr class='minor' /> + +<p>So much, then, for discontinuity of distribution. The next general fact +to be adduced is, that there is no constant correlation between habitats +and animals or plants suited to live upon them. Of course all the +animals and plants living upon any given area are well suited to live +upon that area; for otherwise they could not be there. But the point now +is, that besides the area on which they do live, there are usually many +other areas in different parts of the globe where they might have lived +equally well—as is proved by the fact that when transported by man they +thrive as well, or even better, than in their native country. Therefore, +upon the supposition that all species were separately created in the +countries where they are respectively found, we must conclude that they +were created in only some of the places where they might equally well +have lived. Probably there is at most but a small percentage either of +plants or animals which would not thrive in some <span class="pagenum"><a name="page_211" id="page_211">[211]</a></span> +place, or places, on the earth’s surface other than that in which +they occur; and hence we must say that one of the objects of special +creation—if this be the true theory—was that of depositing +species in only some among the several parts of the earth’s +surface equally well suited to support them.</p> + +<p>Now, I do not contend that this fact in itself raises any difficulty +against the theory of special creation. But I do think that a very +serious difficulty is raised when to this fact we add another—namely, +that on every biological region we encounter species related to other +species in genera, and usually also genera related to other genera in +families. For if each of all the constituent species of a genus, and +even of a family, were separately created, we must hence conclude that +in depositing them there was an unaccountable design manifested to make +areas of distribution correspond to the natural affinities of their +inhabitants. For example, the humming-birds are geographically +restricted to America, and number 120 genera, comprising over 400 +species. Hence, if this betokens 400 separate acts of creation, it +cannot possibly have been due to chance that they were all performed on +the same continent: it must have been design which led to every species +of this large family of birds having been deposited in one geographical +area. Or, to take a case where only the species of a single genus are +concerned. The rats and mice proper constitute a genus which comprises +altogether more than 100 species, and they are all exclusively +restricted to the Old World. In the New World they are represented by +another genus comprising about 70 species, which resemble their<span class="pagenum"><a name="page_212" id="page_212">[212]</a></span> +Old World cousins in form and habits; but differ from them in dentition +and other such minor points. Now, the question is,—Why should all +the 100 species have been separately created on one side of the Atlantic +with one pattern of dentition, and all the 70 species on the other side +with another pattern? What has the Atlantic Ocean got to do with any +“archetypal plan” of rats’ teeth?</p> + +<p>Or again, to recur to Australia, why should all the mammalian forms of +life be restricted to the one group of Marsupials, when we know that not +only the Rodents, such as the rabbit, but all other orders of mammals, +would thrive there equally well. And similarly, of course, in countless +other instances. Everywhere we meet with this same correlation between +areas of distribution and affinities of classification.</p> + +<p>Now, it is at once manifest how completely this general fact harmonizes +with the theory of evolution. If the 400 species of humming-birds, for +instance, are all modified descendants of common ancestors, and if none +of their constituent individuals have ever been large enough to make +their way across the oceans which practically isolate their territory +from all other tropical and sub-tropical regions of the globe, then we +can understand why it is that all the 400 species occupy the same +continent. But on the special-creation theory we can see no reason why +the 400 species should all have been deposited in America. And, as +already observed, we must remember that this correlation between a +geographically restricted habitat and the zoological or botanical +affinities of its inhabitants, is repeated over and over and over again +in the <span class="pagenum"><a name="page_213" id="page_213">[213]</a></span> +faunas and floras of the world, so that merely to enumerate the +instances would require a separate chapter.</p> + +<p>Furthermore, the general argument thus presented in favour of descent +with continuous modification admits of being enormously strengthened by +three different classes of additional facts.</p> + +<p>The first is, that the correlation in question—namely, that between a +geographically restricted habitat and the zoological or botanical +affinities of its inhabitants—is not limited to the now existing +species, but extends also to the extinct. That is to say, the dead +species are allied to the living species, as we should expect that they +must be, if the latter are modified descendants of the former. On the +alternative theory, however, we have to suppose that the policy of +maintaining a correlation between geographical restriction and natural +affinity extends very much further back than even the existing species +of plants and animals; indeed we must suppose that a practically +infinite number of additional acts of separate creation were governed by +the same policy, in the case of long lines of species long since +extinct.</p> + +<p>Thus far, then, the only answer which an advocate of special creation +can adduce is, that for some reason unknown to us such a policy may have +been more wise than it appears: it may have served some inscrutable +purpose that allied products of distinct acts of creation should all be +kept together on the same areas. Well, in answer to this unjustifiable +appeal to the argument from ignorance, I will adduce the second of the +three considerations. This is, that in cases where the geographical +areas are not restricted the <span class="pagenum"><a name="page_214" id="page_214">[214]</a></span> +policy in question fails. In other words, where the inhabitants of an +area are free to migrate to other areas, the policy of correlating +affinity with distribution is most significantly forgotten. In this case +species wander away from their native homes, and the course of their +wanderings is marked by the origination of new species springing up en +route. Now, is it reasonable to suppose that the mere circumstance of +some members of a species being able to leave their native home should +furnish any occasion for creating new and allied species upon the tracts +over which they travel, or the territories to which they go? When the +400 existing species of humming-birds have all been created on the same +continent for some reason supposed to be unknown, why should this reason +give way before the accident of any means of migration being furnished +to humming-birds, so that they should be able to visit, say the +continents of Africa and Asia, there gain a footing beside the +sun-birds, and henceforth determine a new centre for the separate +creation of additional species of humming-birds peculiar to the Old +World—as has happened in the case of the majority of species +which, unlike the humming-birds, have been at any time free to migrate +from their original homes?</p> + +<p>Lastly, my third consideration is, that the supposed policy in question +does not extend to affinities which are wider than those between species +and genera—more rarely to families, scarcely ever to orders, and never +to classes. In other words, nature shows a double correlation in her +geographical distribution of organic types:—first, that which we have +already considered between geographical restriction and <span class="pagenum"><a name="page_215" id="page_215">[215]</a></span> +Natural affinity among inhabitants of the same areas; second, another +of a more detailed character between <i>degrees</i> of geographical +restriction and <i>degrees</i> of natural affinity. The more distant the +affinity, the more general is the extension. This, of course, is what we +should expect on the theory of descent with modification, because the +more distant the affinity, and therefore, <i>ex hypothesi</i>, the larger and +the older the original group of organisms, the greater must be the +chance of dispersal. The 400 species of humming-birds may well be unable +to migrate from their native continent; but it would indeed have been an +unaccountable fact if no other species of all the class of birds had +ever been able to have crossed the atlantic ocean. Thus, on the theory +of evolution, we can well understand the second correlation now before +us—namely, between remoteness of affinity and generality of +dispersal,—so that there is no considerable portion of the +habitable globe without representatives of all the classes of animals, +few portions without representatives of all the orders, but many +portions without many of the families, innumerable portions without +innumerable genera, and, of course, all portions without the great +majority of species. Now, while this general correlation thus obviously +supports the theory of natural descent with progressive modification, it +makes directly against the opposite theory of special creation. For we +have recently seen that when we restrict our view to the case of species +and genera, the theory of special creation is obliged to suppose that +for some inscrutable reason the deity had regard to systematic affinity +while determining on what large areas to <span class="pagenum"><a name="page_216" id="page_216">[216]</a></span> +create his species<a name="FNanchor_20_20" id="FNanchor_20_20"></a><a href="#Footnote_20_20" class="fnanchor">[20]</a>. +but now we see that he must be held to have neglected this inscrutable +reason (whatever it was) when he passed beyond the range of +genera—and this always in proportion to the remoteness of +systematic affinity on the part of the species concerned.</p> + +<p>I cannot well conceive a <i>reductio ad absurdum</i> more complete than this. +But, having now presented these most general facts of geographical +distribution in their relation to the issue before us, we may next +proceed to consider a few illustrations of them in detail, for in this +way I think that their overwhelming weight may become yet more +abundantly apparent.</p> + +<hr class='minor' /> + +<p>It will assist us in dealing with these detailed illustrations if we +begin by considering the means of dispersal of organisms from one place +to another. Of course the most ordinary means is that of continuous +wandering, or emigration; but where geographical barriers of any kind +have to be surmounted, organisms may only be able to pass them by more +exceptional and accidental means. The principal barriers of a +geographical kind are oceans, rivers, mountain-chains, and +desert-tracts, in the case of <span class="pagenum"><a name="page_217" id="page_217">[217]</a></span> +terrestrial organisms; and, in the case of aquatic organisms, the +presence of land. But it is to be observed that, as regards marine +organisms, any considerable difference in the temperature of the water +may constitute a barrier as effectual as the presence of land; and also +that, in the case of all shallow-water faunas, a tract of deep ocean +constitutes almost as complete a barrier as it does to terrestrial +faunas.</p> + +<p>Now, the means whereby barriers admit of being accidentally or +occasionally surmounted are, of course, various; and they differ in the +case of different organisms. Birds, bats, and insects, on account of +their powers of flight, are particularly apt to be blown out great +distances to sea, and hence of all animals are most likely to become the +involuntary colonists of distant shores. Floating timber serves to +convey seeds and eggs of small animals over great distances; and Darwin +has shown that many kinds of seeds are able of themselves to float for +more than a month in sea-water without losing their powers of +germination. For instance, out of 87 kinds, 64 germinated after an +immersion of 28 days, and a few survived an immersion of 137 days. As a +result of all his experiments he concludes, that the seeds of at least +ten per cent. of the species of plants of any country might be floated +by sea-currents during 28 days, without losing their powers of +germination; and this, at the average rate of flow of several Atlantic +currents, would serve to transport the seeds to a distance of at least +900 miles. Again, he proved that even seeds which are quickly destroyed +by contact with sea-water admit of being successfully transported during +30 days, if they be contained within the crop of a dead bird. He also +<span class="pagenum"><a name="page_218" id="page_218">[218]</a></span> +proved that living birds are most active agents in the work of +dissemination, and this not only by taking seeds into their crops +(where, so long as they remain, the seeds are uninjured), but likewise +by carrying seeds (and even young mollusks) attached to their feet and +feathers. In the course of these experiments he found that a small +cup-full of mud, which he gathered from the edges of three ponds in +February, was so charged with seeds that when sown in the ground these +few ounces of mud yielded no less than 537 plants, belonging to many +different species. It is therefore evident what opportunities are thus +afforded for the transportation of seeds on the feet and bills of +wading-birds. Lastly, floating ice is well known to act as a carrier of +any kind of life which may prove able to survive this mode of transit.</p> + +<p>Such being the nature of geographical barriers, and the means that +organisms of various kinds may occasionally have of overcoming them, I +will now give a few detailed illustrations of the argument from +geographical distribution, as previously presented in its general form.</p> + +<p>To begin with aquatic animals. As Darwin remarks, “the marine +inhabitants of the Eastern and Western shores of South America are very +distinct; with extremely few shells, crustacea, or echinodermata in +common.” Again, westward of the shores of America, a wide space of +open ocean extends, which, as we have seen, furnishes as effectual a +barrier as does the land to any emigration of shallow-water animals. +Now, as soon as this reach of deep water is passed, we meet in the +eastern islands of the Pacific with another and totally distinct fauna. +“So that three marine <span class="pagenum"><a name="page_219" id="page_219">[219]</a></span> +faunas range northward and southward in parallel lines not far from +each other, under corresponding climates": they are, however, +“separated from each other by impassable barriers, either of land +or open sea": and it is in exact coincidence with the course of these +barriers that we find so remarkable a differentiation of the +faunas<a name="FNanchor_21_21" id="FNanchor_21_21"></a><a href="#Footnote_21_21" class="fnanchor">[21]</a>. +Obviously, therefore, it is impossible to suggest that this correlation +is accidental. Altogether many thousands of species are involved, and +within this comparatively limited area they are sharply marked off into +three groups as to their natural affinities, and into three groups as to +their several basins. Hence, if all these species were separately +created, there is no escape from the conclusion that for some reason or +another the act of creation was governed by the presence of these +barriers, so that species deposited on the Eastern shores of South +America were formed with one set of natural affinities, while species +deposited on the Western shore were formed with another set; and +similarly with regard to the third set of species in the third basin, +which, extending over a whole hemisphere to the coast of Africa without +any further barrier, nowhere presents, over this vast area, any other +case of a distinct marine fauna. But what conceivable reason can there +have been thus to consult these geographical barriers in the original +creation of specific <span class="pagenum"><a name="page_220" id="page_220">[220]</a></span> +types? Even if such a case stood alone, it would be strongly suggestive +of error on the part of the special creation theory. But let us take +another case, this time from fresh-water faunas.</p> + +<p>Although the geographical distribution of fresh-water fish and +fresh-water shells is often surprisingly extensive and apparently +capricious, this may be explained by the means of dispersal being here +so varied—not only aquatic birds, floods, and whirlwinds, but also +geographical changes of water-shed having all assisted in the process. +Moreover, in some cases it is possible that the habits of more widely +distributed fresh-water fish may have originally been wholly or partly +marine—which, of course, would explain the existing discontinuity of +their existing fresh-water distribution. But, be this as it may (and it +is not a question that affects the issue between special creation and +gradual evolution, since it is only a question as to how a given species +has been dispersed from its original home, whether or not in that home +it was specially created), the point I desire to bring forward is, that +where we find a barrier to the emigration of fresh-water forms which is +more formidable than a thousand miles of ocean—a barrier over which +neither water-fowl nor whirlwinds are likely to pass, and which is above +the reach of any geological changes of water-shed,—where we find such a +barrier, we always find a marked difference in the fresh-water faunas on +either side of it. The kind of barrier to which I allude is a high +mountain-chain. It may be only a few miles wide; yet it exercises a +greater influence on the diversification of specific <span class="pagenum"><a name="page_221" id="page_221">[221]</a></span> +types, where fresh-water faunas are concerned, than almost any other. +But why should this be the case on any intelligible theory of special +creation? Why, in the depositing of species of newly created fresh-water +fish, should the presence of an impassable mountain-chain have +determined so uniformly a difference of specific affinity on either side +of it? The question, so far as I can see, does not admit of an answer +from any reasonable opponent.</p> + +<hr class='minor' /> + +<p>Turning now from aquatic organisms to terrestrial, the body of facts +from which to draw is so large, that I think the space at my disposal +may be best utilized by confining attention to a single division of +them—that, namely, which is furnished by the zoological study of +oceanic islands.</p> + +<p>In the comparatively limited—but in itself extensive—class of facts +thus presented, we have a particularly fair and cogent test as between +the alternative theories of evolution and creation. For where we meet +with a volcanic island, hundreds of miles from any other land, and +rising abruptly from an ocean of enormous depth, we may be quite sure +that such an island can never have formed part of a now submerged +continent. In other words, we may be quite sure that it always has been +what it now is—an oceanic peak, separated from all other land by +hundreds of miles of sea, and therefore an area supplied by nature for +the purpose, as it were, of testing the rival theories of creation and +evolution. For, let us ask, upon these tiny insular specks of land what +kind of life should we expect to find? To this question the theories of +special creation and of gradual evolution would <span class="pagenum"><a name="page_222" id="page_222">[222]</a></span> +agree in giving the same answer up to a certain point. For both +theories would agree in supposing that these islands would, at all +events in large part, derive their inhabitants from accidental or +occasional arrivals of wind-blown or water-floated organisms from other +countries—especially, of course, from the countries least remote. +But, after agreeing upon this point, the two theories must part company +in their anticipations. The special-creation theory can have no reason +to suppose that a small volcanic island in the midst of a great ocean +should be chosen as the theatre of any extraordinary creative activity, +or for any particularly rich manufacture of peculiar species to be found +nowhere else in the world. On the other hand, the evolution theory would +expect to find that such habitats are stocked with more or less peculiar +species. For it would expect that when any organisms chanced to reach a +wholly isolated refuge of this kind, their descendants should forthwith +have started upon an independent course of evolutionary history. +Protected from intercrossing with any members of their parent species +elsewhere, and exposed to considerable changes in their conditions of +life, it would indeed be fatal to the general theory of evolution if +these descendants, during the course of many generations, were not to +undergo appreciable change. It has happened on two or three occasions +that European rats have been accidentally imported by ships upon some of +these islands, and even already it is observed that their descendants +have undergone a slight change of appearance, so as to constitute them +what naturalists call local varieties. The change, of course, is but +<span class="pagenum"><a name="page_223" id="page_223">[223]</a></span> +slight, because the time allowed for it has been so short. But the +longer the time that a colony of a species is thus completely isolated +under changed conditions of life the greater, according to the evolution +theory, should we expect the change to become. Therefore, in all cases +where we happen to know, from independent evidence of a geological kind, +that an oceanic island is of very ancient formation, the evolution +theory would expect to encounter a great wealth of peculiar species. On +the other hand, as I have just observed, the special-creation theory can +have no reason to suppose that there should be any correlation between +the age of an oceanic island and the number of peculiar species which it +may be found to contain.</p> + +<p>Therefore, having considered the principles of geographical distribution +from the widest or most general point of view, we shall pass to the +opposite extreme, and consider exhaustively, or in the utmost possible +detail, the facts of such distribution where the conditions are best +suited to this purpose—that is, as I have already said, upon oceanic +islands, which may be metaphorically regarded as having been formed by +nature for the particular purpose of supplying naturalists with a +crucial test between the theories of creation and evolution. The +material upon which my analysis is to be based will be derived from the +most recent works upon geographical distribution—especially from the +magnificent contributions to this department of science which we owe to +the labours of Mr. Wallace. Indeed, all that follows may be regarded as +a condensed filtrate of the facts which he has collected. Even as thus +restricted, however, our subject-matter <span class="pagenum"><a name="page_224" id="page_224">[224]</a></span> +would be too extensive to be dealt with on the present occasion, were +we to attempt an exhaustive analysis of the floras and faunas of all +oceanic islands upon the face of the globe. Therefore, what I propose to +do is to select for such exhaustive analysis a few of what may be termed +the most oceanic of oceanic islands—that is to say, those oceanic +islands which are most widely separated from mainlands, and which, +therefore, furnish the most unquestionable of test cases as between the +theories of special creation and genetic descent.</p> + +<hr class='minor' /> + +<p><i>Azores.</i>—A group of volcanic islands, nine in number, about 900 miles +from the coast of Portugal, and surrounded by ocean depths of 1,800 to +2,500 fathoms. There is geological evidence that the origin of the group +dates back at least as far as Miocene times. There is a total absence of +all terrestrial Vertebrata, other than those which are known to have +been introduced by man. Flying animals, on the other hand, are abundant; +namely, 53 species of birds, one species of bat, a few species of +butterflies, moths, and hymenoptera, with 74 species of indigenous +beetles. All these animals are unmodified European species, with the +exception of one bird and many of the beetles. Of the 74 indigenous +species of the latter, 36 are not found in Europe; but 19 are natives of +Madeira or the Canaries, and 3 are American, doubtless transplanted by +drift-wood. The remaining 14 species occur nowhere else in the world, +though for the most part they are allied to other European species. +There are 69 known species of land-shells, of which 37 are European, and +32 peculiar, though all allied to European <span class="pagenum"><a name="page_225" id="page_225">[225]</a></span> +forms. Lastly, there are 480 known species of plants, of which 40 are +peculiar, though allied to European species.</p> + +<p><i>Bermudas.</i>—A small volcanic group of islands, 700 miles from North +Carolina. Although there are about 100 islands in the group, their total +area does not exceed 50 square miles. The group is surrounded by water +varying in depth from 2,500 to 3,800 fathoms. The only terrestrial +Vertebrate (unless the rats and mice are indigenous) is a lizard allied +to an American form, but specifically distinct from it, and therefore a +solitary species which does not occur anywhere else in the world. None +of the birds or bats are peculiar, any more than in the case of the +Azores; but, as in that case, a large percentage of the land-shells are +so—namely, at least one quarter of the whole. Neither the botany nor +the entomology of this group has been worked out; but I have said enough +to show how remarkably parallel are the cases of these two volcanic +groups of islands situated in different hemispheres, but at about the +same distance from large continents. In both there is an extraordinary +paucity of terrestrial vertebrata, and of any peculiar species of bird +or beast. On the other hand, there is in both a marvellous wealth of +peculiar species of insects and land-shells. Now these correlations are +all abundantly intelligible. It is a difficult matter for any +terrestrial animal to cross 900, or even 700, miles of ocean: therefore +only one lizard has succeeded in doing so in one of the two parallel +cases; and, living cut off from intercrossing with its parent form, the +descendants of that lizard have become modified so as to constitute a +peculiar species. But it is more easy for large flying animals <span class="pagenum"><a name="page_226" id="page_226">[226]</a></span> +to cross those distances of ocean: consequently, there is only one +instance of a peculiar species of bird or bat—namely, a bull-finch +in the Azores, which, being a small land-bird, is not likely ever to +have had any other visitors from its original parent species coming over +from Europe to keep up the original breed. Lastly, it is very much more +easy for insects and land-mollusca to be conveyed to such islands by +wind and floating timber than it is for terrestrial mammals, or even +than it is for small birds and bats; but yet such means of transit are +not sufficiently sure to admit of much recruiting from the mainland for +the purpose of keeping up the specific types. Consequently, the insects +and the land-shells present a much greater proportion of peculiar +species—namely, one half and one fourth of the land-shells in the +one case, and one eighth of the beetles in the other. All these +correlations, I say, are abundantly intelligible on the theory of +evolution; but who shall explain, on the opposite theory, why orders of +beetles and land-mollusca should have been chosen from among all other +animals for such superabundant creation on oceanic islands, so that in +the Azores alone we find no less than 32 of the one and 14 of the other? +And, in this connexion, I may again allude to the peculiar species of +beetles in the island of Madeira. Here there are an enormous number of +peculiar species, though they are nearly all related to, or included +under the same genera as, beetles on the neighbouring continent. Now, as +we have previously seen, no less than 200 of these species have lost the +use of their wings. Evolutionists explain this remarkable fact by their +general laws of degeneration under disuse, <span class="pagenum"><a name="page_227" id="page_227">[227]</a></span> +and the operation of natural selection, as will be shown later on; but +it is not so easy for special creationists to explain why this enormous +number of peculiar species of beetles should have been deposited on +Madeira, all allied to beetles on the nearest continent, and nearly all +deprived of the use of their wings. And similarly, of course, with all +the peculiar species of the Bermudas and the Azores. For who will +explain, on the theory of independent creation, why all the peculiar +species, both of animals and plants, which occur on the Bermudas should +so unmistakably present American affinities, while those which occur on +the Azores no less unmistakably present European affinities? But to +proceed to other, and still more remarkable, cases.</p> + +<p><i>The Galapagos Islands.</i>—This archipelago is of volcanic origin, +situated under the equator between 500 and 600 miles from the West Coast +of South America. The depth of the ocean around them varies from 2,000 +to 3,000 fathoms or more. This group is of particular interest, from the +fact that it was the study of its fauna which first suggested to +Darwin’s mind the theory of evolution. I will, therefore, begin by +quoting a short passage from his writings upon the zoological relations +of this particular fauna.</p> + +<div class='blockquot'><p>Here almost every product of the land and of the water bears the +unmistakeable stamp of the American continent. There are twenty-six +land birds; of these, twenty-one, or perhaps twenty-three, are +ranked as distinct species, and would commonly be assumed to have +been here created; yet the close affinity of most of these birds to +American species is manifest in every character, in their habits, +gestures, and tones of voice.<span class="pagenum"><a name="page_228" id="page_228">[228]</a></span> +So it is with the other animals, and with a large proportion of the +plants, as shown by Dr. Hooker in his admirable Flora of this +archipelago. The naturalist, looking at the inhabitants of these +volcanic islands in the Pacific, distant several hundred miles from the +continent, feels that he is standing on American land. Why should this +be so? Why should the species which are supposed to have been created in +the Galapagos Archipelago, and nowhere else, bear so plainly the stamp +of affinity to those created in America? There is nothing in the +conditions of life, in the geological nature of the islands, in their +height or climate, or in the proportions in which the several classes +are associated together, which closely resembles the conditions of the +South American coast; in fact, there is a considerable dissimilarity in +all these respects. On the other hand, there is a considerable degree of +resemblance in the volcanic nature of the soil, in the climate, height, +and size of the islands, between the Galapagos and Cape de Verde +Archipelagoes; but what an entire and absolute difference in their +inhabitants! The inhabitants of the Cape de Verde Islands are related to +those of Africa, like those of the Galapagos to America. Facts such as +these admit of no sort of explanation on the ordinary view of +independent creation; whereas on the view here maintained, it is obvious +that the Galapagos Islands would be likely to receive colonists from +America, and the Cape de Verde Islands from Africa; such colonists would +be liable to modification—the principle of inheritance still +betraying their original birthplace<a name="FNanchor_22_22" id="FNanchor_22_22"></a><a href="#Footnote_22_22" class="fnanchor">[22]</a>.</p></div> + +<p>The following is a synopsis of the fauna and flora of this archipelago, +so far as at present known. The only terrestrial vertebrates are two +peculiar species of land-tortoise, and one extinct species; five species +of lizards, all peculiar—two of them so much so as to constitute a +peculiar genus;—and two species of snakes, both closely allied to South +American forms. Of birds there are 57 species, of which no <span class="pagenum"><a name="page_229" id="page_229">[229]</a></span> +less than 38 are peculiar; and all the non-peculiar species, except +one, belong to aquatic tribes. The true land birds are represented by 31 +species, of which all, except one, are peculiar; while more than half of +them go to constitute peculiar genera. Moreover, while they are all +unquestionably allied to South American forms, they present a beautiful +series of gradations, “from perfect identity with the continental +species, to genera so distinct that it is difficult to determine with +what forms they are most nearly allied; and it is interesting to note +that this diversity bears a distinct relation to the probabilities of, +and facilities for, migration to the islands. The excessively abundant +rice-bird, which breeds in Canada, and swarms over the whole United +States, migrating to the West Indies and South America, visiting the +distant Bermudas almost every year, and extending its range as far as +Paraquay, is the only species of land-bird which remains completely +unchanged in the Galapagos; and we may therefore conclude that some +stragglers of the migrating host reach the islands sufficiently often to +keep up the purity of the breed<a name="FNanchor_23_23" id="FNanchor_23_23"></a><a href="#Footnote_23_23" class="fnanchor">[23]</a>.” +Again, of the thirty peculiar land-birds, it is observable that the more +they differ from any other species or genera on the South American +continent, the more certainly are they found to have their nearest +relations among those South American forms which have the more +restricted range, and are therefore the least likely to have found their +way to the islands with any frequency.</p> + +<p>The insect fauna of the Galapagos islands is scanty, and chiefly +composed of beetles. These number 35 <span class="pagenum"><a name="page_230" id="page_230">[230]</a></span> +species, which are nearly all peculiar, and in some cases go to +constitute peculiar genera. The same remarks apply to the twenty species +of land-shells. Lastly, of the total number of flowering plants (332 +species) more than one half (174 species) are peculiar. It is observable +in the case of these peculiar species of plants—as also of the +peculiar species of birds—that many of them are restricted to +single islands. It is also observable that, with regard both to the +fauna and flora, the Galapagos Islands as a whole are very much richer +in peculiar species than either the Azores or Bermudas, notwithstanding +that both the latter are considerably more remote from their nearest +continents. This difference, which at first sight appears to make +against the evolutionary interpretation, really tends to confirm it. For +the Galapagos Islands are situated in a calm region of the globe, +unvisited by those periodic storms and hurricanes which sweep over the +North Atlantic, and which every year convey some straggling birds, +insects, seeds, &c., to the Azores and Bermudas. Notwithstanding +their somewhat greater isolation geographically, therefore, the Azores +and Bermudas are really less isolated biologically than are the +Galapagos Islands; and hence the less degree of peculiarity on the part +of their endemic species. But, on the theory of special creation, it is +impossible to understand why there should be any such correlation +between the prevalence of gales and a comparative inertness of creative +activity. And, as we have seen, it is equally impossible on this theory +to understand why there should be a further correlation between the +<i>degree</i> of peculiarity on the part <span class="pagenum"><a name="page_231" id="page_231">[231]</a></span> +of the isolated species, and the degree in which their nearest allies +on the mainland are there confined to narrow ranges, and therefore less +likely to keep up any biological communication with the islands.</p> + +<p><i>St. Helena.</i>—A small volcanic island, ten miles long by eight wide, +situated in mid-ocean, 1100 miles from Africa, and 1800 from South +America. It is very mountainous and rugged, bounded for the most part by +precipices, rising from ocean depths of 17,000 feet, to a height above +the sea-level of nearly 3,000. When first discovered it was richly +clothed with forests; but these were all destroyed by human agency +during the 16th, 17th and 18th centuries. The records of civilization +present no more lamentable instance of this kind of destruction. From a +merely pecuniary point of view the abolition of these primeval forests +has proved an irreparable loss; but from a scientific point of view the +loss is incalculable. These forests served to harbour countless forms of +life, which extended at least from the Miocene age, and which, having +found there an ocean refuge, survived as the last remnants of a remote +geological epoch. In those days, as Mr. Wallace observes, St. Helena +must have formed a kind of natural museum or vivarium of archaic species +of all classes, the interest of which we can now only surmise from the +few remnants of those remnants, which are still left among the more +inaccessible portions of the mountain peaks and crater edges. These +remnants of remnants are as follows.</p> + +<p>There is a total absence of all indigenous mammals, reptiles, +fresh-water fish, and true land-birds. There is, however, a species of +plover, allied to one <span class="pagenum"><a name="page_232" id="page_232">[232]</a></span> +in South Africa; but it is specifically distinct, and therefore +peculiar to the island. The insect life, on the other hand, is abundant. +Of beetles no less than 129 species are believed to be aboriginal, and, +with one single exception, the whole number are peculiar to the island. +“But in addition to this large amount of specific peculiarity +(perhaps unequalled anywhere else in the world), the beetles of this +island are remarkable for their generic isolation, and for the +altogether exceptional proportion in which the great divisions of the +order are represented. The species belong to 39 genera, of which no less +than 25 are peculiar to the island; and many of these are such isolated +forms that it is impossible to find their allies in any particular +country<a name="FNanchor_24_24" id="FNanchor_24_24"></a><a href="#Footnote_24_24" class="fnanchor">[24]</a>.” +More than two-thirds of all the species belong to the group of +weevils—a circumstance which serves to explain the great wealth of +beetle-population, the weevils being beetles which live in wood, and St. +Helena having been originally a densely wooded island. This circumstance +is also in accordance with the view that the peculiar insect fauna has +been in large part evolved from ancestors which reached the island by +means of floating timber; for, of course, no explanation can be +suggested why special creation of this highly peculiar insect fauna +should have run so disproportionately into the production of weevils. +About two-thirds of the whole number of beetles, or over 80 species, +show no close affinity with any existing insects, while the remaining +third have some relations, though often very remote, with European and +African forms. That this high degree of peculiarity <span class="pagenum"><a name="page_233" id="page_233">[233]</a></span> +is due to high antiquity is further indicated, according to our theory, +by the large number of species which some of the types comprise. Thus, +the 54 species of <span class='sn'>Cossonidæ</span> may be referred to three types; the 11 +species of <span class='sn'>Bembidium</span> form a group by themselves; and the <span class='sn'>Heteromera</span> form +two groups. “Now, each of these types may well be descended from a +single species, which originally reached the island from some other +land; and the great variety of generic and specific forms into which +some of them have diverged is an indication, and to some extent a +measure, of the remoteness of their origin<a name="FNanchor_25_25" id="FNanchor_25_25"></a><a href="#Footnote_25_25" class="fnanchor">[25]</a>.” +But, on the counter-supposition that all these 128 peculiar species were +separately created to occupy this particular island, it is surely +unaccountable that they should thus present such an arborescence of +natural affinities amongst themselves.</p> + +<p>Passing over the rest of the insect fauna, which has not yet been +sufficiently worked out, we next find that there are only 20 species of +indigenous land-shells—which is not surprising when we remember by what +enormous reaches of ocean the island is surrounded. Of these 20 species +no less than 13 have become extinct, three are allied to European +species, while the rest are so highly peculiar as to have no near allies +in any other part of the globe. So that the land-shells tell exactly the +same story as the insects.</p> + +<p>Lastly, the plants likewise tell the same story. The truly indigenous +flowering plants are about 50 in number, besides 26 ferns. Forty of the +former and ten of the latter are peculiar to the island, <span class="pagenum"><a name="page_234" id="page_234">[234]</a></span> +and, as Sir Joseph Hooker tells us, “cannot be regarded as very +close specific allies of any other plants at all” Seventeen of +them belong to peculiar genera, and the others all differ so markedly as +species from their congeners, that not one comes under the category of +being an insular form of a continental species. So that with respect to +its plants no less than with respect to its animals, we find that the +island of St. Helena constitutes a little world of unique species, +allied among themselves, but diverging so much from all other known +forms that in many cases they constitute unique genera.</p> + +<p><i>Sandwich Islands.</i>—These are an extensive group of islands, larger +than any we have hitherto considered—the largest of the group being +about the size of Devonshire. The entire archipelago is volcanic, with +mountains rising to a height of nearly 14,000 feet. The group is +situated in the middle of the North Pacific, at a distance of +considerably over 2,000 miles from any other land, and surrounded by +enormous ocean depths. The only terrestrial vertebrata are two lizards, +one of which constitutes a peculiar genus. There are 24 aquatic birds, +five of which are peculiar; four birds of prey, two of which are +peculiar; and 16 land-birds, all of which are peculiar. Moreover, these +16 land-birds constitute no less than 10 peculiar genera, and even one +peculiar family of five genera. This is an amount of peculiarity far +exceeding that of any other islands, and, of course, corresponds with +the great isolation of this archipelago. The only other animals which +have here been carefully studied are the land-shells, and these tell the +same story as the birds. For there are <span class="pagenum"><a name="page_235" id="page_235">[235]</a></span> +no less than 400 species which are all, without any exception, +peculiar; while about three-quarters of them go to constitute peculiar +genera. Again, of the plants, 620 species are believed to be endemic; +and of these 377 are peculiar, yielding no less than 39 peculiar genera.</p> + +<hr class='minor' /> + +<p>Prejudice apart, I think we must all now agree that it is needless to +continue further this line of proof. I have chosen the smallest and most +isolated islands for the purposes of our present argument, first because +these furnish the most crucial kind of test, and next because they best +admit of being dealt with in a short space. But, if necessary, a vast +amount of additional material could be furnished, not only from other +small oceanic islands, but still more from the largest islands of the +world, such as Australia and New Zealand. However, after the detailed +inventories which have now been given in the case of some of the smaller +islands most remote from mainlands, we may well be prepared to accept it +as a general law, that <i>wherever</i> there is evidence of land-areas having +been for a long time separated from other land-areas, there we meet with +a more or less extraordinary profusion of unique species, often running +up into unique genera. And, in point of fact, so far as naturalists have +hitherto been able to ascertain, <i>there is no exception to this general +law in any region of the globe</i>. Moreover, there is everywhere a +constant correlation between the <i>degree</i> of this peculiarity on the +part of the fauna and flora, and the <i>time</i> during which they have been +isolated. Thus, for instance, among the islands which<span class="pagenum"><a name="page_236" id="page_236">[236]</a></span> +I have called into evidence, those that are at once the most isolated +and give independent proofs of the highest antiquity, are the Galapagos +Islands, the Sandwich Islands, and St. Helena. Now, if we apply the +method of tabular analysis to these three cases, we obtain the following +most astonishing results. For the sake of simplicity I will omit the +enumeration of peculiar genera, and confine attention to peculiar +species. Moreover, I will consider only terrestrial animals; for, as we +have already seen, aquatic animals are so much more likely to reach +oceanic islands that they do not furnish nearly so fair a test of the +evolutionary hypothesis.</p> + +<div class='center mt2'> +<table border="1" cellpadding="4" cellspacing="0" summary=""> +<thead class='c'><tr><td colspan='6'>PECULIAR SPECIES.</td></tr></thead> +<tbody><tr class='i'><td></td><td class='c'>Shells.</td><td class='c'>Insects.</td><td class='c'>Reptiles.</td><td class='c'>Birds.</td><td class='c'>Mammals.</td></tr> +<tr><td class='l'>Sandwich</td><td class='r'>400</td><td class='r'>?</td><td class='r'>2</td><td class='r'>16</td><td class='r'>0</td></tr> +<tr><td class='l'>Galapagos</td><td class='r'>15</td><td class='r'>35</td><td class='r'>10</td><td class='r'>30</td><td class='r'>0</td></tr> +<tr><td class='l'>St. Helena</td><td class='r'>20</td><td class='r'>128</td><td class='r'>0</td><td class='r'>1</td><td class='r'>0</td></tr> +<tr><td class='l'>Totals.</td><td class='r'>435</td><td class='r'>163</td><td class='r'>12</td><td class='r'>47</td><td class='r'>0</td></tr> +</tbody></table></div> + +<div class='center mt2'> +<table border="1" cellpadding="4" cellspacing="0" summary=""> +<thead class='c'><tr><td colspan='6'>NON-PECULIAR SPECIES.</td></tr></thead> +<tbody><tr class='i'><td></td><td class='c'>Shells.</td><td class='c'>Insects.</td><td class='c'>Reptiles.</td><td class='c'>Birds.</td><td class='c'>Mammals.</td></tr> +<tr><td class='l'>Sandwich</td><td class='r'>0</td><td class='r'>?</td><td class='r'>0</td><td class='r'>0</td><td class='r'>0</td></tr> +<tr><td class='l'>Galapagos</td><td class='r'>?</td><td class='r'>?</td><td class='r'>0</td><td class='r'>1</td><td class='r'>0</td></tr> +<tr><td class='l'>St. Helena</td><td class='r'>0</td><td class='r'>?</td><td class='r'>0</td><td class='r'>0</td><td class='r'>0</td></tr> +<tr><td class='l'>Totals.</td><td class='r'>0</td><td class='r'>?</td><td class='r'>0</td><td class='r'>1</td><td class='r'>0</td></tr> +</tbody></table></div> + +<p>From this synopsis we perceive that out of a total of 658 species of +terrestrial animals known to inhabit <span class="pagenum"><a name="page_237" id="page_237">[237]</a></span> +these three oceanic territories, all are peculiar, with the exception +of a single land-bird which is found in the Galapagos Islands. This is +the rice-bird, so very abundant on the American continent that its +representatives must not unfrequently become the involuntary colonists +of the Archipelago. There are, however, a few species of non-peculiar +insects inhabiting the Sandwich and Galapagos Islands, the exact number +of which is doubtful, and on this account are not here quoted. But at +most they would be represented by units, and therefore do not affect the +general result. Lastly, the remarkable fact will be noted, that there is +no single representative of the mammalian class in any of these islands.</p> + +<p>If we turn next to consider the case of plants, we obtain the following +result:—</p> + +<div class='center'> +<table border="0" cellpadding="4" cellspacing="0" summary=""> +<tr class='c i'><td></td><td>Peculiar<br />Species.</td><td> </td><td>Non-peculiar<br />Species.</td></tr> +<tr><td class='l'>Sandwich</td><td class='r'>377</td><td> </td><td class='r'>243</td></tr> +<tr><td class='l'>Galapagos</td><td class='r'>174</td><td> </td><td class='r'>158</td></tr> +<tr><td class='l'>St. Helena</td><td class='r'>50</td><td> </td><td class='r'>26</td></tr> +<tr><td class='l'>Totals.</td><td class='r'>601</td><td> </td><td class='r'>427</td></tr> +</table></div> + +<p>So that by adding together peculiar species both of land-animals and +plants, we find that on these three limited areas alone there are 1258 +forms of life which occur nowhere else upon the globe—not to speak of +the peculiar aquatic species, nor of the presumably large number of +peculiar species of all kinds not hitherto discovered in these +imperfectly explored regions.</p> + +<p>Now let us compare these facts with those which are presented by the +faunas and floras of islands less <span class="pagenum"><a name="page_238" id="page_238">[238]</a></span> +remote from continents, and known from independent geological evidence +to be of comparatively recent origin—that is, to have been +separated from their adjacent mainlands in comparatively recent times, +and therefore as islands to be comparatively young. The British Isles +furnish as good an instance as could be chosen, for they together +comprise over 1000 islands of various sizes, which are nowhere separated +from one another by deep seas, and in the opinion of geologists were all +continuous with the European continent since the glacial period.</p> + +<div class='center mt2'> +<table border="1" cellpadding="4" cellspacing="0" summary=""> +<thead class='c'><tr><td colspan='6'>BRITISH ISLES.</td></tr></thead> +<tbody> +<tr><td class='c sc lc' colspan='6'>NON-PECULIAR SPECIES.</td></tr> +<tr><td class='c'>Plants.</td><td class='c'>Land Shells.</td><td class='c'>Insects.</td><td class='c'>Reptiles and Amphibia.</td><td class='c'>Land Birds.</td><td class='c'>Land Mammals.</td></tr> +<tr><td class='c'>1462</td><td class='c'>83</td><td class='c'>12,551</td><td class='c'>13</td><td class='c'>130</td><td class='c'>40</td></tr> +<tr><td class='c sc lc' colspan='6'>PECULIAR SPECIES.</td></tr> +<tr><td class='c'>Plants.</td><td class='c'>Land Shells.</td><td class='c'>Insects.</td><td class='c'>Reptiles and Amphibia.</td><td class='c'>Land Birds.</td><td class='c'>Land Mammals.</td></tr> +<tr><td class='c'>46</td><td class='c'>4</td><td class='c'>149</td><td class='c'>0</td><td class='c'>1</td><td class='c'>0</td></tr></tbody> +</table></div> + +<div class='center mt2'> +<table border="0" cellpadding="4" cellspacing="0" summary=""> +<tr><td class='l'>Total Peculiar Plants</td><td class='r'>46</td></tr> +<tr><td class='l'>Total Peculiar Animals</td><td class='r'>154</td></tr> +<tr><td></td><td class='r'>——</td></tr> +<tr><td class='l'>Grand Total</td><td class='r'>200</td></tr> +</table></div> + +<p>I have drawn up this table in the most liberal manner possible, +including as peculiar species forms <span class="pagenum"><a name="page_239" id="page_239">[239]</a></span> +which many naturalists regard as merely local varieties. But, even as +thus interpreted, how wonderful is the contrast between the 1000 islands +of Great Britain and the single volcanic rock of St. Helena, where +almost all the animals and about half the plants are peculiar, instead +of about 1/80 of the animals, and 1/30 of the plants. Of course, if no +peculiar species of any kind had occurred in the British Isles, +advocates of special creation might have argued that it was, so to +speak, needless for the Divinity to have added any new species to those +European forms which fully populated the islands at the time when they +were separated from the continent. But, as the matter stands, advocates +of special creation must face the fact that a certain small number of +new and peculiar species have been formed on the British Isles; and, +therefore, that creative activity has not been wholly suspended in their +case. Why, then, has it been so meagre in this case of a thousand +islands, when it has proved so profuse in the case of all single islands +more remote from mainlands, and presenting a higher antiquity? Or why +should the Divinity have thus appeared so uniformly to consult these +merely accidental circumstances of space and time in the depositing of +his unique specific types? Do not such facts rather speak with +irresistible force in favour of the view, that while all ancient and +solitary islands have had time enough, and separation enough, to admit +of distinct histories of evolution having been written in their living +inhabitants, no one of the thousand islands of Great Britain has had +either time enough, or separation enough, to have admitted of more than +some of the first pages of such a history having been commenced?</p> + +<p><span class="pagenum"><a name="page_240" id="page_240">[240]</a></span> +But this allusion to Great Britain introduces us to another point. It +will have been observed that, unlike oceanic islands remote from +mainlands, Great Britain is well furnished both with reptiles (including +amphibia) and mammals. For there is no instance of any oceanic island +situated at more than 300 miles from a continent where any single +species of the whole class of mammals is to be found, excepting species +of the only order which is able to fly—namely, the bats. And the same +has to be said of frogs, toads, and newts, whose spawn is quickly killed +by contact with sea-water, and therefore could never have reached remote +islands in a living state. Hence, on evolutionary principles; it is +quite intelligible why oceanic islands should not present any species of +mammals or batrachians—peculiar or otherwise,—save such species of +mammals as are able to fly. But on the theory of special creation we can +assign no reason why, notwithstanding the extraordinary profusion of +unique types of other kinds which we have seen to occur on oceanic +islands, the Deity should have made this curious exception to the +detriment of all frogs, toads, newts, and mammals, save only such as are +able to fly. Or, if any one should go so far to save a desperate +hypothesis as to maintain that there must have been some hidden reason +why batrachians and quadrupeds were not specially created on oceanic +islands, I may mention another small—but in this relation a most +significant—fact. This is that on some of these islands there occur +certain peculiar species of plants, the seeds of which are provided with +numerous tiny hooks, obviously and beautifully adapted—like those on +the seeds of allied plants elsewhere—to catch the <span class="pagenum"><a name="page_241" id="page_241">[241]</a></span> +wool or hair of moving quadrupeds, and so to further their own +dissemination. But, as we have just seen, there are no quadrupeds in the +islands to meet these beautiful adaptations on the part of the plants; +so that special creationists must resort to the almost impious +supposition that in these cases the Deity has only carried out half his +plan, in that while he made an elaborate provision for these uniquely +created species of plants, which depended for its efficiency on the +presence of quadrupeds, he nevertheless neglected to place any +quadrupeds on the islands where he had placed the plants. Such one-sided +attempts at adaptation surely resolve the thesis of special creation to +a <i>reductio ad absurdum</i>; and hence the only reasonable interpretation +of them is, that while the seeds of allied or ancestral plants were able +to float to the islands, no quadrupeds were ever able over so great a +distance to swim.</p> + +<hr class='minor' /> + +<p>Although much more evidence might still be given under the head of +geographical distribution, I must now close with a brief summary of the +main points that have been adduced.</p> + +<p>After certain preliminary considerations, I began by noticing that the +theory of evolution has a much more intelligible account to give than +has its rival of the facts of discontinuous distribution—the Alpine +flora, for instance, being allied to the Arctic, not because the same +species were separately created in both places, but because during the +glacial period these species extended all over Europe, and were left +behind on the Alps as the Arctic flora receded northwards—which was +sufficiently long ago to explain <span class="pagenum"><a name="page_242" id="page_242">[242]</a></span> +why some of the Alpine species are unique, though closely allied to +Arctic forms.</p> + +<p>Next we saw that, although living things are always adapted to the +climates under which they live (since otherwise they could not live +there at all), it is equally true that, as a rule, besides the area on +which they do live, there are many other areas in different parts of the +globe where they might have lived equally well. Consequently we must +conclude that, if all species were separately created, many species were +severally created on only one among a number of areas where they might +equally well have thrived. Now, although this conclusion in itself may +not seem opposed to the theory of special creation, a most serious +difficulty is raised when it is taken in connexion with another fact of +an equally general kind. This is, that on every biological region we +encounter chains of allied species constituting allied genera, families, +and so on; while we scarcely ever meet with allied species in different +biological regions, notwithstanding that their climates may be similar, +and, consequently, just as well suited to maintain some of the allied +species. Hence we must further conclude, if all species were separately +created, that in the work of creation some unaccountable regard was paid +to making areas of distribution correspond to degrees of structural +affinity. A great many species of the rat genus were created in the Old +World, and a great many species of another, though allied, genus were +created in the New World: yet no reason can be assigned why no one +species of the Old World series should not just as well have been +deposited in the New World, and <i>vice versa</i>. On the other hand, the +theory of evolution <span class="pagenum"><a name="page_243" id="page_243">[243]</a></span> +may claim as direct evidence in its support all the innumerable cases +such as these—cases, indeed, so innumerable that, as Mr. Wallace +remarks, it may be taken as a law of nature that “every species +has come into existence coincident both in space and time with a +pre-existing and closely allied species.” A general law which, +while in itself most strongly suggestive of evolution, is surely +impossible to reconcile with any reasonable theory of special creation. +Furthermore, this law extends backwards through all geological time, +with the result that the extinct species which now occur only as fossils +on any given geological area, resemble the species still living upon +that area, as we should expect that they must, if the former were the +natural progenitors of the latter. On the other hand, if they were not +the natural progenitors, but all the species, both living and extinct, +were the supernatural and therefore independent creations which the +rival theory would suppose, then no reason can be given why the extinct +species should thus resemble the living—any more than why the +living species should resemble one another. For, as we have seen, there +are almost always many other habitats on other parts of the globe, where +any members of any given group of species might equally well have been +deposited; and this, of course, applies to geological no less than to +historical time. Yet throughout all time we meet with this most +suggestive correlation between continuity of a geographical area and +structural affinity between the forms of life which have lived, or are +still living, upon that area.</p> + +<p>Similarly, we find the further, and no less suggestive, <span class="pagenum"><a name="page_244" id="page_244">[244]</a></span> +correlation between the birth of new species and the immediate +pre-existence of closely allied species on the same area—or, at +most, on closely contiguous areas.</p> + +<p>Where a continuous area has long been circumscribed by barriers of any +kind, which prevent the animals from wandering beyond it, then we find +that all the species, both extinct and living, constitute more or less a +world of their own; while, on the other hand, where the animals are free +to migrate from one area to another, the course of their migrations is +marked by the origination of new species springing up <i>en route</i>, and +serving to connect the older, or metropolitan, forms with the younger, +or colonising, forms in the way of a graduated series. This principle, +however, admits of being traced only in certain cases of species +belonging to the same genus, of genera belonging to the same family, or, +at most, of families belonging to the same order. In other words, the +more general the structural affinity, the more general is the +geographical extension—as we should expect to be the case on the theory +of descent with branching modifications, seeing that the larger, the +older, and the more diverse the group of organisms compared, the greater +must be their chances of dispersal.</p> + +<p>These general considerations led us to contemplate more in detail the +correlation between structural affinity and barriers to free migration. +Such barriers, of course, differ in the cases of different organisms. +Marine organisms are stopped by land, unsuitable temperature, or +unsuitable depths; fresh-water organisms by sea and by mountain-chains; +terrestrial organisms chiefly by water. Now it is a matter of <span class="pagenum"><a name="page_245" id="page_245">[245]</a></span> +fact which admits of no dispute, that in each of these cases we meet +with a direct correlation between the kind of barrier and the kind of +organisms whose structural affinities are affected thereby. Where we +have to do with marine organisms, barriers such as the Isthmus of Panama +and the varying depth of the Western Pacific determine three very +distinct faunas, ranging north and south in closely parallel lines, and +under corresponding climates. Where we have to do with fresh-water +organisms, we find that a mountain-chain only a few miles wide has more +influence in determining differences of organic type on either side of +it than is exercised by even thousands of miles of a continuous +land-area, if this be uninterrupted by any mountains high enough to +prevent water-fowl, whirlwinds, &c., from dispersing the ova. Again, +where we have to do with terrestrial organisms, the most effectual +barriers are wide reaches of ocean; and, accordingly, we find that these +exercise an enormous influence on the modification of terrestrial types. +Moreover, we find that the <i>more</i> terrestrial an organism, or the +<i>greater</i> the difficulty it has in traversing a wide reach of ocean, the +<i>greater</i> is the modifying influence of such a barrier upon that type. +In oceanic islands, for example, many of the plants and aquatic birds +usually belong to the same species as those which occur on the nearest +mainlands, and where there are any specific differences, these but +rarely run up to generic differences. But the land-birds, insects, and +reptiles which are found on such islands are nearly always specifically, +and very often generically, distinct from those on the nearest +mainland—although invariably allied with sufficient closeness +<span class="pagenum"><a name="page_246" id="page_246">[246]</a></span> +to leave no manner of doubt as to their affinities with the fauna of +that mainland. Lastly, no amphibians and no mammals (except bats) are +ever found on any oceanic islands. Yet, as we have seen, on the theory +of special creation, these islands must all be taken to have been the +theatres of the most extraordinary creative activity, so that on only +three of them we found no less than 1258 unique species, whereof 657 +were unique species of land animals, to be set against one single +species known to occur elsewhere. Nevertheless, notwithstanding this +prodigious expenditure of creative energy in the case of land-birds, +land-shells, insects, and reptiles, no single new amphibian, or no +single new mammal, has been created on any single oceanic island, if we +except the only kind of mammal that is able to fly, and the ancestors of +which, like those of the land-birds and insects, might therefore have +reached the islands ages ago. Moreover, with regard to mammals, even in +cases where allied forms occur on either side of a sea-channel, it is +found to be a general rule that if the channel is shallow, the species +on either side of it are much more closely related than if it be +deep—and this irrespective of its width. Therefore we can only +conclude, in the words of Darwin—"As the amount of modification +which animals of all kinds undergo partly depends on lapse of time, and +as the islands which are separated from each other or from the mainland +by shallow channels are more likely to have been continuously united +within a recent period than islands separated by deeper channels, we can +understand how it is that a relation exists between the depth of the sea +separating two mammalian <span class="pagenum"><a name="page_247" id="page_247">[247]</a></span> +faunas, and the degree of their affinity—a relation which is +quite inexplicable on the theory of independent acts of creation.”</p> + +<hr class='minor' /> + +<p>Looking to all these general principles of geographical distribution, +and remembering the sundry points of smaller detail relating to oceanic +islands which I will not wait to recapitulate, to my mind it seems that +there is no escape from the following conclusion, with which I will +bring my brief epitome of the evidence to a close. The conclusion to +which, I submit, all the evidence leads is, that if the doctrine of +special creation is taken to be true, then it must be further taken that +the one and only principle which has been consistently followed in the +geographical deposition of species, is that of so depositing them as to +make it everywhere appear that they were not thus deposited at all, but +came into existence where they now occur by way of genetic descent with +perpetual migration and correlative modification. On no other principle, +so far as I can see, would it be possible to account for the fact that +“every species has come into existence coincident both in space +and time with a pre-existing and closely allied species,” together +with the carefully graduated regard to physical barriers which the +Creator must have displayed while depositing his newly formed species on +either sides of them—everywhere making <i>degrees</i> of structural affinity +correspond to <i>degrees</i> of geographical continuity, and <i>degrees</i> of +structural difference correspond to <i>degrees</i> of geographical +separation, whether by mountain-chains in the case of fresh-water +faunas, by land and by deep sea in the case of marine <span class="pagenum"><a name="page_248" id="page_248">[248]</a></span> +faunas, or by reaches of ocean in the case of terrestrial +faunas—stocking oceanic islands with an enormous profusion of +peculiar species all allied to those on the nearest mainlands, yet +everywhere avoiding the creation upon them of any amphibian or mammal, +except an occasional bat. We are familiar with the doctrine that God is +a God who hideth himself; here, however, it seems to me, we should have +but a thinly-veiled insinuation, not merely that in his works he is +hidden, but that in these works he is untrue. Than which I cannot +conceive a stronger condemnation of the theory which it has been my +object fairly to represent and dispassionately to criticise.</p> + +<hr /><p class="pagenum"><a name="page_249" id="page_249">[249]</a></p> +<h2><a name="SECTION_II" id="SECTION_II"></a>SECTION II<br /><br /> +<span class='i'>SELECTION</span></h2> +<hr class='minor' /><p class="pagenum"><a name="page_251" id="page_251">[251]</a></p> +<h2><a name="CHAPTER_VII" id="CHAPTER_VII"></a>CHAPTER VII.<br /><br /> +<span class="sc">The Theory of Natural Selection.</span></h2> + +<p>Thus far we have been considering the main evidences of organic +evolution considered as a fact. We now enter a new field, namely, the +evidences which thus far have been brought to light touching the causes +of organic evolution considered as a process.</p> + +<p>As was pointed out in the opening chapter, this is obviously the +methodical course to follow: we must have some reasonable assurance that +a fact is a fact before we endeavour to explain it. Nevertheless, it is +not necessary that we should actually demonstrate a fact to be a fact +before we endeavour to explain it. Even if we have but a reasonable +presumption as to its probability, we may find it well worth while to +consider its explanation; for by so doing we may obtain additional +evidence of the fact itself. And this because, if it really is a fact, +and if we hit upon the right explanation of it, by proving the +explanation probable, we may thereby greatly increase our evidence of +the fact. In the very case before us, for example, the evidence of +evolution as a fact has from the first been largely derived from testing +Darwin’s theory concerning its method. It was this theoretical +explanation of its <span class="pagenum"><a name="page_252" id="page_252">[252]</a></span> +method which first set him seriously to enquire into the evidences of +evolution as a fact; and ever since he published his results, the +evidences which he adduced in favour of natural selection as a method +have constituted some of the strongest reasons which scientific men have +felt for accepting evolution as a fact. Of course the evidence in favour +of this fact has gone on steadily growing, quite independently of the +assistance which was thus so largely lent to it by the distinctively +Darwinian theory of its method; and, indeed, so much has this been the +case, that in the present treatise we have been able to consider such +direct evidence of the fact itself, without any reference at all to the +indirect or accessory evidence which is derived from that of natural +selection as a method. From which it follows that in most of what I am +about to say in subsequent chapters on the evidences of natural +selection as a method, there will be furnished a large addition to the +evidences which have already been detailed of evolution as a fact. But, +as a matter of systematic treatment, I have thought it desirable to keep +these two branches of our subject separate. Which means that I have made +the evidences of evolution as a fact to stand independently on their own +feet—feet which in my opinion are amply strong enough to bear any +weight of adverse criticism that can be placed upon them.</p> + +<p>Our position, then, is this. On the foundation of the previous chapters, +I will henceforth assume that we all accept organic evolution as a fact, +without requiring any of the accessory evidence which is gained by +independent proof of natural selection as a method. But in making this +assumption—namely, that we are all now firmly persuaded of the fact of +evolution—I do <span class="pagenum"><a name="page_253" id="page_253">[253]</a></span> +not imagine that such is really the case. I make the assumption for the +purposes of systematic exposition, and in order that different parts of +the subject may be kept distinct. I confess it does appear to me +remarkable that there should still be a doubt in any educated mind +touching the general fact of evolution; while it becomes to me +unaccountable that such should be the case with a few still living men +of science, who cannot be accused of being ignorant of the evidences +which have now been accumulated. But in whatever measure we may +severally have been convinced—or remained unconvinced—on +this matter, for the purposes of exposition I must hereafter assume that +we are all agreed to the extent of regarding the process of evolution +as, at least, sufficiently probable to justify enquiry touching its +causes on supposition of its truth.</p> + +<p>Now, the causes of evolution have been set forth in a variety of +different hypotheses, only the chief of which need be mentioned here. +Historically speaking the first of these was that which was put forward +by Erasmus Darwin, Lamarck, and Herbert Spencer. It consists in putting +together the following facts and inferences.</p> + +<p>We know that, in the lifetime of the individual, increased use of +structures leads to an increase of their functional efficiency; while, +on the other hand, disuse leads to atrophy. The arms of a blacksmith, +and the legs of a mountaineer, are familiar illustrations of the first +principle: our hospital wards are full of illustrations of the second. +Again, we know that the characters of parents are transmitted to their +progeny by means of heredity. Now the hypothesis <span class="pagenum"><a name="page_254" id="page_254">[254]</a></span> +in question consists in supposing that if any particular organs in a +species are habitually used for performing any particular action, they +must undergo a structural improvement which would more and more adapt +them to the performance of that action; for in each generation constant +use would better and better adapt the structures to the discharge of +their functions, and they would then be bequeathed to the next +generation in this their improved form by heredity. So that, for +instance, if there had been a thousand generations of blacksmiths, we +might expect the sons of the last of them to inherit unusually strong +arms, even if these young men had themselves taken to some other trade +not requiring any special use of their arms. Similarly, if there had +been a thousand generations of men who used their arms but slightly, we +should expect their descendants to show but a puny development of the +upper extremities. Now let us apply all this to the animal kingdom in +general. The giraffe, for instance, is a ruminant whose entire frame has +been adapted to support an enormously long neck, which is of use to the +animal in reaching the foliage of trees. The ancestors of the giraffe, +having had ordinary necks, were supposed by Lamarck to have gradually +increased the length of them, through many successive generations, by +constantly stretching to reach high foliage; and he further supposed +that, when the neck became so long as to require for its support special +changes in the general form of the animal as a whole, these special +changes would have brought about the dwindling of other parts from which +so much activity was no longer required—the general result being +that the whole organization of the animal became more and more +<span class="pagenum"><a name="page_255" id="page_255">[255]</a></span> +adapted to browsing on high foliage. And so in the cases of other +animals, Lamarck believed that the adaptation of their forms to their +habits could be explained by this simple hypothesis that the habits +created the forms, through the effects of use and disuse, coupled with +heredity.</p> + +<p>Such is what is ordinarily known as Lamarck’s theory of evolution. +We may as well remember, however, that it really constitutes only one +part of his theory; for besides this hypothesis of the cumulative +inheritance of functionally-produced modifications—to which we may add +the inherited effects of any direct action exercised by surrounding +conditions of life,—Lamarck believed in some transcendental principle +tending to produce gradual improvement in pre-determined lines of +advance. Therefore it would really be more correct to designate the +former hypothesis by the name either of Erasmus Darwin, or, still +better, of Herbert Spencer. Nevertheless, in order to avoid confusion, I +will follow established custom, and subsequently speak of this +hypothesis as the Lamarckian hypothesis—understanding, however, that in +employing this designation I am not referring to any part or factor of +Lamarck’s general theory of evolution other than the one which has +just been described—namely, the hypothesis of the cumulative +transmission of functionally-produced, or otherwise +“acquired,” modifications.</p> + +<p>This, then, was the earliest hypothesis touching the causes of organic +evolution. But we may at once perceive that it is insufficient to +explain all that stands to be explained. In the first place, it refers +in chief part only to the higher animals, which are actuated to +<span class="pagenum"><a name="page_256" id="page_256">[256]</a></span> +effort by intelligence. Its explanatory power in the case of most +invertebrata—as well as in that of all plants—is extremely +limited, inasmuch as these organisms can never be moved to a greater or +less use of their several parts by any discriminating volition, such as +that which leads to the continued straining of a giraffe’s neck +for the purpose of reaching foliage. In the second place, even among the +higher animals there are numberless tissues and organs which +unquestionably present a high degree of adaptive evolution, but which +nevertheless cannot be supposed to have fallen within the influence of +Lamarckian principles. Of such are the shells of crustacea, tortoises, +&c., which although undoubtedly of great use to the animals +presenting them, cannot ever have been <i>used</i> in the sense required by +Lamarck’s hypothesis, i. e. actively exercised, so as to increase +a flow of nutrition to the part. Lastly, in the third place, the +validity of Lamarck’s hypothesis in any case whatsoever has of +late years become a matter of serious question, as will be fully shown +and discussed in the next volume. Meanwhile it is enough to observe +that, on account of all these reasons, the theory of Lamarck, even if it +be supposed to present any truth at all, is clearly insufficient as a +full or complete theory of organic evolution.</p> + +<hr class='minor' /> + +<p>In historical order the next theory that was arrived at was the theory +of natural selection, simultaneously published by Darwin and Wallace on +July 1st, 1858.</p> + +<p>If we may estimate the importance of an idea by the change of thought +which it effects, this idea of natural selection is unquestionably the +most important <span class="pagenum"><a name="page_257" id="page_257">[257]</a></span> +idea that has ever been conceived by the mind of man. Yet the wonder is +that it should not have been hit upon long before. Or rather, I should +say, the wonder is that its immense and immeasurable importance should +not have been previously recognised. For, since the publication of this +idea by Darwin and Wallace, it has been found that its main features had +already occurred to at least two other minds—namely, Dr. Wells in +1813, and Mr. Patrick Matthew in 1831. But neither of these writers +perceived that in the few scattered sentences which they had written +upon the subject they had struck the key-note of organic nature, and +resolved one of the principal chords of the universe. Still more +remarkable is the fact that Mr. Herbert Spencer—notwithstanding +his great powers of abstract thought and his great devotion of those +powers to the theory of evolution, when as yet this theory was scorned +by science—still more remarkable, I say, is the fact that Mr. +Herbert Spencer should have missed what now appears so obvious an idea. +But most remarkable of all is the fact that Dr. Whewell, with all his +stores of information on the history of the inductive sciences, and with +all his acumen on the matter of scientific method, should not only have +conceived the idea of natural selection, but expressly stated it as a +logically possible explanation of the origin of species, and yet have so +stated it merely for the purpose of dismissing it with contempt<a +name="FNanchor_26_26" id="FNanchor_26_26"></a><a href="#Footnote_26_26" class="fnanchor">[26]</a>. +This, I think, is most remarkable, because it serves to prove how very +far men’s minds at that time must have been from entertaining, as +in any way antecedently probable, the doctrine of transmutation. In +order to show this I will here quote one <span class="pagenum"><a name="page_258" id="page_258">[258]</a></span> +passage from the writings of Whewell, and another from a distinguished +French naturalist referred to by him.</p> + +<p>In 1846 Whewell wrote:—</p> + +<div class='blockquot'><p>Not only is the doctrine of the transmutation of species in itself +disproved by the best physiological reasonings, but the additional +assumptions which are requisite to enable its advocates to apply it +to the explanation of the geological and other phenomena of the +earth, are altogether gratuitous and fantastical<a name="FNanchor_27_27" id="FNanchor_27_27"></a><a href="#Footnote_27_27" class="fnanchor">[27]</a>.</p></div> + +<p>Then he quotes with approval the following opinion:—</p> + +<div class='blockquot'><p>Against this hypothesis, which, up to the present time, I regard as +purely gratuitous, and likely to turn geologists out of the sound +and excellent road in which they now are, I willingly raise my +voice, with the most absolute conviction of being in the right<a name="FNanchor_28_28" id="FNanchor_28_28"></a><a href="#Footnote_28_28" class="fnanchor">[28]</a>.</p></div> + +<p>And, after displaying the proof rendered by Lyell of uniformitarianism +in geology, and cordially subscribing thereto, Whewell adds:—</p> + +<div class='blockquot'><p>We are led by our reasonings to this view, that the present order +of things was commenced by an act of creative power entirely +different to any agency which has been exerted since. None of the +influences which have modified the present races of animals and +plants since they were placed in their habitations on the +earth’s surface can have had any efficacy in producing them +at first. We are necessarily driven to assume, as the beginning of +the present cycle of organic nature, an event not included in the +course of nature<a name="FNanchor_29_29" id="FNanchor_29_29"></a><a href="#Footnote_29_29" class="fnanchor">[29]</a>.</p></div> + +<p>So much, then, for the state of the most enlightened and representative +opinions on the question of evolution <span class="pagenum"><a name="page_259" id="page_259">[259]</a></span> +before the publication of Darwin’s work; and so much, likewise, +for the only reasonable suggestions as to the causes of evolution which +up to that time had been put forward, even by those few individuals who +entertained any belief in evolution as a fact. It was the theory of +natural selection that changed all this, and created a revolution in the +thought of our time, the magnitude of which in many of its far-reaching +consequences we are not even yet in a position to appreciate; but the +action of which has already wrought a transformation in general +philosophy, as well as in the more special science of biology, that is +without a parallel in the history of mankind.</p> + +<hr class='minor' /> + +<p>Although every one is now more or less well acquainted with the theory +of natural selection, it is necessary, for the sake of completeness, +that I should state the theory; and I will do so in full detail.</p> + +<p>It is a matter of observable fact that all plants and animals are +perpetually engaged in what Darwin calls a “struggle for +existence.” That is to say, in every generation of every species a +great many more individuals are born than can possibly survive; so that +there is in consequence a perpetual battle for life going on among all +the constituent individuals of any given generation. Now, in this +struggle for existence, which individuals will be victorious and live? +Assuredly those which are best fitted to live, in whatever respect, or +respects, their superiority of fitness may consist. Hence it follows +that Nature, so to speak, <i>selects</i> the best individuals out of each +generation to live. And not only so; but as these favoured individuals +transmit their favourable qualities to their offspring, according to +<span class="pagenum"><a name="page_260" id="page_260">[260]</a></span> +the fixed laws of heredity, it further follows that the individuals +composing each successive generation have a general tendency to be +better suited to their surroundings than were their forefathers. And +this follows, not merely because in every generation it is only the +“flower of the flock” that is allowed to breed, but also +because, if in any generation some new and beneficial qualities happen +to arise as slight variations from the ancestral type, they will (other +things permitting) be seized upon by natural selection, and, being +transmitted by heredity to subsequent generations, will be added to the +previously existing type. Thus the best idea of the whole process will +be gained by comparing it with the closely analogous process whereby +gardeners, fanciers, and cattle-breeders create their wonderful +productions; for just as these men, by always “<i>selecting</i>” +their best individuals to breed from, slowly but continuously improve +their stock, so Nature, by a similar process of +“<i>selection</i>” slowly but continuously makes the various +species of plants and animals better and better suited to the conditions +of their life.</p> + +<p>Now, if this process of continuously adapting organisms to their +environment takes place in nature at all, there is no reason why we +should set any limits on the extent to which it is able to go, up to the +point at which a complete and perfect adaptation is achieved. Therefore +we might suppose that all species would eventually reach this condition +of perfect harmony with their environment, and then remain fixed. And +so, according to the theory, they would, if the environment were itself +unchanging. But forasmuch as the environment (i. e. the sum total of the +<span class="pagenum"><a name="page_261" id="page_261">[261]</a></span> +external conditions of life) of almost every organic type alters more +or less from century to century—whether from astronomical, +geological, and geographical changes, or from the immigrations and +emigrations of other species living on contiguous areas, and so +on—it follows that the process of natural selection need never +reach a terminal phase. And forasmuch as natural selection may thus +continue, <i>ad infinitum</i>, slowly to alter a specific type in adaptation +to a gradually changing environment, if in any case the alteration thus +effected is sufficient in amount to lead naturalists to name the result +as a distinct species, it follows that natural selection has transmuted +one specific type into another. Similarly, by a continuation of the +process, specific types would become transmuted into generic, generic +into family types, and so on. Thus the process is supposed to go on +throughout all the countless forms of life continuously and +simultaneously—the world of organic types being thus regarded as +in a state of perpetual, though gradual, flux.</p> + +<hr class='minor' /> + +<p>Now, the first thing we have to notice about this theory is, that in all +its main elements it is merely a statement of observable facts. It is an +observable fact that in all species of plants and animals a very much +larger number of individuals are born than can possibly survive. Thus, +for example, it has been calculated that if the progeny of a single pair +of elephants—which are the slowest breeding of animals—were all +allowed to reach maturity and propagate, in 750 years there would be +living 19,000,000 descendants. Again, in the case of vegetables, if a +species of annual plant produces only two seeds a <span class="pagenum"><a name="page_262" id="page_262">[262]</a></span> +year, if these in successive years were all allowed to reproduce their +kind, in twenty years there would be 11,000,000 plants from a single +ancestor. Yet we know that nearly all animals and plants produce many +more young at a time than in either of these two supposed cases. Indeed, +as individuals of many kinds of plants, and not a few kinds of animals, +produce every year several thousand young, we may make a rough estimate +and say, that over organic nature as a whole probably not one in a +thousand young are allowed to survive to the age of reproduction. How +tremendous, therefore, must be the struggle for existence! It is thought +a terrible thing in battle when one half the whole number of combatants +perish. But what are we to think of a battle for life where only one in +a thousand survives?</p> + +<p>This, then, is the first fact. The second is the fact so long ago +recognised, that the battle is to the strong, the race to the swift. The +thousandth individual which does survive in the battle for +existence—which does win the race for life—is, without question, one +of the individuals best fitted to do so; that is to say, best fitted to +the conditions of its existence considered as a whole. Nature is, +therefore, always picking out, or selecting, such individuals to live +and to breed.</p> + +<p>The third fact is, that the individuals so selected transmit their +favourable qualities to their offspring by heredity. There is no doubt +about this fact, so far as we are concerned with it. For although, as I +have already hinted, considerable doubt has of late years been cast upon +Lamarck’s doctrine of the hereditary transmission of <i>acquired</i> +characters, it remains as impossible as ever it was to question the +<span class="pagenum"><a name="page_263" id="page_263">[263]</a></span> +hereditary transmission of what are called <i>congenital</i> characters. And +this is all that Darwin’s theory necessarily requires.</p> + +<p>The fourth fact is, that although heredity as a whole produces a +wonderfully exact copy of the parent in the child, there is never a +precise reduplication. Of all the millions of human beings upon the face +of the earth, no one is so like another that we cannot see some +difference; the resemblance is everywhere specific, nowhere individual. +Now this same remark applies to all specific types. The only reason why +we notice individual differences in the case of the human type more than +we do in the case of any other types, is because our attention is here +more incessantly focussed upon these differences. We are compelled to +notice them in the case of our own species, however small they may +appear to a naturalist, because, unless we do so, we should not +recognise the members of our own family, or be able to distinguish +between a man whom we know is ready to do us an important service, and +another man whom we know is ready to cut our throats. But our common +mother Nature is able thus to distinguish between all her children. Her +eyes are much more ready to detect small individual peculiarities than +are the eyes of any naturalist. No slight variations in the cast of +feature or disposition of parts, no minute difference in the arrangement +of microscopical cells, can escape her ever vigilant attention. And, +consequently, when among all the innumerable multitudes of individual +variations any one arises which—no matter in how slight a degree—gives +to that individual a better chance of success in the struggle for life, +Nature chooses that individual <span class="pagenum"><a name="page_264" id="page_264">[264]</a></span> +to survive, and so to perpetuate the improvement in his or her progeny.</p> + +<p>Now I say that all these several component parts of Darwinian doctrine +are not matters of theory, but matters of fact. The only element of +theory in his doctrine of evolution by natural selection has reference +to the degree in which these observable facts, when thus brought +together, are adequate to account for the process of evolution.</p> + +<hr class='minor' /> + +<p>So much, then, as a statement of the theory of natural selection. But +from this statement—i. e. from the theory of natural selection +itself—there follow certain matters of general principle which it is +important to bear in mind. These, therefore, I shall here proceed to +mention.</p> + +<p>First of all, it is evident that the theory is applicable as an +explanation of organic changes in specific types only in so far as these +changes are of <i>use</i>, or so far as such changes endow the species with +better chances of success in the general struggle for existence. This is +the only sense in which I shall always employ the terms use, utility, +service, benefit, and so forth—that is to say, in the sense of +life-preserving.</p> + +<hr class='minor' /> + +<p>Next, it must be clearly understood that the life which it is the +object, so to speak, of natural selection to preserve, is primarily the +life of the <i>species</i>; not that of the <i>individual</i>. Natural selection +preserves the life of the individual only in so far as this is conducive +to that of the species. Wherever the life-interests of the individual +clash with those of the species, that individual is sacrificed in favour +of others <span class="pagenum"><a name="page_265" id="page_265">[265]</a></span> +who happen better to subserve the interests of the species. For +example, in all organisms a greater or less amount of vigour is wasted, +so far as individual interests are concerned, in the formation and the +nourishment of progeny. In the great majority of plants and animals an +enormous amount of physiological energy is thus expended. Look at the +roe or the milt of a herring, for instance, and see what a huge drain +has been made upon the individual for the sake of its species. Again, +all unselfish instincts have been developed for the sake of the species, +and usually against the interests of the individual. An ant which will +allow her head to be slowly drawn from her body rather than relinquish +her hold upon a pupa, is clearly acting in response to an instinct which +has been developed for the benefit of the hive, though fatal to the +individual. And, in a lesser degree, the parental instincts, wherever +they occur, are more or less detrimental to the interests of the +individual, though correspondingly essential to those of the race.</p> + +<p>These illustrations will serve to show that natural selection always +works primarily for the life-interests of the species—and, indeed, only +works for those of the individual at all in so far as the latter happen +to coincide with the former. Or, otherwise stated, the object of natural +selection is always that of producing and maintaining specific types in +the highest degree of efficiency, no matter what may become of the +constituent individuals. Which is a striking republication by Science of +a general truth previously stated by Poetry:—</p> + +<div class="poem"><div class="stanza"> +<span class="i0">So careful of the type she seems,<br /></span> +<span class="i0">So careless of the single life.<br /></span> +</div></div><p><span class="pagenum"><a name="page_266" id="page_266">[266]</a></span></p> + +<p class='noin'>Tennyson thus noted the fact, and a few years later Darwin supplied the +explanation.</p> + +<p>But of course in many, if not in the majority of cases, anything that +adds to the life-sustaining power of the single life thereby ministers +also to the life-sustaining power of the type; and thus we can +understand why all mechanisms and instincts which minister to the single +life have been developed—namely, because the life of the species is +made up of the lives of all its constituent individuals. It is only +where the interests of the one clash with those of the other that +natural selection works against the individual. So long as the interests +are coincident, it works in favour of both.</p> + +<p>Natural selection, then, is a theory which seeks to explain by natural +causes the occurrence of every kind of adaptation which is to be met +with in organic nature, on the assumption that adaptations of every kind +have primary reference to the preservation of species, and therefore +also, as a general rule, to the preservation of their constituent +individuals. And from this it follows that where it is for the benefit +of a species to change its type, natural selection will effect that +change, thus leading to a specific transmutation, or the evolution of a +new species. In such cases the old species may or may not become +extinct. If the transmutation affects the species as a whole, or +throughout its entire range, of course <i>that</i> particular type becomes +extinct, although it does so by becoming changed into a still more +suitable type in the course of successive generations. If, on the other +hand, the transmutation affects only a part of the original species, or +not throughout its entire range, then the <span class="pagenum"><a name="page_267" id="page_267">[267]</a></span> +other parts of that species may survive for any number of ages as they +originally were. In the one case there is a ladder-like transmutation of +species in time; in the other case a possibly tree-like multiplication +of species in space. But whether the evolution of species be thus serial +in time or divergent in space, the object of natural selection, so to +speak, is in either case the same—namely, that of preserving all +types which prove best suited to the conditions of their existence.</p> + +<hr class='minor' /> + +<p>Once more, the term “struggle for existence” must be +understood to comprehend, not only a competition for life among +contemporary individuals of the same species, but likewise a struggle by +all such individuals taken collectively for the continuance of their own +specific type. Thus, on the one hand, while there is a perpetual civil +war being waged between members of the same species, on the other hand +there is a foreign war being waged by the species as a whole against its +world as a whole. Hence it follows that natural selection does not +secure survival of the fittest as regards individuals only, but also +survival of the fittest as regards types. This is a most important point +to remember, because, as a general rule, these two different causes +produce exactly opposite effects. Success in the civil war, where each +is fighting against all, is determined by <i>individual</i> fitness and +<i>self-reliance</i>. But success in the foreign war is determined by what +may be termed <i>tribal</i> fitness and <i>mutual dependence</i>. For example, +among social insects the struggle for existence is quite as great +between different tribes or communities, as it is between different +individuals of <span class="pagenum"><a name="page_268" id="page_268">[268]</a></span> +the same community; and thus we can understand the extraordinary degree +in which not only co-operative instincts, but also largely intelligent +social habits, have here been developed<a name="FNanchor_30_30" id="FNanchor_30_30"></a><a href="#Footnote_30_30" class="fnanchor">[30]</a>. +Similarly, in the case of mankind, we can understand the still more +extraordinary development of these things—culminating in the moral +sense. I have heard a sermon, preached at one of the meetings of the +British Association, entirely devoted to arguing that the moral sense +could not have been evolved by natural selection, seeing that the +altruism which this sense involves is the very opposite of selfishness, +which alone ought to have been the product of survival of the fittest in +a struggle for life. And, of course, this argument would have been +perfectly sound had Darwin limited the struggle for existence to +individuals, without extending it to communities. But if the preacher +had ever read Darwin’s works he would have found that, when thus +extended, the principle of natural selection is bound to work in favour +of the co-operative instincts in the case of so highly social an animal +as man; and that of these instincts conscience is the highest imaginable +exhibition.</p> + +<p>What I have called tribal fitness—in contradistinction to individual +fitness—begins with the family, developes in the community (herd, hive, +clan, &c.), and usually ends with the limits of the species. On the one +hand, however, it is but seldom that it extends so far as to embrace the +entire species; while, on the other hand, it may in some cases, and as +it were <span class="pagenum"><a name="page_269" id="page_269">[269]</a></span> +sporadically, extend beyond the species. In these latter cases members +of different species mutually assist one another, whether in the way of +what is called symbiosis, or in a variety of other ways which I need not +wait to mention. For the only point which I now desire to make clear is, +that all cases of mutual aid or co-operation, whether within or beyond +the limits of species, are cases which fall under the explanatory sweep +of the Darwinian theory<a name="FNanchor_31_31" id="FNanchor_31_31"></a><a href="#Footnote_31_31" class="fnanchor">[31]</a>.</p> + +<hr class='minor' /> + +<p>Another important point to notice is, that it constitutes no part of the +theory of natural selection to suppose that survival of the fittest must +invariably lead to <i>improvement</i> of type, in the sense of superior +organization. On the contrary, if from change of habits or conditions of +life an organic type ceases to have any use for previously useful +organs, natural selection will not only allow these organs in successive +generations to deteriorate—by no longer placing any selective premium +upon their maintenance—but may even proceed to assist the agencies +engaged in their destruction. For, being now useless, they may become +even deleterious, by absorbing nutriment, causing weight, occupying +space, &c., without conferring any compensating benefit. Thus we can +understand why it is that parasites, for example, present the phenomena +of what is called <i>degeneration</i>, i. e. showing by their whole structure +that they have descended from a possibly very much higher type of +organization than that which they now exhibit. Having for innumerable +<span class="pagenum"><a name="page_270" id="page_270">[270]</a></span> +generations ceased to require their legs, their eyes, and so forth, all +such organs of high elaboration have either disappeared or become +vestigial, leaving the parasite as a more or less effete representative +of its ancestry.</p> + +<p>These facts of degeneration, as we have previously seen, are of very +general occurrence, and it is evident that their importance in the field +of organic evolution as a whole has been very great. Moreover, it ought +to be particularly observed that, as just indicated, the facts may be +due either to a passive <i>cessation</i> of selection, or to an active +<i>reversal</i> of it. Or, more correctly, these facts are probably <i>always</i> +due to the cessation of selection, although in most cases where species +in a state of nature are concerned, the process of degeneration has been +both hastened and intensified by the super-added influence of the +reversal of selection. In the next volume I shall have occasion to recur +to this distinction, when it will be seen that it is one of no small +importance to the general theory of descent.</p> + +<hr class='minor' /> + +<p>We may now proceed to consider certain misconceptions of the Darwinian +theory which are largely, not to say generally, prevalent among +supporters of the theory. These misconceptions, therefore, differ from +those which fall to be considered in the next chapter, i. e. +misconceptions which constitute grounds of objection to the theory.</p> + +<hr class='minor' /> + +<p>Of all the errors connected with the theory of natural selection, +perhaps the one most frequently met with—especially among supporters of +the theory—is that of employing the theory to explain all cases of +<span class="pagenum"><a name="page_271" id="page_271">[271]</a></span> +Phyletic modification (or inherited change of type) indiscriminately, +without waiting to consider whether in particular cases its application +is so much as logically possible. The term “natural +selection” thus becomes a magic word, or sesame, at the utterance +of which every closed door is supposed to be immediately opened. Be it +observed, I am not here alluding to that merely blind faith in natural +selection, which of late years has begun dogmatically to force this +principle as the sole cause of organic evolution in every case where it +is <i>logically possible</i> that the principle can have come into play. Such +a blind faith, indeed, I hold to be highly inimical, not only to the +progress of biological science, but even to the true interests of the +natural selection theory itself. As to this I shall have a good deal to +say in the next volume. Here, however, the point is, that the theory in +question is often invoked in cases where it is not even logically +possible that it can apply, and therefore in cases where its application +betokens, not merely an error of judgment or extravagance of dogmatism, +but a fallacy of reasoning in the nature of a logical contradiction. +almost any number of examples might be given; but one will suffice to +illustrate what is meant. And I choose it from the writings of one of +the authors of the selection theory itself, in order to show how easy it +is to be cheated by this mere juggling with a phrase—for of course +I do not doubt that a moment’s thought would have shown the writer +the untenability of his statement.</p> + +<p>In his most recent work Mr. Wallace advances an interesting hypothesis +to the effect that differences of colour between allied species, which +are apparently <span class="pagenum"><a name="page_272" id="page_272">[272]</a></span> +too slight to serve any other purpose, may act as “recognition +marks,” whereby the opposite sexes are enabled at once to +distinguish between members of their own and of closely resembling +species. Of course this hypothesis can only apply to the higher animals; +but the point here is that, supposing it to hold for them, Mr. Wallace +proceeds to argue thus:—Recognition marks “have in all +probability been acquired in the process of differentiation for the +purpose of checking the intercrossing of allied forms,” because +“one of the first needs of a new species would be to keep separate +from its nearest allies, and this could be more readily done by some +easily seen external mark<a name="FNanchor_32_32" id="FNanchor_32_32"></a><a href="#Footnote_32_32" class="fnanchor">[32]</a>.” +Now, it is clearly not so much as logically possible that these +recognition-marks (supposing them to be such) can have been acquired by +natural selection, “for the purpose of checking intercrossing of +allied forms.” For the theory of natural selection, from its own +essential nature as a theory, is logically exclusive of the supposition +that survival of the fittest ever provides changes in anticipation of +future uses. Or, otherwise stated, it involves a contradiction of the +theory itself to say that the colour-changes in question were originated +by natural selection, in order to meet “one of the <i>first</i> needs +of a <i>new</i> species,” or for the purpose of <i>subsequently</i> +preventing intercrossing with allied forms. If it had been said that +these colour-differentiations were originated by some cause other than +natural selection (or, if by natural selection, still with regard to +some <i>previous</i>, instead of <i>prophetic</i>, “purpose"), and, when so +“acquired,” <i>then</i> began to serve the “purpose” +assigned, <span class="pagenum"><a name="page_273" id="page_273">[273]</a></span> +the argument would not have involved the fallacy which we are now +considering. But, as it stands, the argument reverts to the teleology of +pre-Darwinian days—or the hypothesis of a “purpose” in +the literal sense which sees the end from the beginning, instead of a +“purpose” in the metaphorical sense of an adaptation that is +evolved by the very modifications which subserve it<a name="FNanchor_33_33" id="FNanchor_33_33"></a><a href="#Footnote_33_33" class="fnanchor">[33]</a>.</p> + +<hr class='minor' /> + +<p>Another very prevalent, and more deliberate, fallacy connected with the +theory of natural selection is, <i>that it follows deductively from the +theory itself</i> that the principle of natural selection must be the sole +means of modification in all cases where modification is of an +<i>adaptive</i> kind,—with the consequence that no other principle can ever +have been concerned in the production of structures or instincts which +are of any use to their possessors. Whether or not natural selection +actually has been the sole means of adaptive modification in the race, +as distinguished from the individual, is a question of biological +fact<a name="FNanchor_34_34" id="FNanchor_34_34"></a><a href="#Footnote_34_34" class="fnanchor">[34]</a>; but it <span class="pagenum"><a name="page_274" id="page_274">[274]</a></span>involves a grave error of reasoning to suppose that +this question can be answered deductively from the theory of natural +selection itself, as I shall show at some length in the next volume.</p> + +<hr class='minor' /> + +<p>A still more extravagant, and a still more unaccountable fallacy is the +one which represents it as following deductively from the theory of +natural selection itself, that all <i>hereditary</i> characters are +“necessarily” due to natural selection. In other words, not +only all adaptive, but likewise all non-adaptive hereditary characters, +it is said, <i>must</i> be due to natural selection. For non-adaptive +characters are taken to be due to “correlation of growth,” +in connexion with some of the adaptive ones—natural selection being +thus the <i>indirect</i> means of producing the former <i>wherever</i> they may +occur, on account of its being the <i>direct</i> and the <i>only</i> means of +producing the latter. Thus it is deduced from the theory of natural +selection itself,—1st, that the principle of natural selection is the +only possible cause of adaptive modification: 2nd, that non-adaptive +modifications can only occur in the race as correlated appendages to the +adaptive: 3rd, that, consequently, natural selection is the only +possible cause of modification, whether adaptive or non-adaptive. Here +again, therefore, we must observe that none of these sweeping +generalizations can possibly be justified by deductive reasoning from +the theory of natural selection itself. Any attempt at such deductive +reasoning must necessarily end in circular reasoning, as I shall +likewise show in the <span class="pagenum"><a name="page_275" id="page_275">[275]</a></span> +Second volume, where this whole “question of utility” will +be thoroughly dealt with.</p> + +<hr class='minor' /> + +<p>Once more, there is an important oversight very generally committed by +the followers of Darwin. For even those who avoid the fallacies above +mentioned often fail to perceive, that natural selection can only begin +to operate if the <i>degree</i> of adaptation is already given as +sufficiently high to count for something in the struggle for +<i>existence</i>. Any adaptations which fall below this level of importance +cannot possibly have been produced by survival of the fittest. Yet the +followers of Darwin habitually speak of adaptative characters, which <i>in +their own opinion</i> are subservient merely to comfort or convenience, as +having been produced by such means. Clearly this is illogical; for it +belongs to the essence of Darwin’s theory to suppose, that natural +selection can have no jurisdiction beyond the line where structures or +instincts already present a sufficient degree of adaptational value to +increase, in some measure, the expectation of life on the part of their +possessors. We cannot speak of adaptations as due to natural selection, +without thereby affirming that they present what I have elsewhere termed +a “selection value.”</p> + +<hr class='minor' /> + +<p>Lastly, as a mere matter of logical definition, it is well-nigh +self-evident that the theory of natural selection is a theory of the +origin, and cumulative development, of <i>adaptations</i>, whether these be +distinctive of species, or of genera, orders, families, classes, and +sub-kingdoms. It is only when the adaptations happen to be distinctive +of the first (or lowest) of these <span class="pagenum"><a name="page_276" id="page_276">[276]</a></span> +taxonomic divisions, that the theory which accounts for <i>these</i> +adaptations accounts also for the forms which present them,—i. e. +becomes <i>also</i> a theory of the origin of species. This, however, is +clearly but an accident of particular cases; and, therefore, even in +them the theory is <i>primarily</i> a theory of adaptations, while it is but +secondarily a theory of the species which present them. Or, otherwise +stated, the theory is no more a theory of the origin of species than it +is of the origin of genera, families, and the rest; while, on the other +hand, it is <i>everywhere</i> a theory of the adaptive modifications whereby +each of these taxonomic divisions has been differentiated as such. Yet, +sufficiently obvious as the accuracy of this definition must appear to +any one who dispassionately considers it, several naturalists of high +standing have denounced it in violent terms. I shall therefore have to +recur to the subject at somewhat greater length hereafter. At present it +is enough merely to mention the matter, as furnishing another and a +curious illustration of the not infrequent weakness of logical +perception on the part of minds well gifted with the faculty of +observation. It may be added, however, that the definition in question +is in no way hostile to the one which is virtually given by Darwin in +the title of his great work. <i>The Origin of Species by means of Natural +Selection</i> is beyond doubt the best title that could have been given, +because at the time when the work was published the <i>fact</i>, no less than +the <i>method</i>, of organic evolution had to be established; and hence the +most important thing to be done at that time was to prove the +transmutation of species. But now that this has been done to the +satisfaction of naturalists in general, it is <span class="pagenum"><a name="page_277" id="page_277">[277]</a></span> +as I have said, curious to find some of them denouncing a wider +definition of the principle of natural selection, merely because the +narrower (or included) definition is invested with the charm of verbal +associations<a name="FNanchor_35_35" id="FNanchor_35_35"></a><a href="#Footnote_35_35" class="fnanchor">[35]</a>.</p> + +<hr class='minor' /> + +<p>So much for fallacies and misconceptions touching Darwin’s theory, +which are but too frequently met with in the writings of its supporters. +We must now pass on to mention some of the still greater fallacies and +misconceptions which are prevalent in the writings of its opponents. +And, in order to do this thoroughly, I shall begin by devoting the +remainder of the present chapter to a consideration of the antecedent +standing of the two theories of natural selection and supernatural +design. This having been done, in the succeeding chapters I shall deal +with the evidences for, and the objections against, the former theory.</p> + +<hr class='minor' /> + +<p>Beginning, then, with the antecedent standing of these alternative +theories, the first thing to be noticed is, that they are both concerned +with the same subject-matter, which it is their common object to +explain. Moreover, this subject-matter is clearly and sharply divisible +into two great classes of facts in organic nature—namely, those of +Adaptation and those of Beauty. Darwin’s theory of descent +explains the former by his doctrine of natural selection, and the latter +by his doctrine of sexual selection. In the first instance, therefore, I +shall have to deal only with the facts of <span class="pagenum"><a name="page_278" id="page_278">[278]</a></span> +adaptation, leaving for subsequent consideration the facts of beauty.</p> + +<p>Innumerable cases of the adaptation of organisms to their surroundings +being the facts which now stand before us to be explained either by +natural selection or by supernatural intention, we may first consider a +statement which is frequently met with—namely, that even if all such +cases of adaptation were proved to be fully explicable by the theory of +descent, this would constitute no disproof of the theory of design: all +the cases of adaptation, it is argued, might still be due to design, +even though they admit of being hypothetically accounted for by the +theory of descent. I have heard an eminent Professor tell his class that +the many instances of mechanical adaptation discovered and described by +Darwin as occurring in orchids, seemed to him to furnish better proof of +supernatural contrivance than of natural causes; and another eminent +Professor has informed me that, although he had read the <i>Origin of +Species</i> with care, he could see in it no evidence of natural selection +which might not equally well have been adduced in favour of intelligent +design. But here we meet with a radical misconception of the whole +logical attitude of science. For, be it observed, this exception <i>in +limine</i> to the evidence which we are about to consider does not question +that natural selection <i>may</i> be able to do all that Darwin ascribes to +it. The objection is urged against his interpretation of the facts +merely on the ground that these facts might <i>equally well</i> be ascribed +to intelligent design. And so undoubtedly they might, if we were all +simple enough to adopt a supernatural explanation whenever a natural one +is found sufficient to <span class="pagenum"><a name="page_279" id="page_279">[279]</a></span> +account for the facts. Once admit the irrational principle that we may +assume the operation of higher causes where the operation of lower ones +is sufficient to explain the observed phenomena, and all our science and +all our philosophy are scattered to the winds. For the law of logic +which Sir William Hamilton called the law of parsimony—or the law +which forbids us to assume the operation of higher causes when lower +ones are found sufficient to explain the observed effects—this law +constitutes the only barrier between science and superstition. It is +always possible to give a hypothetical explanation of any phenomenon +whatsoever, by referring it immediately to the intelligence of some +supernatural agent; so that the only difference between the logic of +science and the logic of superstition consists in science recognising a +validity in the law of parsimony which superstition disregards. +Therefore one can have no hesitation in saying that this way of looking +at the evidence in favour of natural selection is not a scientific or a +reasonable way of looking at it, but a purely superstitious way. Let us +take, as an illustration, a perfectly parallel case. When Kepler was +unable to explain by any known causes the paths described by the +planets, he resorted to a supernatural explanation, and supposed that +every planet was guided in its movements by some presiding angel. But +when Newton supplied a beautifully simple physical explanation, all +persons with a scientific habit of mind at once abandoned the +metaphysical one. Now, to be consistent, the above-mentioned Professors, +and all who think with them, ought still to adhere to Kepler’s +hypothesis in preference to Newton’s explanation; for, excepting +the law of parsimony, there is certainly <span class="pagenum"><a name="page_280" id="page_280">[280]</a></span> +no other logical objection to the statement, that the movements of the +planets afford as good evidence of the influence of guiding angels as +they do of the influence of gravitation.</p> + +<p>So much, then, for the illogical position that, granting the evidence in +favour of natural descent and supernatural design to be equal and +parallel, we should hesitate in our choice between the two theories. +But, of course, if the evidence is supposed <i>not</i> to be equal and +parallel—i. e. if it is supposed that the theory of natural selection +is not so good a theory whereby to explain the facts of adaptation as is +that of supernatural design,—then the objection is no longer the one +which we are considering. It is quite another objection, and one which +is not <i>prima facie</i> absurd. Therefore let us state clearly the distinct +question which thus arises.</p> + +<p>Innumerable cases of adaptation of organisms to their environments are +the observed facts for which an explanation is required. To supply this +explanation, two, and only two, hypotheses are in the field. Of these +two hypotheses one is intelligent design manifested directly in special +creation; the other is natural causation operating through countless +ages of the past. Now, the adaptations in question involve an +innumerable multitude of special mechanisms, in most cases even within +the limits of any one given species; but when we consider the sum of all +these mechanisms presented by organic nature as a whole, the mind must +indeed be dull which does not feel astounded. For, be it further +observed, these mechanical contrivances<a name="FNanchor_36_36" id="FNanchor_36_36"></a><a href="#Footnote_36_36" class="fnanchor">[36]</a> +are, for the most part, no merely simple <span class="pagenum"><a name="page_281" id="page_281">[281]</a></span> +arrangements, which might reasonably be supposed due, like the +phenomena of crystallization, to comparatively simple physical causes. +On the contrary, they everywhere and habitually exhibit so deep-laid, so +intricate, and often so remote an adaptation of means to ends, that no +machinery of human contrivance can properly be said to equal their +perfection from a mechanical point of view. Therefore, without question, +the hypothesis which first of all they suggest—or suggest most +readily—is the hypothesis of design. And this hypothesis becomes +virtually the only hypothesis possible, if it be assumed—as it +generally was assumed by natural theologians of the past,—that all +species of plants and animals were introduced into the world <i>suddenly</i>. +For it is quite inconceivable that any known cause, other than +intelligent design, could be competent to turn out instantaneously any +one of these intricate pieces of machinery, already adapted to the +performance of its special function. But, on the other hand, if there is +any evidence to show that one species becomes slowly transformed into +another—or that one set of adaptations becomes slowly changed into +another set as changing circumstances require,—then it becomes +quite possible to imagine that a strictly natural causation may have had +something to do with the matter. And this suggestion becomes greatly +more probable when we discover, from geological evidence and +embryological research, that in the history both of races and of +individuals the <span class="pagenum"><a name="page_282" id="page_282">[282]</a></span> +various mechanisms in question have themselves had a +history—beginning in the forms of most uniformity and simplicity, +gradually advancing to forms more varied and complex, nowhere exhibiting +any interruptions in their upward progress, until the world of organic +machinery as we now have it is seen to have been but the last phase of a +long and gradual growth, the ultimate roots of which are to be found in +the soil of undifferentiated protoplasm.</p> + +<p>Lastly, when there is supplied to us the suggestion of natural selection +as a cause presumably adequate to account for this continuous growth in +the number, the intricacy, and the perfection of such mechanisms, it is +only the most unphilosophical mind that can refuse to pause as between +the older hypothesis of design and the newer hypothesis of descent.</p> + +<p>Thus it is clear that the <i>a priori</i> standing of the rival hypotheses of +naturalism and supernaturalism in the case of all these pieces of +organic machinery, is profoundly affected by the question whether they +came into existence suddenly, or whether they did so gradually. For, if +they all came into existence suddenly, the fact would constitute +well-nigh positive proof in favour of supernaturalism, or creation by +design; whereas, if they all came into existence gradually, this fact +would in itself constitute presumptive evidence in favour of naturalism, +or of development by natural causes. And, as shown in the previous +chapters, the proof that all species of plants and animals came into +existence gradually—or the proof of evolution as a fact—is simply +overwhelming.</p> + +<p>From a still more general point of view I may state the case in another +way, by borrowing and somewhat <span class="pagenum"><a name="page_283" id="page_283">[283]</a></span> +expanding an illustration which, I believe, was first used by Professor +Huxley. If, when the tide is out, we see lying upon the shore a long +line of detached sea-weed, marking the level which is reached by full +tide, we should be free to conclude that the separation of the sea-weed +from the sand and the stones was due to the intelligent work of some one +who intended to collect the sea-weed for manure, or for any other +purpose. But, on the other hand, we might explain the fact by a purely +physical cause—namely, the separation by the sea-waves of the +sea-weed from the sand and stones, in virtue of its lower specific +gravity. Now, thus far the fact would be explained equally well by +either hypothesis; and this fact would be the fact of <i>selection</i>. But +whether we yielded our assent to the one explanation or to the other +would depend upon a due consideration of all collateral circumstances. +The sea-weed might not be of a kind that is of any use to man; there +might be too great a quantity of it to admit of our supposing that it +had been collected by man; the fact that it was all deposited on the +high-water-mark would in itself be highly suggestive of the agency of +the sea; and so forth. Thus, in such a case any reasonable observer +would decide in favour of the physical explanation, or against the +teleological one.</p> + +<p>Now the question whether organic evolution has been caused by physical +agencies or by intelligent design is in precisely the same predicament. +There can be no logical doubt that, theoretically at all events, the +physical agencies which the present chapter is concerned with, and which +are conveniently summed up in the term natural selection, are as +competent to produce these so-called mechanical contrivances, and the +other <span class="pagenum"><a name="page_284" id="page_284">[284]</a></span> +cases of adaptation which are to be met with in organic nature, as +intelligent design could be. Hence, our choice as between these two +hypotheses must be governed by a study of all collateral circumstances; +that is to say, by a study of the evidences in favour of the physical +explanation. To this study, therefore, we shall now address ourselves, +in the course of the following chapters.</p> + +<hr /><p class="pagenum"><a name="page_285" id="page_285">[285]</a></p> +<h2><a name="CHAPTER_VIII" id="CHAPTER_VIII"></a>CHAPTER VIII.<br /><br /> +<span class="sc">Evidences of the Theory of Natural Selection.</span></h2> + +<p>I will now proceed to state the main arguments in favour of the theory +of natural selection, and then, in the following chapter, the main +objections which have been urged against it.</p> + +<p>In my opinion, the main arguments in favour of the theory are three in +number.</p> + +<p>First, it is a matter of observation that the struggle for existence in +nature does lead to the extermination of forms less fitted for the +struggle, and thus makes room for forms more fitted. This general fact +may be best observed in cases where an exotic species proves itself +better fitted to inhabit a new country than is some endemic species +which it exterminates. In Great Britain, for example, the so-called +common rat is a comparatively recent importation from Norway, and it has +so completely supplanted the original British rat, that it is now +extremely difficult to procure a single specimen of the latter: the +native black rat has been all but exterminated by the foreign brown rat. +The same thing is constantly found in the case of imported species of +plants. I have seen the river at Cambridge so choked with the inordinate +propagation of a species <span class="pagenum"><a name="page_286" id="page_286">[286]</a></span> +Of water-weed which had been introduced from america, that considerable +expense had to be incurred in order to clear the river for traffic. In +new zealand the same thing has happened with the european water-cress, +and in australia with the common rabbit. So it is doubtless true, as one +of the natives is said to have philosophically remarked, “the +white man’s rat has driven away our rat, the european fly drives +away our fly, his clover kills our grass, and so will the maoris +disappear before the white man himself.” innumerable other cases +to the same effect might be quoted; and they all go to establish the +fact that forms less fitted to survive succumb in their competition with +forms better fitted.</p> + +<hr class='minor' /> + +<p>Secondly, there is a general consideration of the largest possible +significance in the present connexion—namely, that among all the +millions of structures and instincts which are so invariably, and for +the most part so wonderfully, adapted to the needs of the species +presenting them, we cannot find a single instance, either in the +vegetable or animal kingdom, of a structure or an instinct which is +developed for the exclusive benefit of another species. Now this great +and general fact is to my mind a fact of the most enormous, not to say +overwhelming, significance. The theory of natural selection has now been +before the world for more than thirty years, and during that time it had +stood a fire of criticism such as was never encountered by any +scientific theory before. From the first Darwin invited this criticism +to adduce any single instance, either in the vegetable or animal +kingdom, of a structure or an instinct which should unquestionably +<span class="pagenum"><a name="page_287" id="page_287">[287]</a></span> +be proved to be of exclusive use to any species other than the one +presenting it. He even went so far as to say that if any one such +instance could be shown he would surrender his whole theory on the +strength of it—so assured had he become, by his own prolonged +researches, that natural selection was the true agent in the production +of adaptive structures, and, as such, could never have permitted such a +structure to occur in one species for the benefit of another. Now, as +this invitation has been before the world for so many years, and has not +yet been answered by any naturalist, we may by this time be pretty +confident that it never will be answered. How tremendous, then, is the +significance of this fact in its testimony to Darwin’s theory! The +number of animal and vegetable species, both living and extinct, is to +be reckoned by millions, and every one of these species presents on an +average hundreds of adaptive structures,—at least one of which in +many, possibly in most, if not actually in all cases, is peculiar to the +species that presents it. In other words, there are millions of adaptive +structures (not to speak of instincts) which are peculiar to the species +presenting them, and also many more which are the common property of +allied species: yet, notwithstanding this inconceivable profusion of +adaptive structures in organic nature, there is no single instance that +has been pointed out of the occurrence of such a structure save for the +benefit of the species that presents it. Therefore, I say that this +immensely large and general fact speaks with literally immeasurable +force in favour of natural selection, as at all events one of the main +causes of organic evolution. For the fact is precisely what we should +expect if this theory is true, while <span class="pagenum"><a name="page_288" id="page_288">[288]</a></span> +upon no other theory can its universality and invariability be rendered +intelligible. On the beneficent design theory, for instance, it is +inexplicable that no species should ever be found to present a structure +or an instinct having primary reference to the welfare of another +species, when, <i>ex hypothesi</i>, such an endless amount of thought has +been displayed in the creation of structures and instincts having +primary reference to the species which present them. For how magnificent +a display of divine beneficence would organic nature have afforded, if +all—or even some—species had been so inter-related as to +have ministered to each others wants. Organic species might then have +been likened to a countless multitude of voices, all singing in one +great harmonious psalm. But, as it is, we see absolutely no vestige of +such co-ordination: every species is for itself, and for itself +alone—an outcome of the always and everywhere fiercely raging +struggle for life.</p> + +<p>In order that the force of this argument may not be misapprehended, it +is necessary to bear in mind that it is in no way affected by cases +where a structure or an instinct is of primary benefit to its possessor, +and then becomes of secondary benefit to some other species on account +of the latter being able in some way or another to utilise its action. +Of course organic nature is full of cases of this kind; but they only go +to show the readiness which all species display to utilise for +themselves everything that can be turned to good account in their own +environments, and so, among other things, the structures and instincts +of other animals. For instance, it would be no answer to Darwin’s +challenge if any one were to point to a hermit-crab inhabiting the +<span class="pagenum"><a name="page_289" id="page_289">[289]</a></span> +cast-off shell of a mollusk; because the shell was primarily of use to +the mollusk itself, and, so far as the mollusk is concerned, the fact of +its shell being afterwards of a secondary use to the crab is quite +immaterial. What Darwin’s challenge requires is, that some +structure or instinct should be shown which is not merely of such +secondary or accidental benefit to another species, but clearly adapted +to the needs of that other species in the first instance—such, for +example, as would be the case if the tail of a rattle-snake were of no +use to its possessor, while serving to warn other animals of the +proximity of a dangerous creature; or, in the case of instincts, if it +were true that a pilot-fish accompanies a shark for the purpose of +helping the shark to discover food. Both these instances have been +alleged; but both have been shown untenable. And so it has proved of all +the other cases which thus far have been put forward.</p> + +<p>Perhaps the most remarkable of all the allegations which ever have been +put forward in this connexion are those that were current with regard to +instincts before the publication of Darwin’s work. These +allegations are the most remarkable, because they serve to show, in a +degree which I do not believe could be shown anywhere else, the warping +power of preconceived ideas. A short time ago I happened to come across +the 8th edition of the <i>Encyclopædia Britannica</i>, and turned up the +article on “Instinct” there, in order to see what amount of +change had been wrought with regard to our views on this subject by the +work of Darwin—the 8th edition of the <i>Encyclopædia Britannica</i> having +been published shortly before <i>The Origin of Species by means of Natural +Selection</i>.<span class="pagenum"><a name="page_290" id="page_290">[290]</a></span> +I cannot wait to give any lengthy quotations from this representative +exponent of scientific opinion upon the subject at that time; but its +general drift may be appreciated if I transcribe merely the short +concluding paragraph, wherein he sums up his general results. Here he +says:—</p> + +<div class='blockquot'><p>It thus only remains for us to regard instinct as a mental faculty, +<i>sui generis</i>, the gift of God to the lower animals, that man in +his own person, and by them, might be relieved from the meanest +drudgery of nature.</p></div> + +<p>Now, here we have the most extraordinary illustration that is imaginable +of the obscuring influence of a preconceived idea. Because he started +with the belief that instincts <i>must</i> have been implanted in animals for +the benefit of man, this writer, even when writing a purely scientific +essay, was completely blinded to the largest, the most obvious, and the +most important of the facts which the phenomena of instinct display. +For, as a matter of fact, among all the many thousands of instincts +which are known to occur in animals, there is no single one that can be +pointed to as having any special reference to man; while, on the other +hand, it is equally impossible to point to one which does not refer to +the welfare of the animal presenting it. Indeed, when the point is +suggested, it seems to me surprising how few in number are the instincts +of animals which have proved to be so much as of secondary or accidental +benefit to man, in the same way as skins, furs, and a whole host of +other animal products are thus of secondary use to him. Therefore, this +writer not only failed to perceive the most obvious truth that every +instinct, without any single exception, has reference to the animal +which <span class="pagenum"><a name="page_291" id="page_291">[291]</a></span> +presents it; but he also conceived a purely fictitious inversion of +this truth, and wrote an essay to prove a statement which all the +instincts in the animal kingdom unite in contradicting.</p> + +<p>This example will serve to show, in a striking manner, not only the +distance that we have travelled in our interpretation of organic nature +between two successive editions of the <i>Encyclopædia Britannica</i>, but +also the amount of verification which this fact furnishes to the theory +of natural selection. For, inasmuch as it belongs to the very essence of +this theory that all adaptive characters (whether instinctive or +structural) must have reference to their own possessors, we find +overpowering verification furnished to the theory by the fact now before +us—namely, that immediately prior to the enunciation of this theory, +the truth that all adaptive characters have reference only to the +species which present them was not perceived. In other words, it was the +testing of this theory by the facts of nature that <i>revealed</i> to +naturalists the general law which the theory, as it were, predicted—the +general law that all adaptive characters have primary reference to the +species which present them. And when we remember that this is a kind of +verification which is furnished by millions of separate cases, the whole +mass of it taken together is, as I have before said, overwhelming.</p> + +<p>It is somewhat remarkable that the enormous importance of this argument +in favour of natural selection as a prime factor of organic evolution +has not received the attention which it deserves. Even Darwin himself, +with his characteristic reserve, has not presented its incalculable +significance; nor do I <span class="pagenum"><a name="page_292" id="page_292">[292]</a></span> +know any of his followers who have made any approach to an adequate use +of it in their advocacy of his views. In preparing the present chapter, +therefore, I have been particularly careful not to pitch too high my own +estimate of its evidential value. That is to say, I have considered, +both in the domain of structures and of instincts, what instances admit +of being possibly adduced <i>per contra</i>, or as standing outside the +general law that adaptive structures and instincts are of primary use +only to their possessors. In the result I can only think of two such +instances. These, therefore, I will now dispose of.</p> + +<p>The first was pointed out, and has been fully discussed, by Darwin +himself. Certain species of ants are fond of a sweet fluid that is +secreted by aphides, and they even keep the aphides as we keep cows for +the purpose of profiting by their “milk.” Now the point is, +that the use of this sweet secretion to the aphis itself has not yet +been made out. Of course, if it is of no use to the aphis, it would +furnish a case which completely meets Darwin’s own challenge. But, +even if this supposition did not stand out of analogy with all the other +facts of organic nature, most of us would probably deem it prudent to +hold that the secretion must primarily be of some use to the aphis +itself, although the matter has not been sufficiently investigated to +inform us of what this use is. For, in any case, the secretion is not of +any vital importance to the ants which feed upon it: and I think but few +impartial minds would go so far to save an hypothesis as to maintain, +that the Divinity had imposed this drain upon the internal resources of +one species of insect for the sole purpose of supplying a luxury to +another.<span class="pagenum"><a name="page_293" id="page_293">[293]</a></span> +On the whole, it seems most probable that the fluid is of the nature of +an excretion, serving to carry off waste products. Such, at all events, +was the opinion at which Darwin himself arrived, as a result of +observing the facts anew, and in relation to his theory.</p> + +<hr class='minor' /> + +<p>The other instance to which I have alluded as seeming at first sight +likely to answer Darwin’s challenge is the formation of vegetable +galls. The great number and variety of galls agree in presenting a more +or less elaborate structure, which is not only foreign to any of the +uses of plant-life, but singularly and specially adapted to those of the +insect-life which they shelter. Yet they are produced by a growth of the +plant itself, when suitably stimulated by the insects’ +inoculation—or, according to recent observations, by emanations from +the bodies of the larvæ which develop from the eggs deposited in the +plant by the insect. Now, without question, this is a most remarkable +fact; and if there were many more of the like kind to be met with in +organic nature, we might seriously consider whether the formation of +galls should not be held to make against the ubiquitous agency of +natural selection. But inasmuch as the formation of galls stands out as +an exception to the otherwise universal rule of every species for +itself, and for itself alone, we are justified in regarding this one +apparent exception with extreme suspicion. Indeed, I think we are +justified in regarding the peculiar pathological effect produced in the +plant by the secretions of the insect as having been in the first +instance accidentally beneficial to the insects. Thus, if any other +effect than that of a growing tumour had been produced in <span class="pagenum"><a name="page_294" id="page_294">[294]</a></span> +the first instance, or if the needs of the insect progeny had not been +such as to have derived profit from being enclosed in such a tumour, +then, of course, the inoculating instinct of these animals could not +have been developed by natural selection. But, given these two +conditions, and it appears to me there is nothing very much more +remarkable about an accidental correlation between the effects of a +parasitic larva on a plant and the needs of that parasite, than there is +between the similarly accidental correlation between a hydated parasite +and the nutrition furnished to it by the tissues of a warm-blooded +animal. Doubtless the case of galls is somewhat more remarkable, +inasmuch as the morbid growth of the plant has more concern in the +correlation—being, in many instances, a more specialized structure +on the part of a host than occurs anywhere else, either in the animal or +vegetable world. But here I may suggest that although natural selection +cannot have acted upon the plant directly, so as to have produced galls +ever better and better adapted to the needs of the insect, it may have +so acted upon the plants indirectly <i>though the insects</i>. For it may +very well have been that natural selection would ever tend to preserve +those individual insects, the quality of whose emanations tended to +produce the form of galls best suited to nourish the insect progeny; and +thus the character of these pathological growths may have become ever +better and better adapted to the needs of the insects. Lastly, looking +to the enormous number of relations and inter-relations between all +organic species, it is scarcely to be wondered at that even so +extraordinary an instance of correlation as this should have arisen thus +by accident, and then <span class="pagenum"><a name="page_295" id="page_295">[295]</a></span> +have been perfected by such an <i>indirect</i> agency of natural selection +as is here suggested<a name="FNanchor_37_37" id="FNanchor_37_37"></a><a href="#Footnote_37_37" class="fnanchor">[37]</a>.</p> + +<hr class='minor' /> + +<p>The third general class of facts which tell so immensely in favour of +natural selection as an important cause of organic evolution, are those +of domestication. The art of the horticulturist, the fancier, the +cattle-breeder, &c., consists in producing greater and greater +deviations from a given wild type of plant or animal, in any particular +direction that may be desired for purposes either of use or of beauty. +Cultivated cereals, fruits, and flowers are known to have been all +derived from wild species; and, of course, the same applies to all our +domesticated varieties of animals. Yet if we compare a cabbage rose with +a wild rose, a golden pippin apple with a crab, a toy terrier with any +species of wild dog, not to mention any number of other instances, there +can be no question that, if such differences had appeared in nature, the +organisms presenting them would have been entitled to rank as distinct +species—or even, in many cases, as distinct genera. Yet we know, as a +matter of fact, that all these differences have been produced by a +process of artificial selection, or pairing, which has been continuously +practised by horticulturists and breeders through a number of +generations. It is the business of these men to note the individual +organisms which show most variation in the directions required, and then +to propagate from these individuals, in order that the progeny shall +inherit the qualities desired. The results thus become cumulative from +generation to generation, until we now have an astonishing manifestation +<span class="pagenum"><a name="page_296" id="page_296">[296]</a></span> +of useful qualities on the one hand, and of beautiful qualities on the +other, according as the organisms have been thus bred for purposes of +use or for those of beauty.</p> + +<p>Now it is immediately obvious that in these cases the process of +artificial selection is precisely analogous to that of natural selection +(and of sexual selection which will be considered later on), in all +respects save one: the utility or the beauty which it is the aim of +artificial selection continually to enhance, is utility or beauty in +relation to the requirements or to the tastes of man; whereas the +utility or the beauty which is produced by natural selection and sexual +selection has reference only to the requirements or the tastes of the +organisms themselves. But, with the exception of this one point of +difference, the processes and the products are identical in kind. +Persevering selection by man is thus proved to be capable of creating +what are virtually new specific types, and this in any required +direction. Hence, when we remember how severe is the struggle for +existence in nature, it becomes impossible to doubt that selection by +nature is able to do at least as much as artificial selection in the way +of thus creating new types out of old ones. Artificial selection, +indeed, notwithstanding the many and marvellous results which it has +accomplished, can only be regarded as but a feeble imitation of natural +selection, which must act with so much greater vigilance and through +such immensely greater periods of time. In a word, the proved +capabilities of artificial selection furnish, in its best conceivable +form, what is called an argument <i>a fortiori</i> in favour of natural +selection.</p> + +<p><span class="pagenum"><a name="page_297" id="page_297">[297]</a></span> +Or, to put it in another way, it may be said that for thousands of years +mankind has been engaged in making a gigantic experiment to test, as it +were by anticipation, the theory of natural selection. For, although +this prolonged experiment has been carried on without any such intention +on the part of the experimenters, it is none the less an experiment in +the sense that its results now furnish an overwhelming verification of +Mr. Darwin’s theory. That is to say, they furnish overwhelming +proof of the efficacy of the selective principle in the modification of +organic types, when once this principle is brought steadily and +continuously to bear upon a sufficiently long series of generations.</p> + +<p>In order to furnish ocular evidence of the value of this line of +verification, I have had the following series of drawings prepared. +Another and equally striking series might be made of the products of +artificial selection in the case of plants; but it seems to me that the +case of animals is more than sufficient for the purpose just stated. +Perhaps it is desirable to add that considerable care has been bestowed +upon the execution of these portraits; and that in every case the latter +have been taken from the most typical specimens of the artificial +variety depicted. Those of them which have not been drawn directly from +life are taken from the most authoritative sources; and, before being +submitted to the engraver, they were all examined by the best judges in +each department. In none of the groups, however, have I aimed at an +exhaustive representation of all the varieties: I have merely introduced +representatives of as many as the page would in each case accommodate.</p> + +<p class="pagenum"><a name="page_298" id="page_298">[298]</a></p> +<div class="figcenter" style="width: 300px;"> +<img src="images/i_315_091.jpg" width="300" height="500" alt="Pigeons." title="" /> +<span class="caption"><span class="sc">Fig.</span> 91.—Pigeons. Drawn from life (prize specimens).</span></div> + +<p class="pagenum"><a name="page_299" id="page_299">[299]</a></p> +<div class="figcenter" style="width: 303px;"> +<img src="images/i_316_092.jpg" width="303" height="500" alt="Pigeons, continued." title="" /> +<span class="caption"><span class="sc">Fig.</span> 92.—Pigeons, continued. Drawn from life (prize +specimens).</span></div> + +<p class="pagenum"><a name="page_300" id="page_300">[300]</a></p> +<div class="figcenter" style="width: 296px;"> +<img src="images/i_317_093.jpg" width="296" height="500" alt="Fowls." title="" /> +<span class="caption"><span class="sc">Fig.</span> 93.—Fowls. Drawn from life (prize specimens).</span></div> + +<p class="pagenum"><a name="page_301" id="page_301">[301]</a></p> +<div class="figcenter" style="width: 295px;"> +<img src="images/i_318_094.jpg" width="295" height="500" alt="Fowls, continued." title="" /> +<span class="caption">Fig. 94.—Fowls, continued. Drawn from life (prize +specimens).</span></div> + +<p class="pagenum"><a name="page_302" id="page_302">[302]</a></p> +<div class="figcenter" style="width: 295px;"> +<img src="images/i_319_095.jpg" width="295" height="500" alt="Pair of Japanese Fowls, long-tailed breed." title="" /> +<span class="caption"><span class="sc">Fig.</span> 95.—Pair of Japanese Fowls, long-tailed breed. +Drawn from stuffed specimens in the British Museum.</span></div> + +<p class="pagenum"><a name="page_303" id="page_303">[303]</a></p> +<div class="figcenter" style="width: 307px;"> +<img src="images/i_320_096.jpg" width="307" height="500" alt="Canaries." title="" /> +<span class="caption"><span class="sc">Fig.</span> 96.—Canaries. Drawn from life (prize specimens).</span></div> + +<p class="pagenum"><a name="page_304" id="page_304">[304]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_321_097.jpg" width="500" height="401" alt="Sebastopol, or Frizzled Goose." title="" /> +<span class="caption"><span class="sc">Fig.</span> 97.—Sebastopol, or Frizzled Goose. Drawn from a +photograph.</span></div> + +<div class="figcenter" style="width: 400px;"> +<img src="images/i_321_098.jpg" width="400" height="375" alt="The Dingo, or wild dog of Australia." title="" /> +<span class="caption"><span class="sc">Fig.</span> 98.—The Dingo, or wild dog of Australia, 1/10 nat. +size. Drawn from life (<i>Zoological Gardens</i>).</span></div> + +<p class="pagenum"><a name="page_305" id="page_305">[305]</a></p> +<div class="figcenter" style="width: 298px;"> +<img src="images/i_322_099.jpg" width="298" height="500" alt="Dogs." title="" /> +<span class="caption"><span class="sc">Fig.</span> 99.—Dogs. Drawn from life (prize specimens).</span></div> + +<p class="pagenum"><a name="page_306" id="page_306">[306]</a></p> +<div class="figcenter" style="width: 300px;"> +<img src="images/i_323_100.jpg" width="300" height="500" alt="Dogs, continued." title="" /> +<span class="caption"><span class="sc">Fig.</span> 100.—Dogs, continued. Drawn from life (prize +specimens).</span></div> + +<p class="pagenum"><a name="page_307" id="page_307">[307]</a></p> +<div class="figcenter" style="width: 400px;"> +<img src="images/i_324_101.jpg" width="400" height="375" alt="The Hairless Dog of Japan." title="" /> +<span class="caption"><span class="sc">Fig.</span> 101.—The Hairless Dog of Japan, 1/10 nat. size. +Drawn from a photograph kindly lent for the purpose by the proprietor.</span></div> + +<div class="figcenter" style="width: 400px;"> +<img src="images/i_324_102.jpg" width="400" height="385" alt="The skull of a Bull-dog compared with that of +a Deerhound." title="" /> +<span class="caption"><span class="sc">Fig.</span> 102.—The skull of a Bull-dog compared with that of +a Deerhound. Drawn from nature.</span></div> + +<p class="pagenum"><a name="page_308" id="page_308">[308]</a></p> +<div class="figcenter" style="width: 296px;"> +<img src="images/i_325_103.jpg" width="296" height="500" alt="Rabbits." title="" /> +<span class="caption"><span class="sc">Fig.</span> 103. Rabbits. Drawn from life (prize specimens).</span></div> + +<p class="pagenum"><a name="page_309" id="page_309">[309]</a></p> +<div class="figcenter" style="width: 289px;"> +<img src="images/i_326_104.jpg" width="289" height="500" alt="Horses." title="" /> +<span class="caption"><span class="sc">Fig.</span> 104.—Horses. Drawn from life (prize specimens).</span></div> + +<p class="pagenum"><a name="page_310" id="page_310">[310]</a></p> +<div class="figcenter" style="width: 300px;"> +<img src="images/i_327_105.jpg" width="300" height="500" alt="Sheep. British breeds." title="" /> +<span class="caption"><span class="sc">Fig.</span> 105.—Sheep. The illustrations are confined to +British breeds. Drawn from life (prize specimens).</span></div> + +<p class="pagenum"><a name="page_311" id="page_311">[311]</a></p> +<div class="figcenter" style="width: 297px;"> +<img src="images/i_328_106.jpg" width="297" height="500" alt="Cattle. British breeds." title="" /> +<span class="caption"><span class="sc">Fig.</span> 106.—Cattle. The illustrations are confined to +British breeds. Drawn from life (prize specimens).</span></div> + +<p class="pagenum"><a name="page_312" id="page_312">[312]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_329_107.jpg" width="500" height="410" alt="Wild Boar contrasted with a modern +Domesticated Pig." title="" /> +<span class="caption"><span class="sc">Fig.</span> 107.—Wild Boar contrasted with a modern +Domesticated Pig. Drawn from life (<i>Zoological Gardens</i>, and prize +specimen).</span></div> + +<p>The exigencies of space have prevented, in some of the groups, strict +adherence to a uniform scale—with the result that contrasts between +different breeds in respect of size are not adequately rendered. This +remark applies especially to the dogs; for although the artist has +endeavoured to draw them in perspective, unless the distance between +those in the foreground and those in the background is understood to be +more considerable than it appears, an inadequate idea is given of the +relative differences of size. The most instructive of the groups, I +think, is that of the Canaries; because the many and great changes in +<span class="pagenum"><a name="page_313" id="page_313">[313]</a></span> +different directions must in this case have been produced by artificial +selection in so comparatively short a time—the first mention of this +bird that I can find being by Gesner, in the sixteenth century.</p> + +<hr class='minor' /> + +<p>Now, it is surely unquestionable that in these typical proofs of the +efficacy of artificial selection in the modification of specific types, +we have the strongest conceivable testimony to the power of natural +selection in the same direction. For it thus appears that wherever +mankind has had occasion to operate by selection for a sufficiently long +time—that is to say, on whatever species of plant or animal he chooses +thus to operate for the purpose of modifying the type in any required +direction,—the results are always more or less the same: he finds that +all specific types lend themselves to continuous deflection in any +particulars of structure, colour, &c., that he may desire to modify.</p> + +<p>Nevertheless, to this parallel between the known effects of artificial +selection, and the inferred effects of natural selection, two objections +have been urged. The first is, that in the case of artificial selection +the selecting agent is a voluntary intelligence, while in the case of +natural selection the selecting agent is Nature herself; and whether or +not there is any counterpart of man’s voluntary intelligence in +nature is a question with which Darwinism has nothing to do. Therefore, +it is alleged, the analogy between natural selection and artificial +selection fails <i>ab initio</i>, or at the fountain-head of the causes which +are taken by the analogy to be respectively involved.</p> + +<p>The second objection to the analogy is, that the products of artificial +selection, closely as they may <span class="pagenum"><a name="page_314" id="page_314">[314]</a></span> +resemble natural species in all other respects, nevertheless present +one conspicuous and highly important point of difference: they rarely, +if ever, present the physiological character of mutual infertility, +which is a character of extremely general occurrence in the case of +natural species, even when these are most nearly allied.</p> + +<p>I will deal with these two objections in the next chapter, where I shall +be concerned with the meeting of all the objections which have ever been +urged against the theory of natural selection. Meanwhile I am engaged +only in presenting the general arguments which support the theory, and +therefore mention these objections to one of them merely <i>en passant</i>. +And I do so in order to pledge myself effectually to dispose of them +later on, so that for the purposes of my present argument both these +objections may be provisionally regarded as non-existent; which means, +in other words, that we may provisionally regard the analogy between +artificial selection and natural selection as everywhere logically +intact.</p> + +<hr class='minor' /> + +<p>To sum up, then, the results of the foregoing exposition thus far, what +I hold to be the three principal, or most general, arguments in favour +of the theory of natural selection, are as follows.</p> + +<p>First, there is the <i>a priori</i> consideration that, if on independent +grounds we believe in the theory of evolution at all, it becomes obvious +that natural selection <i>must</i> have had <i>some</i> part in the process. For +no one can deny the potent facts of heredity, variability, the struggle +for existence, and survival of the fittest. But to admit these facts is +to admit <span class="pagenum"><a name="page_315" id="page_315">[315]</a></span> +natural selection as a principle which must be, at any rate, one of the +factors of organic evolution, supposing such evolution to have taken +place. Next, when we turn from these <i>a priori</i> considerations, which +thus show that natural selection <i>must</i> have been concerned to some +extent in the process of evolution, we find in organic nature evidence +<i>a posteriori</i> of the extent to which this principle <i>has</i> been thus +concerned. For we find that among all the countless millions of adaptive +structures which are to be met with in organic nature, it is an +invariable rule that they exist in relation to the needs of the +particular species which present them: they never have any primary +reference to the needs of other species. And as this extraordinarily +large and general fact is exactly what the theory of natural selection +would expect, the theory is verified by the fact in an extraordinarily +cogent manner. In other words, the fact goes to prove that in <i>all</i> +cases where adaptive structures or instincts are concerned, natural +selection must have been either the sole cause at work, or, at the +least, an influence controlling the operation of all other causes.</p> + +<p>Lastly, an actually experimental verification of the theory has been +furnished on a gigantic scale by the operations of breeders, fanciers, +and horticulturists. For these men, by their process of selective +accumulation, have empirically proved what immense changes of type may +thus be brought about; and so have verified by anticipation, and in a +most striking manner, the theory of natural selection—which, as now so +fully explained, is nothing more than a theory of cumulative +modifications by means of selective breeding.</p> + +<p><span class="pagenum"><a name="page_316" id="page_316">[316]</a></span> +So much, then, by way of generalities. But perhaps the proof of natural +selection as an agency of the first importance in the transmutation of +species may be best brought home to us by considering a few of its +applications in detail. I will therefore devote the rest of the present +chapter to considering a few cases of this kind.</p> + +<p>There are so many large fields from which such special illustrations may +be supplied, that it is difficult to decide which of them to draw upon. +For instance, the innumerable, always interesting, and often astonishing +adaptations on the part of flowers to the fertilising agency of insects, +has alone given rise to an extensive literature since the time when +Darwin himself was led to investigate the subject by the guidance of his +own theory. The same may be said of the structures and movements of +climbing plants, and in short, of all the other departments of natural +history where the theory of natural selection has led to the study of +the phenomena of adaptation. For in all these cases the theory of +natural selection, which first led to their discovery, still remains the +only scientific theory by which they can be explained. But among all the +possible fields from which evidences of this kind may be drawn, I think +the best is that which may be generically termed defensive colouring. To +this field, therefore, I will restrict myself. But, even so, the cases +to be mentioned are but mere samples taken from different divisions of +this field; and therefore it must be understood at the outset that they +could easily be multiplied a hundred-fold.</p><p> + +<span class="pagenum"><a name="page_317" id="page_317">[317]</a></span></p> +<h3 class='i'>Protective Colouring.</h3> + +<div class="figcenter" style="width: 500px;"> +<img src="images/i_334_108.jpg" width="500" height="416" alt="Seasonal changes of colour in Ptarmigan." title="" /> +<div class="captionh"><span class="sc">Fig.</span> 108.—Seasonal changes of colour in Ptarmigan (<span +class='sn'>Lagopus mutus</span>). Drawn from stuffed specimens in the British +Museum, 1/6 nat. size, with appropriate surroundings supplied.</div></div> + +<p>A vast number of animals are rendered more or less inconspicuous by +resembling the colours of the surfaces on which they habitually rest. +Such, for example, are grouse, partridges, rabbits, &c. Moreover, there +are many cases in which, if the needs of the creature be such that it +must habitually frequent surfaces of different colours, it has acquired +the power of changing its colour accordingly—e. g. cuttle-fish, +flat-fish, frogs, chameleons, &c. The physiological mechanism whereby +these adaptive changes of colour are produced <span class="pagenum"><a name="page_318" id="page_318">[318]</a></span> +differs in different animals; but it is needless for our purposes to go +into this part of the subject. Again, there are yet other cases where +protective colouring which is admirably suited to conceal an animal +through one part of the year, would become highly conspicuous during +another part of it—namely, when the ground is covered with snow. +Accordingly, in these cases the animals change their colour in the +winter months to a snowy white: witness stoats, mountain hares, +ptarmigan, &c. (Fig. 108.)</p> + +<p>Now, it is sufficiently obvious that in all these classes of cases the +concealment from enemies or prey which is thus secured is of advantage +to the animals concerned; and, therefore, that in the theory of natural +selection we have a satisfactory theory whereby to explain it. And this +cannot be said of any other theory of adaptive mechanisms in nature that +has ever been propounded. The so-called Lamarckian theory, for instance, +cannot be brought to bear upon the facts at all; and on the theory of +special creation it is unintelligible why the phenomena of protective +colouring should be of such general occurrence. For, in as far as +protective colouring is of advantage to the species which present it, it +is of corresponding disadvantage to those other species against the +predatory nature of which it acts as a defence. And, of course, the same +applies to yet other species, if they serve as prey. Moreover, the more +minutely this subject is investigated in all its details, the more +exactly is it found to harmonise with the naturalistic +interpretation<a name="FNanchor_38_38" id="FNanchor_38_38"></a><a href="#Footnote_38_38" class="fnanchor">[38]</a>.</p> + +<p><span class="pagenum"><a name="page_319" id="page_319">[319]</a></span> +In the first place, we always find a complete correspondence between +imitative colouring and instinctive endowment. If a caterpillar exactly +resembles the colour of a twig, it also presents the instinct of +habitually reposing in the attitude which makes it most resemble a +twig—standing out from the branch on which it rests at the same angle +as is presented by the real twigs of the tree on which it lives.</p> + +<p>Here, again, is a bird protectively coloured so as to resemble stones +upon the rough ground where it habitually lives; and the drawing shows +the attitude in which the bird instinctively reposes, so as still +further to increase its resemblance to a stone. (Fig. 109.)</p> + +<p class="pagenum"><a name="page_320i" id="page_320i">[320i.]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_337_109.jpg" width="500" height="435" alt="Œdicnemus crepitans." title="" /> +<div class="captionh"><span class="sc">Fig.</span> 109.—<span class='sn'>Œdicnemus crepitans</span>, showing the +instinctive attitude of concealment. Drawn from a stuffed specimen in +the British Museum, 1/6 nat. size, with appropriate surroundings +supplied.</div></div> + +<p><span class="pagenum"><a name="page_319c" id="page_319c">[319c.]</a></span> +To take only one other instance, hares and rabbits, like grouse and +partridges—or like the plover just alluded to,—instinctively crouch +upon those surfaces the colours of which they resemble; and I have often +remarked that if, on account of any individual peculiarity of +coloration, the animal is not able thus <span class="pagenum"><a name="page_320" id="page_320">[320]</a></span> +to secure concealment, it nevertheless exhibits the instinct of +crouching which is of benefit to all its kind, although, from the +accident of its own abnormal colouring, this instinct is then actually +detrimental to the animal itself. For example, every sportsman must have +noticed that the somewhat rare melanic variety of the common rabbit will +crouch as steadily as the normal brownish-gray type, notwithstanding +that, owing to its abnormal colour, a “nigger-rabbit” thus +renders itself the most conspicuous object in the <span class="pagenum"><a name="page_321" id="page_321">[321]</a></span> +landscape. In all such cases, of course, there has been a deviation +from the normal type in respect of colour, with the result that the +inherited instinct is no longer in tune with the other endowments of the +animal. Such a variation of colour, therefore, will tend to be +suppressed by natural selection; while any variations which may bring +the animal still more closely to resemble its habitual surroundings will +be preserved. Thus we can understand the truly wonderful extent to which +this principle of protective colouring has been carried in many cases +where the need of it has been most urgent.</p> + +<p>Not only colour, but structure, may be profoundly modified for the +purposes of protective concealment. Thus, caterpillars which resemble +twigs do so not only in respect of colour, but also of shape; and this +even down to the most minute details in cases where the adaptation is +most complete: certain butterflies and leaf-insects so precisely +resemble the leaves upon which, or among which, they live, that it is +almost impossible to detect them in the foliage—not only the colour, +the shape, and the venation being all exactly imitated, but in some +cases even the defects to which the leaves are liable, in the way of +fungoid growths, &c. There are other insects which with similar +exactness resemble moss, lichens, and so forth. A species of fish +secures a complete resemblance to bunches of sea-weed by a frond-like +modification of all its appendages, and so on through many other +instances. Now, in all such cases where there is so precise an +imitation, both in colour and structure, it seems impossible to suggest +any other explanation of the facts than the one which is supplied by +<span class="pagenum"><a name="page_323" id="page_323">[323]</a></span> +Mr. Darwin’s theory—namely, that the more perfect the resemblance +is caused to become through the continuous influence of natural +selection always picking out the best imitations, the more highly +discriminative becomes the perception of those enemies against the +depredations of which this peculiar kind of protection is developed; so +that, in virtue of this action and re-action, eventually we have a +degree of imitation which renders it almost impossible for a naturalist +to detect the animal when living in its natural environment.</p> + +<p class="pagenum"><a name="page_322" id="page_322">[322]</a></p> +<div class="figcenter" style="width: 295px;"> +<img src="images/i_339_110.jpg" width="295" height="500" alt="Imitative forms and colours in insects." title="" /> +<span class="caption"><span class="sc">Fig.</span> 110.—Imitative forms and colours in insects. Drawn +from nature (<i>R. Coll. Surg. Mus.</i>).</span></div> + +<p class="pagenum"><a name="page_323c" id="page_323c">[323c.]</a></p> +<h3 class='i'>Warning Colours.</h3> + +<p>In strange and glaring contrast to all these cases of protective +colouring, stand other cases of conspicuous colouring. Thus, for +example, although there are numberless species of caterpillars which +present in an astonishing degree the phenomena of protective colouring, +there are numberless other species which not only fail to present these +phenomena in any degree, but actually go to the opposite extreme of +presenting colours which appear to have been developed for the sake of +their conspicuousness. At all events, these caterpillars are usually the +most conspicuous objects in their surroundings, and therefore in the +early days of Darwinism they were regarded by Darwin himself as +presenting a formidable difficulty in the way of his theory. To Mr. +Wallace belongs the merit of having cleared up this difficulty in an +extraordinarily successful manner. He virtually reasoned thus. If the +<i>raison d’être</i> of protective colouring be that of concealing +agreeably flavoured caterpillars from the eye-sight of birds, may not +<span class="pagenum"><a name="page_324" id="page_324">[324]</a></span> +the <i>raison d’être</i> of conspicuous colouring be that of +protecting disagreeably flavoured caterpillars from any possibility of +being mistaken by birds? Should this be the case, of course the more +conspicuous the colouring the better would it be for the caterpillars +presenting it. Now as soon as this suggestion was acted upon +experimentally, it was found to be borne out by facts. Birds could not +be induced to eat caterpillars of the kinds in question; and there is +now no longer any doubt that their conspicuous colouring is correlated +with their distastefulness to birds, in the same way as the +inconspicuous or imitative colouring of other caterpillars is correlated +with their tastefulness to birds. Here then is yet another instance, +added to those already given, of the verification yielded to the theory +of natural selection by its proved competency as a guide to facts in +nature; for assuredly this particular class of facts would never have +been suspected but for its suggestive agency.</p> + +<p>As in the case of protective imitation, so in this case of warning +conspicuousness, not only colour, but structure may be greatly modified +for the purpose of securing immunity from attack. Here, of course, the +object is to assume, as far as possible, a touch-me-not appearance; so +that, although destitute of any real means of offence, the creatures in +question present a fictitiously dangerous aspect. As the +Devil’s-coach-horse turns up his stingless tail when threatened by +an enemy, so in numberless ways do many harmless animals of all classes +pretend to be formidable. But the point now is that these instincts of +self-defence are often helped out by structural <span class="pagenum"><a name="page_325" id="page_325">[325]</a></span> +modifications, expressly and exclusively adapted to this end. For +example, what a remarkable series of protective adjustments occurs in +the life-history of the Puss Moth—culminating with so comical an +instance of the particular device now under consideration as the +following. I quote the facts from Mr. E. B. Poulton’s admirable +book on <i>The Colours of Animals</i> (pp. 269-271).</p> + +<div class='blockquot'><div class="figcenter" style="width: 400px;"> +<img src="images/i_342_111.jpg" width="400" height="125" alt="The larva of Puss Moth when +undisturbed." title="" /> +<span class="caption"><span class="sc">Fig.</span> 111.—The larva of Puss Moth (<span class='sn'>C. vinula</span>) when +undisturbed; full-fed; natural size.</span></div> + +<p>The larva of the Puss Moth (Cerura vinula) is very common upon +poplar and willow. The circular dome-like eggs are laid, either +singly or in little groups of two or three, upon the upper side of +the leaf, and being of a reddish colour strongly suggest the +appearance of little galls, or the results of some other injury to +the leaf. The youngest larvæ are black, and also rest upon the +upper surface of the leaf, resembling the dark patches which are +commonly seen in this position. As the larva grows, the apparent +black patch would cover too large a space, and would lead to +detection if it still occupied the whole surface of the body. The +latter gains a green ground-colour which harmonises with the leaf, +while the dark marking is chiefly confined to the back. As growth +proceeds the relative amount of green increases, and the dark mark +is thus prevented from attaining a size which would render it too +conspicuous. In the last stage of growth the green larva becomes +very large, and usually rests on the twigs of its food-plant (Fig. +111). The dark colour is still present on the back but is softened +to a purplish tint, which tends to be replaced by a combination of +white and green in many of the largest larvæ. Such a larva is well +concealed by General Protective<span class="pagenum"><a name="page_326" id="page_326">[326]</a></span> +Resemblance, and one may search a long time before finding it, although +assured of its presence from the stripped branches of the food-plant and +the fæces on the ground beneath.</p> + +<p>As soon as a large larva is discovered and disturbed it withdraws +its head into the first body-ring, inflating the margin, which is +of a bright red colour.<span class="figleft" style="width: 220px;"> +<img src="images/i_343_112.jpg" width="220" height="179" alt="The larva of Puss Moth in its terrifying +attitude after being disturbed." title="" /> +<span class="caption"><span class="sc">Fig.</span> 112.—The larva of Puss Moth in its terrifying +attitude after being disturbed; full-fed; natural size.</span></span> +There are two intensely black spots on this +margin in the appropriate position for eyes, and the whole +appearance is that of a large flat face extending to the outer edge +of the red margin (see Fig. 112). The effect is an intensely +exaggerated caricature of a vertebrate face, which is probably +alarming to the vertebrate enemies of the caterpillar. The +terrifying effect is therefore mimetic. The movements entirely +depend on tactile impressions: when touched ever so lightly a +healthy larva immediately assumes the terrifying attitude, and +turns so as to present its full face towards the enemy; if touched +on the other side or on the back it instantly turns its face in the +appropriate direction. The effect is also greatly strengthened by +two pink whips which are swiftly protruded from the prongs of the +fork in which the body terminates. The prongs represent the last +pair of larval legs which have been greatly modified from their +ordinary shape and use. The end of the body is at the same time +curved forward over the back (generally much further than in Fig. +112), so that the pink filaments are brandished above the head.</p></div> + +<h3 class='i'>Mimicry.</h3> + +<p>Lastly, these facts as to imitative and conspicuous colouring lead on to +the yet more remarkable facts of what is called mimicry. By mimicry is +meant the imitation in form and colour of one species by another, +<span class="pagenum"><a name="page_327" id="page_327">[327]</a></span> +in order that the imitating species may be mistaken for the imitated, +and thus participate in some advantage which the latter enjoys. For +instance, if, as in the case of the conspicuously-coloured caterpillars, +it is of advantage to an ill-savoured species that it should hold out a +warning to enemies, clearly it may be of no less advantage to a +well-savoured species that it should borrow this flag, and thus be +mistaken for its ill-savoured neighbour. Now, the extent to which this +device of mimicry is carried is highly remarkable, not only in respect +of the number of its cases, but also in respect of the astonishing +accuracy which in most of these cases is exhibited by the imitation. +There need be little or virtually no zoological affinity between the +imitating and the imitated forms; that is to say, in some cases the +zoological affinity is not closer than ordinal, and therefore cannot +possibly be ascribed to kinship. Like all the other branches of the +general subject of protective resemblance in form or colouring, this +branch has already been so largely illustrated by previous writers, +that, as in the previous cases, I need only give one or two examples. +Those which I choose are chosen on account of the colours concerned not +being highly varied or brilliant, and therefore lending themselves to +less ineffectual treatment by wood-engraving than is the case where +attempts are made to render by this means even more remarkable +instances. (Figs. 113, 114, 115.)</p> + +<p class="pagenum"><a name="page_328" id="page_328">[328]</a></p> +<div class="figcenter" style="width: 295px;"> +<img src="images/i_345_113.jpg" width="295" height="500" alt="Three cases of mimicry." title="" /> +<span class="caption"><span class="sc">Fig.</span> 113.—Three cases of mimicry. Drawn from nature: +first two pairs nat. size, last pair 2/3 (<i>R. Coll. Surg. Mus.</i>).</span></div> + +<p><span class="pagenum"><a name="page_327c" id="page_327c">[327c.]</a></span> +It is surely apparent, without further comment, that it is impossible to +imagine stronger evidence in favour of natural selection as a true cause +in nature, than is furnished by this culminating fact in the matter of +<span class="pagenum"><a name="page_329" id="page_329">[329]</a></span> +protective resemblance, whereby it is shown that a species of one +genus, family, or even order, will accurately mimic the appearance of a +species belonging to another genus, family, or order, so as to deceive +its natural enemies into mistaking it for a creature of so totally +different a kind. And it must be added that while this fact of mimicry +is of extraordinarily frequent occurrence, there can be no possibility +of our mistaking its purpose. For the fact is never observable except in +the case of species which occupy the same area or district.</p> + +<div class="figcenter" style="width: 400px;"> +<img src="images/i_346_114.jpg" width="400" height="335" alt="Two flies resembling +a wasp and a bee." title="" /> +<div class="captionh"><span class="sc">Fig.</span> 114.—Two further cases of mimicry; flies resembling +a wasp in the one and a bee in the other. Drawn from nature: nat. size +(<i>R. Coll. Surg. Mus.</i>).</div></div> + +<p class="pagenum"><a name="page_330" id="page_330">[330]</a></p> +<div class="figcenter" style="width: 295px;"> +<img src="images/i_347_115.jpg" width="295" height="500" alt="A non-venomous species +of snake resembling a venomous one." title="" /> +<span class="caption"><span class="sc">Fig.</span> 115.—A case of mimicry where a non-venomous species +of snake resembles a venomous one. Drawn from nature: 1/3 nat. size (<i>R. +Coll. Surg. Mus.</i>).</span></div> + +<p><span class="pagenum"><a name="page_329c" id="page_329c">[329c.]</a></span> +Such being what appears to me the only reasonable view of the matter, I +will now conclude this chapter on the evidences of natural selection as +at all events the <span class="pagenum"><a name="page_331" id="page_331">[331]</a></span> +main factor of organic evolution, by simply adding illustrations of +two further cases of mimicry, which are perhaps even more remarkable +than any of the foregoing examples. The first of the two (Fig. 115) +speaks for itself. The second will be rendered intelligible by the +following few words of explanation.</p> + +<p>There are certain ants of the Amazons which present the curious instinct +of cutting off leaves from trees, and carrying them like banners over +their heads to the hive, as represented in Fig. 116, B, where one ant is +shown without a leaf, and the others each with a leaf. Their object in +thus collecting leaves is probably that of growing a fungus upon the +“soil” which is furnished by the leaves when decomposing. +But, be this as it may<a name="FNanchor_39_39" id="FNanchor_39_39"></a><a href="#Footnote_39_39" class="fnanchor">[39]</a>, +the only point we are now concerned with is the appearance which these +ants present when engaged in their habitual operation of carrying +leaves. For it has been recently observed by Mr. W. L. Sclater, that in +the localities where these hymenopterous insects occur, there occurs +also a <span class='sn'>homopterous</span> insect which mimics the ant, leaf and all, in a +wonderfully deceptive manner. The leaf is imitated by the thin flattened +body of the insect, “which in its dorsal aspect is so compressed +laterally that it is no thicker than a leaf, and terminates in a sharp +jagged edge.” The colour is exactly the same as that of a leaf, +and the brown legs show themselves beneath the green body in just the +same way as those of the ant show themselves beneath the leaf. So that +both the form and the colouring of the homopterous insect has been +brought to resemble, with singular exactness, <span class="pagenum"><a name="page_332" id="page_332">[332]</a></span> +those belonging to a different order of insect, when the latter is +engaged in its peculiar avocation. A glance at the figure is enough to +show the means employed and the result attained. In A, an ant and its +mimic are represented as about 2½ times their natural size, and +both proceeding in the same direction. It ought to be mentioned, +however, that in reality the margin of the leaf is seldom allowed to +retain its natural serrations as here depicted: the ants usually gnaw +the edge of the real leaf, so that the margin of the false one bears an +even closer resemblance to it than the illustration represents. B is a +drawing from life of a group of five ants carrying leaves, and their +mimic walking beside them<a name="FNanchor_40_40" id="FNanchor_40_40"></a><a href="#Footnote_40_40" class="fnanchor">[40]</a>.</p> + +<div class="figcenter" style="width: 369px;"> +<img src="images/i_350_116.jpg" width="369" height="500" alt="Protective Mimicry" title="" /> +<span class="caption"><span class="sc">Fig. 116. <span class='lc'>PROTECTIVE MIMICRY</span></span></span></div> + +<hr /><p class="pagenum"><a name="page_333" id="page_333">[333]</a></p> +<h2><a name="CHAPTER_IX" id="CHAPTER_IX"></a>CHAPTER IX.<br /><br /> +<span class="sc">Criticisms of the Theory of Natural Selection.</span></h2> + +<p>I will now proceed to consider the various objections and difficulties +which have hitherto been advanced against the theory of natural +selection.</p> + +<p>Very early in the day Owen hurled the weight of his authority against +the new theory, and this with a strength of onslaught which was only +equalled by its want of judgment. Indeed, it is painfully apparent that +he failed to apprehend the fundamental principles of the Darwinian +theory. For he says:—</p> + +<div class='blockquot'><p>Natural Selection is an explanation of the process [of +transmutation] of the same kind and value as that which has been +proffered of the mystery of “secretion.” For example, a +particular mass of matter in a living animal takes certain elements +out of the blood, and rejects them as “bile.” +Attributes were given to the liver which can only be predicated of +the whole animal; the “appetency” of the liver, it was +said, was for the elements of bile, and “biliosity,” or +the “hepatic sensation,” guided the gland to their +secretion. Such figurative language, I need not say, explains +absolutely nothing of the nature of bilification<a name="FNanchor_41_41" id="FNanchor_41_41"></a><a href="#Footnote_41_41" class="fnanchor">[41]</a>.</p></div> + +<p>Assuredly, it was needless for Owen to say that figurative language of +this kind explains nothing; but <span class="pagenum"><a name="page_334" id="page_334">[334]</a></span> +it was little less than puerile in him to see no more in the theory of +natural selection than such a mere figure of speech. To say that the +liver selects the elements of bile, or that nature selects specific +types, may both be equally unmeaning re-statements of facts; but when it +is explained that the term natural selection, unlike that of +“hepatic sensation,” is used as a shorthand expression for a +whole group of well-known natural causes—struggle, variation, +survival, heredity,—then it becomes evidence of an almost childish +want of thought to affirm that the expression is figurative and nothing +more. The doctrine of natural selection may be a huge mistake; but, if +so, this is not because it consists of any unmeaning metaphor: it can +only be because the combination of natural causes which it suggests is +not of the same adequacy in fact as it is taken to be in theory.</p> + +<p>Owen further objected that the struggle for existence could only act as +a cause of the extinction of species, not of their origination—a view +of the case which again shows on his part a complete failure to grasp +the conception of Darwinism. Acting alone, the struggle for existence +could only cause extermination; but acting together with variation, +survival, and heredity, it may very well—for anything that Owen, or +others who followed in this line of criticism, show to the +contrary—have produced every species of plant and animal that has ever +appeared upon the face of the earth.</p> + +<p>Another and closely allied objection is, that the theory of natural +selection “personifies an abstraction.” Or, as the Duke of +Argyll states it, the theory is “essentially the image of +mechanical necessity <span class="pagenum"><a name="page_335" id="page_335">[335]</a></span> +concealed under the clothes, and parading in the mask, of mental +purpose. The word ‘natural’ suggests Matter, and the +physical forces. The word ‘selection’ suggests Mind, and the +powers of choice.” This, however, is a mere quarrelling about +words. Darwin called the principle which he had discovered by the name +natural selection in order to mark the analogy between it and artificial +selection. No doubt in this analogy there is not necessarily supposed to +be in nature any counterpart to the mind of the breeder, nor, therefore, +to his powers of intelligent choice. But there is no need to limit the +term <i>selection</i> (<i>se</i> and <i>lego</i>, Gr. λέγω) to powers of +intelligent choice. As previously remarked, a bank of sea-weed on the +sea-shore may be said to have been selected by the waves from all the +surrounding sand and stones. Similarly, we may say that grain is +selected from chaff by the wind in the process of winnowing corn. Or, if +it be thought that there is any ambiguity involved in such a use of the +term in the case of “Natural Selection,” there is no +objection to employing the phrase which has been coined by Mr. Spencer +as its equivalent—namely, “Survival of the Fittest.” +The point of the theory is, that those organisms which are best suited +to their surroundings are allowed to live and to propagate, while those +which are less suited are eliminated; and whether we call this process a +process of selection, or call it by any other name, is clearly +immaterial.</p> + +<p>A material question is raised only when it is asked whether the process +is one that can be ascribed to causation strictly natural. It is often +denied that such is the case, on the ground that natural selection does +not originate the variations which it favours, <span class="pagenum"><a name="page_336" id="page_336">[336]</a></span> +but depends upon the variations being supplied by some other means. +For, it is said, all that natural selection does is to preserve the +suitable variations <i>after they have arisen</i>. Natural selection does not +<i>cause</i> these suitable variations; and therefore, it is argued, Darwin +and his followers are profoundly mistaken in representing the principle +as one which <i>produces</i> adaptations. Now, although this objection has +been put forward by some of the most intelligent minds in our +generation, it appears to me to betoken some extraordinary failure to +appreciate the very essence of Darwinian doctrine. No doubt it is +perfectly true that natural selection does not produce variations of any +kind, whether beneficial or otherwise. But if it be granted that +variations of many kinds are occurring in every generation, and that +natural selection is competent to preserve the more favourable among +them, then it appears to me unquestionable that this principle of +selection deserves to be regarded as, in the full sense of the word, a +natural cause. The variations being expressly regarded by the theory as +more or less promiscuous<a name="FNanchor_42_42" id="FNanchor_42_42"></a><a href="#Footnote_42_42" class="fnanchor">[42]</a>, +<span class="pagenum"><a name="page_337" id="page_337">[337]</a></span> +Survival of the fittest becomes the winnowing fan, whose function it is +to eliminate all the less fit in each generation, in order to preserve +the good grain, out of which to constitute the next generation. And as +this process is supposed to be continuous through successive +generations, its action is supposed to be cumulative, till from the eye +of a worm there is gradually developed the eye of an eagle. Therefore it +follows from these suppositions (which are not disputed by the present +objection), that if it had not been for the process of selection, such +development would never have been begun; and that in the exact measure +of its efficiency will the development proceed. But any agency without +the operation of which a result cannot take place may properly be +designated the cause of that result: it is the agency which, in +co-operation with all the other agencies in the cosmos, produces that +result.</p> + +<p><span class="pagenum"><a name="page_338" id="page_338">[338]</a></span> +Take any analogous case. The selective agency of specific gravity which +is utilised in gold-washing does not create the original differences +between gold-dust and dust of all other kinds. But these differences +being presented by as many different bodies in nature, the gold-washer +takes advantage of the selective agency in question, and, by using it as +a cause of segregation, is enabled to separate the gold from all the +earths with which it may happen to be mixed. So far as the objects of +the gold-washer are concerned, it is immaterial with what other earths +the gold-dust may happen to be mixed. For although gold-dust may occur +in intimate association with earths of various kinds in various +proportions, and although in each case the particular admixture which +occurs must have been due to definite causes, these things, in relation +to the selective process of the washer, are what is called accidental: +that is to say, they have nothing to do with the causative action of the +selective process. Now, in precisely the same sense Darwin calls the +multitudinous variations of plants and animals accidental. By so calling +them he expressly says he does not suppose them to be accidental in the +sense of not all being due to definite causes. But they are accidental +in relation to the sifting process of natural selection: all that they +have to do is to furnish the promiscuous material on which this sifting +process acts.</p> + +<p>Or let us take an even closer analogy. The power of selective breeding +by man is so wonderful, that in the course of successive generations all +kinds of peculiarities as to size, shape, colour, special appendages or +abortions, &c., can be produced at pleasure, <span class="pagenum"><a name="page_339" id="page_339">[339]</a></span> +as we saw in the last chapter. Now all the promiscuous variations which +are supplied to the breeder, and out of which, by selecting only those +that are suited to his purpose, he is able to produce the required +result—all those promiscuous variations, in relation to that +purpose, are accidental. Therefore the selective agency of the breeder +deserves to be regarded as the cause of that which it produces, or of +that which could not have been produced but for the operation of such +agency. But where is the difference between artificial and natural +selection in this respect? And, if there is no difference, is not +natural selection as much entitled to be regarded as a true cause of the +origin of natural species, as artificial selection is to be regarded as +a true cause of our domesticated races? Here, as in the case of the +previous illustration, if there be any ambiguity in speaking of +variations as accidental, it arises from the incorrect or undefined +manner in which the term “accidental” is used by +Darwin’s critics. In its original and philosophically-correct +usage, the term “accident” signifies a property or quality +not essential to our conception of a substance: hence, it has come to +mean anything that happens as a result of unforeseen causes—or, +lastly, that which is causeless. But, as we know that nothing can happen +without causes of some kind, the term “accident” is divested +of real meaning when it is used in the last of these senses. Yet this is +the sense that is sought to be placed upon it by the objection which we +are considering. If the objectors will but understand the term in its +correct philosophical sense—or in the only sense in which it +presents any meaning at all,—they will see that<span class="pagenum"><a name="page_340" id="page_340">[340]</a></span> +Darwinians are both logically and historically justified in employing +the word “accidental” as the word which serves most properly +to convey the meaning that they intend—namely, variations due to +causes accidental to the struggle for existence. Similarly, when it is +said that variations are “spontaneous,” or even +“fortuitous,” nothing further is meant than that we do not +know the causes which lead to them, and that, so far as the principle of +selection is concerned, it is immaterial what these causes may be. Or, +to revert to our former illustration, the various weights of different +kinds of earths are no doubt all due to definite causes; but, in +relation to the selective action of the gold-washer, all the different +weights of whatever kinds of earth he may happen to include in his +washing-apparatus are, <i>strictly speaking</i>, accidental. And as at +different washings he meets with different proportions of heavy earths +with light ones, and as these “variations” are immaterial to +him, he may colloquially speak of them as “fortuitous,” or +due to “chance,” even though he knows that at each washing +they must have been determined by definite causes.</p> + +<p>More adequately to deal with this merely formal objection, however, +would involve more logic-chopping than is desirable on the present +occasion. But I have already dealt with it fully elsewhere,—viz. in +<i>The Contemporary Review</i> for June, 1888, to which therefore I may refer +any one who is interested in dialectics of this kind<a name="FNanchor_43_43" id="FNanchor_43_43"></a><a href="#Footnote_43_43" class="fnanchor">[43]</a>.</p> + +<p><span class="pagenum"><a name="page_341" id="page_341">[341]</a></span> +I will now pass on to consider another misconception of the Darwinian +theory, which is very prevalent in the public mind. It is virtually +asked, If some species are supposed to have been improved by natural +selection, why have not all species been similarly improved? Why should +not all invertebrated <span class="pagenum"><a name="page_342" id="page_342">[342]</a></span> +animals have risen into vertebrated? Or why should not all monkeys have +become men?</p> + +<p>The answers are manifold. In the first place, it by no means follows +that because an advance in organization has proved itself of benefit in +the case of one form of life, therefore any or every other form would +have been similarly benefited by a similar advance. The business of +natural selection is to bring this and that form of life into the +closest harmony with its environment that all the conditions of the case +permit. Sometimes it will happen that the harmony will admit of being +improved by an improvement of organization. But just as often it will +happen that it will be best secured by leaving matters as they are. If, +therefore, an organism has already been brought into a tolerably full +degree of harmony with its environment, natural selection will not try +to change it so long as the environment remains unchanged; and this, no +doubt, is the reason why some species have survived through enormous +periods of geological time without having undergone any change. Again, +as we saw in a previous chapter, there are yet other cases where, on +account of some change in the environment or even in the habits of the +organisms themselves, adaption will be best secured by an active +<i>reversal</i> of natural selection, with the result of causing +<i>degeneration</i>.</p> + +<p>But, it is sometimes further urged, there are cases where we cannot +doubt that improvement of organization would have been of benefit to +species; and yet such improvement has not taken place—as, for instance, +in the case all monkeys not turning into men. Here, however, we must +remember that the operation <span class="pagenum"><a name="page_343" id="page_343">[343]</a></span> +of natural selection in any case depends upon a variety of highly +complex conditions; and, therefore, that the fact of all those +conditions having been satisfied in one instance is no reason for +concluding that they must also have been satisfied in other instances. +Take, for example, the case of monkeys passing into men. The wonder to +me appears to be that this improvement should have taken place in even +one line of descent; not that, having taken place in one line, it should +not also have taken place in other lines. For how enormously complex +must have been the conditions—physical, anatomical, physiological, +psychological, sociological—which by their happy conjunction first +began to raise the inarticulate cries of an ape into the rational speech +of a man. Therefore, the more that we appreciate the superiority of a +man to an ape, the less ought we to countenance this supposed objection +to Darwin’s theory—namely, that natural selection has not +effected the change in more than one line of descent.</p> + +<p>Even in the case of two races of mankind where one has risen higher in +the scale of civilization than another, it is now generally impossible +to assign the particular causes of the difference; much more, then, must +this be impossible in the case of still more remote conditions which +have led to the divergence of species. The requisite variations may not +have arisen in the one line of descent which did arise in the other; or +if they did arise in both, some counterbalancing disadvantages may have +attended their initial development in the one case which did not obtain +in the other. In short, where so exceedingly complex a play of +conditions are <span class="pagenum"><a name="page_344" id="page_344">[344]</a></span> +concerned, the only wonder would be if two different lines of descent +<i>had</i> happened to present two independent and yet perfectly parallel +lines of history.</p> + +<p>These general considerations would apply equally to the great majority +of other cases where some types have made great advances upon others, +notwithstanding that we can see no reason why the latter should not in +this respect have imitated the former. But there is yet a further +consideration which must be taken into account. The struggle for +existence is always most keen between closely allied species, because, +from the similarity of their forms, habits, needs, &c., they are in +closest competition. Therefore it often happens that the mere fact of +one species having made an advance upon others of itself precludes the +others from making any similar advance: the field, so to speak, has +already been occupied as regards that particular improvement, and where +the struggle for existence is concerned possession is emphatically nine +points of the law. For example, to return to the case of apes becoming +men, the fact of one rational species having been already evolved (even +if the rational faculty were at first but dimly nascent) must make an +enormous change in the conditions as regards the possibility of any +other such species being subsequently evolved—unless, of course, it be +by way of descent from the rational one. Or, as Sir Charles Lyell has +well put it, two rational species can never <i>coexist</i> on the globe, +although the descendants of one rational species may in time become +<i>transformed</i> into another single rational species<a name="FNanchor_44_44" id="FNanchor_44_44"></a><a href="#Footnote_44_44" class="fnanchor">[44]</a>.</p> + +<p>In view of such considerations, another and exactly <span class="pagenum"><a name="page_345" id="page_345">[345]</a></span> +opposite objection has sometimes been urged—viz. that we ought +never to find inferior forms of organization in company with superior, +because in the struggle for existence the latter ought to have +exterminated the former. Or, to quote the most recent expression of this +view, “in every locality there would only be one species, and that +the most highly organized; and thus a few superior races would partition +the earth amongst them to the entire exclusion of the innumerable +varieties, species, genera, and orders which now inhabit it<a +name="FNanchor_45_45" id="FNanchor_45_45"></a><a href="#Footnote_45_45" class="fnanchor">[45]</a>.” +Of course to this statement it would be sufficient to enquire, On what +would these few supremely organized species subsist? Unless manna fell +from heaven for their especial benefit, it would appear that such forms +could under no circumstances be the most improved forms; in +exterminating others on such a scale as this, they would themselves be +quickly, and very literally, improved off the face of the earth. But +even when the statement is not made in so extravagant a form as this, it +must necessarily be futile as an objection unless it has first been +shown that we know exactly all the conditions of the complex struggle +for existence between the higher and lower forms in question. And this +it is impossible that we ever can know. The mere fact that one form has +been changed in virtue of this struggle must in many cases of itself +determine a change in the conditions of the struggle. Again, the other +and closely allied forms (and these furnish the best grounds for the +objection) may also have undergone defensive changes, although these may +be less conspicuous to our observation, or perhaps less <span class="pagenum"><a name="page_346" id="page_346">[346]</a></span> +suggestive of “improvement” to our imperfect means of +judging. Lastly, not to continue citing an endless number of such +considerations, there is the broad fact that it is only to those cases +where, for some reason or another, the lower forms have not been exposed +to a struggle of fatal intensity, that the objection applies. But we +know that in millions of other cases the lower (i. e. less fitted) forms +<i>have</i> succumbed, and therefore I do not see that the objection has any +ground to stand upon. That there is a general tendency for lower forms +to yield their places to higher is shown by the gradual advance of +organization throughout geological time; for if <i>all</i> the inferior forms +had survived, the earth could not have contained them, unless she had +been continually growing into something like the size of Jupiter. And if +it be asked why any of the inferior forms have survived, the answer has +already been given, as above.</p> + +<p>There is only one other remark to be made in this connexion. Mr. Syme +chooses two cases as illustrations of the supposed difficulty. These are +sufficiently diverse—viz. Foraminifera and Man. Touching the former, +there is nothing that need be added to the general answer just given. +But with regard to the latter it must be observed that the dominion of +natural selection as between different races of mankind is greatly +restricted by the presence of rationality. Competition in the human +species is more concerned with wits and ideas than with nails and teeth; +and therefore the “struggle” between man and man is not so +much for actual <i>being</i>, as for <i>well-being</i>. Consequently, in regard to +the present objection, the <span class="pagenum"><a name="page_347" id="page_347">[347]</a></span> +Human species furnishes the worst example that could have been chosen.</p> + +<hr class='minor' /> + +<p>Hitherto I have been considering objections which arise from +misapprehensions of Darwin’s theory. I will now go on to consider +a logically sound objection, which nevertheless is equally futile, +because, although it does not depend on any misapprehension of the +theory, it is not itself supported by fact.</p> + +<p>The objection is the same as that which we have already considered in +relation to the general theory of descent—namely, that similar organs +or structures are to be met with in widely different branches of the +tree of life. Now this would be an objection fatal to the theory of +natural selection, supposing these organs or structures in the cases +compared are not merely analogous, but also homologous. For it would be +incredible that in two totally different lines of descent one and the +same structure should have been built up independently by two parallel +series of variations, and that in these two lines of descent it should +always and independently have ministered to the same function. On the +other hand, there would be nothing against the theory of natural +selection in the fact that two structures, <i>not</i> homologous, should come +by independent variation in two different lines of descent to be adapted +to perform the same function. For it belongs to the very essence of the +theory of natural selection that a useful function should be secured by +favourable variations of whatever structural material may happen to be +presented by different organic types. Flying, for instance, is a very +useful function, and it has been developed independently in at least +<span class="pagenum"><a name="page_348" id="page_348">[348]</a></span> +four different lines of descent—namely, the insects, reptiles, +birds, and mammals. Now if in all, or indeed in any, of these four cases +the wings had been developed on the same anatomical pattern, so as not +only to present the analogical resemblance which it is necessary that +they should present in order to discharge their common function of +flying, but likewise an homologous or structural resemblance, showing +that they had been formed on the same anatomical “plan,"—if +such has been the case, I say, the theory of natural selection would +certainly be destroyed.</p> + +<p>Now it has been alleged by competent naturalists that there are several +such cases in organic nature. We have already noticed in a previous +chapter (pp. <a href="#page_58">58</a>, <a href="#page_59">59</a>), that Mr. Mivart has instanced the eye of the +cuttle-fish as not only analogous to, but also homologous with, the eye +of a true fish—that is to say, the eye of a mollusk with the eye of a +vertebrate. And he has also instanced the remarkable resemblance of a +shrew to a mouse—that is, of an insectivorous mammal to a rodent—not +to mention other cases. In the chapter alluded to these instances of +homology, alleged to occur in different branches of the tree of life, +were considered with reference to the process of organic evolution as a +fact: they are now being considered with reference to the agency of +natural selection as a method. And just as in the former case it was +shown, that if any such alleged instances could be proved, the proof +would be fatal to the general theory of organic evolution by physical +causes, so in the present case, if this could be proved, it would be +equally fatal to the more special theory of natural selection. But, as +we have <span class="pagenum"><a name="page_349" id="page_349">[349]</a></span> +before seen, no single case of this kind has ever been made out; and, +therefore, not only does this supposed objection fall to the ground, but +in so doing it furnishes an additional argument in favour of natural +selection. For in the earlier chapter just alluded to I showed that this +great and general fact of our nowhere being able to find two homologous +structures in different branches of the tree of life, was the strongest +possible testimony in favour of the theory of evolution. And, by parity +of reasoning, I now adduce it as equally strong evidence of natural +selection having been the cause of <i>adaptive</i> structures, independently +developed in all the different lines of descent. For the alternative is +between adaptations having been caused by natural selection or by +supernatural design. Now, if adaptations were caused by natural +selection, we can very well understand why they should never be +homologous in different lines of descent, even in cases where they have +been brought to be so closely analogous as to have deceived so good a +naturalist as Mr. Mivart. Indeed, as I have already observed, so well +can we understand this, that any single instance to the contrary would +be sufficient to destroy the theory of natural selection <i>in toto</i>, +unless the structure be one of a very simple type. But on the other +hand, it is impossible to suggest any rational explanation why, if all +adaptations are due to supernatural design, such scrupulous care should +have been taken never to allow homologous adaptations to occur in +different divisions of the animal or vegetable kingdoms. Why, for +instance, should the eye of a cuttle-fish <i>not</i> have been constructed on +the same ideal pattern as that of vertebrate? Or why, <span class="pagenum"><a name="page_350" id="page_350">[350]</a></span> +among the thousands of vertebrated species, should no one of their eyes +be constructed on the ideal pattern that was devised for the +cuttle-fish? Of course it may be answered that perhaps there was some +hidden reason why the design should never have allowed an adaptation +which it had devised for one division of organic nature to appear in +another—even in cases where the new design necessitated the +closest possible resemblance in everything else, save in the matter of +anatomical homology. Undoubtedly such may have been the case—or +rather such <i>must</i> have been the case—if the theory of special +design is true. But where the question is as to the truth of this +theory, I think there can be no doubt that its rival gains an enormous +advantage by being able to <i>explain</i> why the facts are such as they are +instead of being obliged to take refuge in hypothetical possibilities of +a confessedly unsubstantiated and apparently unsubstantial kind.</p> + +<p>Therefore, as far as this objection to the theory of natural selection +is concerned—or the allegation that homologous structures occur in +different divisions of organic nature—not only does it fall to the +ground, but positively becomes itself converted into one of the +strongest arguments in favour of the theory. As soon as the allegation +is found to be baseless, the very fact that it cannot be brought to bear +upon any one of all the millions of adaptive structures in organic +nature becomes a fact of vast significance on the opposite side.</p> + +<hr class='minor' /> + +<p>The next difficulty to which I shall allude is that of explaining by the +theory of natural selection the preservation of the first beginnings of +structures which <span class="pagenum"><a name="page_351" id="page_351">[351]</a></span> +are then useless, though afterwards, when more fully developed, they +become useful. For it belongs to the very essence of the theory of +natural selection, that a structure must be supposed already useful +before it can come under the influence of natural selection: therefore +the theory seems incapable of explaining the origin and conservation of +<i>incipient</i> organs, or organs which are not yet sufficiently developed +to be of any service to the organisms presenting them.</p> + +<p>This objection is one that has been advanced by all the critics of +Darwinism; but has been presented with most ability and force by the +Duke of Argyll. I will therefore state it in his words.</p> + +<div class='blockquot'><p>If the doctrine of evolution be true—that is to say, if all +organic creatures have been developed by ordinary generation from +parents—then it follows of necessity that the primæval germs must +have contained potentially the whole succeeding series. Moreover, +if that series has been developed gradually and very slowly, it +follows, also as a matter of necessity, that every modification of +structure must have been functionless at first, when it began to +appear.... Things cannot be selected until they have first been +produced. Nor can any structure be selected by utility in the +struggle for existence until it has not only been produced, but has +been so far perfected as to actually be used.</p></div> + +<p>The Duke proceeds to argue that all adaptive structures must therefore +originally have been due to special design: in the earlier stages of +their development they must all have been what he calls “prophetic +germs.” Not yet themselves of any use, and therefore not yet +capable of being improved by natural selection, both in their origin and +in the first stages (at all events) of their development, they must +<span class="pagenum"><a name="page_352" id="page_352">[352]</a></span> +be regarded as intentionally preparatory to the various uses which they +subsequently acquire.</p> + +<p>Now this argument, forcible as it appears at first sight, is really at +fault both in its premiss and in its conclusion. By which I mean that, +in the first place the premiss is not true, and, in the next place, that +even if it were, the conclusion would not necessarily follow. The +premiss is, “that every modification of structure must have been +functionless at first, when it began to appear;” and the +conclusion is, that, <i>quâ</i> functionless, such a modification cannot have +been caused by natural selection. I will consider these two points +separately.</p> + +<p>First as to the premiss, it is not true that every modification of +structure must necessarily be functionless when it first begins to +appear. There are two very good reasons why such should not be the case +in all instances, even if it should be the case in some. For, as a +matter of observable fact, a very large proportional number of incipient +organs are useful from the very moment of their inception. Take, for +example, what is perhaps the most wonderful instance of refined +mechanism in nature—the eye of a vertebrated animal. Comparative +anatomy and embryology combine to testify that this organ had its origin +in modifications of the endings of the ordinary nerves of the skin. Now +it is evident that from the very first any modification of a cutaneous +nerve whereby it was rendered able, in however small a degree, to be +differently affected by light and by darkness would be of benefit to the +creature presenting it; for the creature would thus be able to seek the +one and shun the other according to the requirements of its life. +<span class="pagenum"><a name="page_353" id="page_353">[353]</a></span> +And being thus useful from the very moment of its inception, it would +afterwards be gradually improved as variations of more and more utility +presented themselves, until not only would finer and finer degrees of +difference between light and shade become perceptible, but even the +outlines of solid bodies would begin to be appreciated. And so on, stage +by stage, till from an ordinary nerve-ending in the skin is evolved the +eye of an eagle.</p> + +<p>Moreover, in this particular instance there is very good reason to +suppose that the modification of the cutaneous nerves in question began +by a progressive increase in their sensitiveness to temperature. +Wherever dark pigment happened to be deposited in the skin—and we know +that in all animals it is apt to be deposited in points and patches, as +it were by accident, or without any “prophecy” as to future +uses,—the cutaneous nerves in its vicinity would be better able to +appreciate the difference between sun and shade in respect of +temperature, even though as yet there were no change at all in these +cutaneous nerves tending to make them responsive to light. Now it is +easy to see how, from such a purely accidental beginning, natural +selection would have had from the first sufficient material to act upon. +It being of advantage to a lowly creature that it should distinguish +with more and more delicacy, or with more and more rapidity, between +light and darkness by means of its thermal sensations, the pigment spots +in the skin would be rendered permanent by natural selection, while the +nerves in that region would by the same agency be rendered more and more +specialized as organs adapted to perceive changes of temperature, until +<span class="pagenum"><a name="page_354" id="page_354">[354]</a></span> +from the stage of responding to the thermal rays of the non-luminous +spectrum alone, they become capable of responding also to luminous.</p> + +<p>So much, then, for the first consideration which serves to invalidate +the Duke’s premiss. The second consideration is, that very often +an organ which began by being useful for the performance of one +function, after having been fully developed for the performance of that +function, finds itself, so to speak, accidentally fitted to the +performance of some other and even more important function, which it +thereupon begins to discharge, and so to undergo a new course of +adaptive development. In such cases, and so far as the new function is +concerned, the difficulty touching the first inception of an organ does +not apply; for here the organ has already been built up by natural +selection for one purpose, before it begins to discharge the other. As +an example of such a case we may take the lung of an air-breathing +animal. Originally the lung was a swim-bladder, or float, and as such it +was of use to the aquatic ancestors of terrestrial animals. But as these +ancestors gradually became more and more amphibious in their habits, the +swim-bladder began more and more to discharge the function of a lung, +and so to take a wholly new point of departure as regards its +developmental history. But clearly there is here no difficulty with +regard to the inception of its new function, because the organ was +already well developed for one purpose before it began to serve another. +Or, to take only one additional example, there are few structures in the +animal kingdom so remarkable in respect of adaptation as is the wing of +a bird or a bat; and at first sight it might well appear <span class="pagenum"><a name="page_355" id="page_355">[355]</a></span> +that a wing could be of no conceivable use until it had already +acquired enormous proportional dimensions, as well as an immense amount +of special elaboration as to its general form, size of muscle, amount of +blood-supply, and so on. For, obviously, not until it had attained all +these things could it even begin to raise the animal in the air. But +observe how fallacious is this argument. Although it is perfectly true +that a wing could be of no use <i>as a wing</i> until sufficiently developed +to serve the purpose of flight, this is merely to say that until it has +become a wing it is no use as a wing. It does not, however, follow that +on this account it was of no prior use for any other purpose. The first +modifications of the fore-limb which ended in its becoming an organ of +flight may very well have been due to adapting it as an organ for +increased rapidity of locomotion of other kinds—whether on land as +in the case of its now degenerated form in the ostrich, or in water as +in the case of the expanded fins of fish. Indeed, we may see the actual +process of transition from the one function to the other in the case of +“flying-fish.” Here the progressive expansion of the +pectoral fins must certainly have been always of use for continuously +promoting rapidity of locomotion through water; and thus natural +selection may have continuously increased their development until they +now begin to serve also as wings for carrying the animal a short +distance through air. Again, in the case of the so-called flying +squirrels we find the limbs united to the body by means of large +extensions of the skin, so-that when jumping from one tree to another +the animal is able to sustain itself through a long distance in the air +by merely spreading out its <span class="pagenum"><a name="page_356" id="page_356">[356]</a></span> +limbs, and thus allowing the skin-extensions to act after the manner of +a parachute. Here, of course, we have not yet got a wing, any more than +we have in the case of the flying-fish; but we have the foundations laid +for the possible development of a future wing, upon a somewhat similar +plan as that which has been so wonderfully perfected in the case of +bats. And through all the stages of progressive expansion which the skin +of the squirrel has undergone, the expansion has been of use, even +though it has not yet so much as begun to acquire the distinctive +functions of a wing. Here, then, there is obviously nothing +“prophetic” in the matter, any more than there was in the +case of the swim-bladder and the lung, or in that of the nerve-ending +and the eye. In short, it is the business of natural selection to secure +the highest available degree of adaptation for the time being; and, in +doing this, it not unfrequently happens that an extreme development of a +structure in one direction (produced by natural selection for the sake +of better and better adapting the structure to perform some particular +function) ends by beginning to adapt it to the performance of some other +function. And, whenever this happens to be the case, natural selection +forthwith begins to act upon the structure, so to speak, from a new +point of departure.</p> + +<p>So much, then, for the Duke’s premiss—namely, that “every +modification of structure <i>must</i> have been functionless <i>at first</i>, when +it began to appear.” This premiss is clearly opposed to observable +fact. But now, the second position is that, even if this were not so, +the Duke’s conclusion would not follow. This conclusion, it will +be remembered, is, that if incipient <span class="pagenum"><a name="page_357" id="page_357">[357]</a></span> +structures are useless, it necessarily follows that natural selection +can have had no part whatever in their inception. Now, this is a +conclusion which does not “necessarily” follow. Even if it +be granted that there are structures which in their first beginnings are +not of any use at all for any purpose, it is still possible that they +may owe their origin to natural selection—not indeed directly, but +indirectly. This possibility arises from the occurrence in nature of a +principle which has been called the Correlation of Growth.</p> + +<p>Mr. Darwin, who has paid more attention to this matter than any other +writer, has shown, in considerable detail, that all the parts of any +given organism are so intimately bound together, or so mutually +dependent upon each other, that when one part is caused to change by +means of natural selection, some other parts are very likely to undergo +modification as a consequence. For example, there are several kinds of +domesticated pigeons and fowls, which grow peculiar wing-like feathers +on the feet. These are quite unlike all the other feathers in the +animal, except those of the wing, to which they bear a very remarkable +resemblance. Mr. Darwin records the case of a bantam where these +wing-like feathers were nine inches in length, and I have myself seen a +pigeon where they reproduced upon the feet a close imitation of the +different kinds of feathers which occupy homologous positions in the +wing—primaries, secondaries, and tertiaries all being distinctly +repeated in their proper anatomical relations. Furthermore, in this +case, as in most cases where such wing-feathers occur upon the feet, the +third and fourth toes were partly united by skin; and, as is well known, +in the wing <span class="pagenum"><a name="page_358" id="page_358">[358]</a></span> +of a bird the third and fourth digits are completely united by skin; +“so that in feather-footed pigeons, not only does the exterior +surface support a row of long feathers, like wing-feathers [which, as +just stated, may in some cases be obviously differentiated into +primaries, secondaries and tertiaries], but the very same digits which +in the wing are completely united by skin become partially united by +skin in the feet; and thus by the law of correlated variation of +homologous parts, we can understand the curious connexion of feathered +legs and membrane between the two outer toes<a name="FNanchor_46_46" id="FNanchor_46_46"></a><a href="#Footnote_46_46" class="fnanchor">[46]</a>.” The +illustration is drawn from the specimen to which I have referred.</p> + +<p class="pagenum"><a name="page_359i" id="page_359i">[359i.]</a></p> +<div class="figcenter" style="width: 392px;"> +<img src="images/i_378_117.jpg" width="392" height="500" alt="Feather-footed pigeon." title="" /> +<span class="caption"><span class="sc">Fig.</span> 117.—Feather-footed pigeon. Drawn from nature.</span></div> + +<p><span class="pagenum"><a name="page_358c" id="page_358c">[358c.]</a></span> +Many similar instances of the same law are to be met with throughout +organic nature; and it is evident that in this principle we find a +conceivable explanation of the origin of such adaptive structures as +could not have been originated by natural selection acting directly upon +themselves: they may have been originated by natural selection +developing other adaptive structures elsewhere in the organism, the +gradual evolution of which has entailed the production of these by +correlation of growth. And, if so, when once started in this way, these +structures, because thus accidentally useful, will now themselves come +under the <i>direct</i> action of natural selection, and so have their +further evolution determined with or without the correlated association +which first led to their inception.</p> + +<p>Of course it must be understood that in thus applying the principle of +correlated growth, to explain the origin of adaptive structures where it +is impossible to explain such origin by natural selection having from +<span class="pagenum"><a name="page_359" id="page_359">[359]</a></span> +the first acted directly upon these structures themselves, Darwinists +do not suppose that in all—or even in most—cases of +correlated growth the correlated structures are of use. On the contrary, +it is well known that structures due to correlated growth are, as a +rule, useless. Being only the by-products of adaptive changes going on +elsewhere, in any given case the chances are against these correlated +effects being themselves of any utilitarian significance; and, +therefore, as a matter of fact, correlated growths appear to be usually +meaningless from the point of <span class="pagenum"><a name="page_360" id="page_360">[360]</a></span> +view of adaptation. Still, on the doctrine of chances, it is to be +expected that sometimes a change of structure which has thus been +indirectly produced by correlation of growth might happen to prove +useful for some purpose or another; and in as many cases as such +indirectly produced structures do prove useful, they will straightway +begin to be improved by the direct action of natural selection. In all +such cases, therefore, we should have an explanation of the <i>origin</i> of +such a structure, which is the only point that we are now considering.</p> + +<p>I think, then, that all this effectually disposes of the doctrine of +“prophetic germs.” But, before leaving the subject, I should +like to make one further statement of greater generality than any which +I have hitherto advanced. This statement is, that we must remember how +large a stock of meaningless structures are always being produced in the +course of specific transmutations, not only by correlation of growth, +which we have just been considering, but also by the direct action of +external conditions, together with the constant play of all the many and +complex forces internal to organisms themselves. In other words, +important as the principle of correlation undoubtedly is, we must +remember that even this is very far from being the only principle which +is concerned in the origination of structures that may or may not chance +to be useful. Therefore, it is not only natural selection when operating +indirectly through the correlation of growth that is competent to +produce new structures without reference to utility. In all the complex +action and reaction of internal and external forces, new variations are +perpetually arising <span class="pagenum"><a name="page_361" id="page_361">[361]</a></span> +without any reference to utility, either present or future. Among all +this multitude of promiscuous variations, the chances must be that some +percentage will prove of some service, either from the first moment of +their appearance, or else after they have undergone some amount of +development. Such development prior to utility may be due, either to +correlation of growth, to the structure having previously performed some +other function, as already explained, or else to a continued operation +of the causes which were concerned in the first appearance of originally +useless characters. In a series of chapters which will be devoted to the +whole question of utility in the next volume, I shall hope to give very +good reasons for concluding that useless characters are not only of +highly frequent occurrence, but are due to a variety of other causes +besides correlation of growth. And, if so, the possibility of originally +useless characters happening in some cases to become, by increased +development, useful characters, is correspondingly increased. Among a +hundred varietal or specific characters which are directly produced in +as many different species by a change of climate, for example, some five +or six may be <i>potentially</i> useful: that is to say, characters thus +adventitiously produced in an incipient form may only require to be +further developed by a continuance of the same causes as first +originated them, in order that some percentage of the whole number shall +become of some degree of use. Those professed followers of Darwin, +therefore, who without any reason—or, as it appears to me, against +all reason—deny the possibility of useless specific characters in +any case or in any degree (unless correlated with useful characters), +<span class="pagenum"><a name="page_362" id="page_362">[362]</a></span> +are playing into the hands of Darwin’s critics by indirectly +countenancing the difficulty which we are now considering. For, if +correlation of growth is unreasonably supposed to be the only possible +cause of the origin of incipient structures which are not useful from +the first moment of their inception, clearly the field is greatly +narrowed as regards the occurrence of incipient characters sufficient in +amount—and, still more, in constancy of appearance and persistency +of transmission—to admit of furnishing material for the working of +natural selection. But in the measure that incipient +characters—whether varietal or specific—are recognised as +not always or “necessarily” useful from the moment of their +inception, and yet capable of being developed to a certain extent by the +causes which first led to their occurrence, in that measure is this line +of criticism closed. For of all the variations which thus occur, it is +only those which afterwards prove of any use that are laid hold upon and +wrought up by natural selection into adaptive structures, or working +organs. And, therefore, what we see in organic nature is the net outcome +of the development of all the happy chances. So it comes that the +appearance presented by organic nature as a whole is that of a continual +fulfilment of structural prophecies, when, in point of fact, if we had a +similar record of all the other variations it would be seen that +possibly not one such prophecy in a thousand is ever destined to be +fulfilled.</p> + +<hr class='minor' /> + +<p>Here, then, I feel justified in finally taking leave of the difficulty +from the uselessness of incipient organs, as this difficulty has been +presented, in varying degrees <span class="pagenum"><a name="page_363" id="page_363">[363]</a></span> +of emphasis, by the Duke of Argyll, Mr. Mivart, Professors Nägeli, +Bronn, Broca, Eimer, and, indeed, by all other writers who have hitherto +advanced it. For, as thus presented, I think I have shown that it admits +of being adequately met. But now, I must confess, to me individually it +does appear that behind this erroneous presentation of the difficulty +there lies another question, which is deserving of much more serious +attention. For although it admits of being easily shown—as I have +just shown—that the difficulty as ordinarily presented fails on +account of its extravagance, the question remains whether, if stated +with more moderation, a real difficulty might not be found to remain.</p> + +<p>My quarrel with the conclusion, like my quarrel with the premiss, is due +to its universality. By saying in the premiss that <i>all</i> incipient +organs are <i>necessarily</i> useless at the time of their inception, these +writers admit of being controverted by fact; and by saying in the +conclusion that, <i>if</i> all incipient organs are useless, it necessarily +follows that in <i>no</i> case can natural selection have been the cause of +building up an organ until it becomes useful, they admit of being +controverted by logic. For, even if the premiss were true in +fact—namely, that all incipient organs are useless at the time of their +inception,—it would not necessarily follow that in no case could +natural selection build up a useless structure into a useful one; +because, although it is true that in no case can natural selection do +this by acting on a useless structure <i>directly</i>, it may do so by acting +on the useless structure <i>indirectly</i>, through its direct action on some +other part of the organism with which the useless structure happens to +be correlated. Moreover, as I believe, and <span class="pagenum"><a name="page_364" id="page_364">[364]</a></span> +will subsequently endeavour to prove, there is abundant evidence to +show that incipient characters are often developed to a large extent by +causes other than natural selection (or apart from any reference to +utility), with the result that some of them thus happen to become of +use, when, of course, the supposed difficulty is at an end.</p> + +<p>But although it is thus easy to dispose of both the propositions in +question, on account of their universality, stated more carefully they +would require, as I have said, more careful consideration. Thus, if it +had been said that some incipient organs are <i>presumably</i> useless at the +time of their inception, and that in <i>some of these cases</i> it is +difficult, or impossible, to conceive how the principle of correlation, +or any other principle hitherto suggested, can apply—then the question +would have been raised from the sphere of logical discussion to that of +biological fact. And the new question thus raised would have to be +debated, no longer on the ground of general or abstract principles, but +on that of special or concrete cases. Now until within the last year or +two it has not been easy to find such a special or concrete case—that +is to say, a case which can be pointed to as apparently excluding the +possibility of natural selection having had anything to do with the +genesis of an unquestionably adaptive structure. But eventually such a +case has arisen, and the Duke of Argyll has not been slow in perceiving +its importance. This case is the electric organ in the tail of the +skate. No sooner had Professor Cossar Ewart published an abstract of his +first paper on this subject, than the Duke seized upon it as a case for +which, as he said, he had long been waiting—namely, <span class="pagenum"><a name="page_365" id="page_365">[365]</a></span> +the case of an <i>adaptive</i> organ the genesis of which <i>could not +possibly</i> be attributed to natural selection, and must therefore be +attributed to supernatural design. Now, I do not deny that he is here in +possession of an admirable case—a case, indeed, so admirable that +it almost seems to have been specially designed for the discomfiture of +Darwinians. Therefore, in order to do it full justice, I will show that +it is even more formidable than the Duke of Argyll has represented.</p> + +<p>Electric organs are known to occur in several widely different kinds of +fish—such as the <span class='sn'>Gymnotus</span> and <span class='sn'>Torpedo</span>. +Wherever these organs do occur, they perform the function of electric +batteries in storing and discharging electricity in the form of more or +less powerful shocks. Here, then, we have a function which is of obvious +use to the fish for purposes both of offence and defence. These organs +are everywhere composed of a transformation of muscular, together with +an enormous development of nervous tissue; but inasmuch as they occupy +different positions, and are also in other respects dissimilar in the +different zoological groups of fishes where they occur, no difficulty +can be alleged as to these analogous organs being likewise homologous in +different divisions of the aquatic vertebrata.</p> + +<p>Now, in the particular case of the skate, the organ is situated in the +tail, where it is of a spindle-like form, measuring, in a large fish, +about two feet in length by about an inch in diameter at the middle of +the spindle. Although its structure is throughout as complex and perfect +as that of the electric organ in <span class='sn'>Gymnotus</span> or <span class='sn'>Torpedo</span>, +its smaller size does not admit <span class="pagenum"><a name="page_366" id="page_366">[366]</a></span> +of its generating a sufficient amount of electricity to yield a +discharge that can be felt by the hand. Nevertheless, that it does +discharge under suitable stimulation has been proved by Professor Burdon +Sanderson by means of a telephone; for he found that every time he +stimulated the animal its electrical discharge was rendered audible by +the telephone. Here, then, the difficulty arises. For of what +conceivable use is such an organ to its possessor? We can scarcely +suppose that any aquatic animal is more sensitive to electric shocks +than is the human hand; and even if such were the case, a discharge of +so feeble a kind taking place in water would be short-circuited in the +immediate vicinity of the skate itself. So there can be no doubt that +such weak discharges as the skate is able to deliver must be wholly +imperceptible alike to prey and to enemies. Yet for the delivery of such +discharges there is provided an organ of such high peculiarity and huge +complexity, that, regarded as a piece of living mechanism, it deserves +to rank as at once the most extremely specialized and the most highly +elaborated structure in the whole animal kingdom. Thousands of +separately formed elements are ranged in row after row, all electrically +insulated one from another, and packed away into the smallest possible +space, with the obvious end, or purpose, of conspiring together for the +simultaneous delivery of an electric shock. Nevertheless, the shock when +delivered is, as we have just seen, too slight to be of any conceivable +use to the skate. Therefore it appears impossible to suggest how this +astonishing structure—much more astonishing, in my opinion, than +the human eye or the human hand—can ever <span class="pagenum"><a name="page_367" id="page_367">[367]</a></span> +have been begun, or afterwards developed, by means of natural +selection. For if it be not even yet of any conceivable use to its +possessor, clearly thus far survival of the fittest can have had nothing +to do with its formation. On the other hand, seeing that electric organs +when of larger size, as in the <span class='sn'>Gymnotus</span> and <span class='sn'>Torpedo</span>, +are of obvious use to their possessors, the facts of the case, so far as +the skate is concerned, assuredly do appear to sanction the doctrine of +“prophetic germs.” The organ in the skate seems to be on its +way towards becoming such an organ as we meet with in these other +animals; and, therefore, unless we can show that it is now, and in all +previous stages of its evolution has throughout been, of use to the +skate, the facts do present a serious difficulty to the theory of +natural selection, while they readily lend themselves to the +interpretation of a disposing or fore-ordaining mind, which knows how to +construct an electric battery by thus transforming muscular tissue into +electric tissue, and is now actually in process of constructing such an +apparatus for the prospective benefit of future creatures.</p> + +<p>Should it be suggested that possibly the electric organ of the skate may +be in process of degeneration, and therefore that it is now the +practically functionless remnant of an organ which in the ancestors of +the skate was of larger size and functional use—against so obvious a +suggestion there lie the whole results of Professor Ewart’s +investigations, which go to indicate that the organ is here not in a +stage of degeneration, but of evolution. For instance, in <span class='sn'>Raia radiata</span>, +it does not begin to be formed out of the muscular tissue until some +time after the animal has left the egg-capsule, <span class="pagenum"><a name="page_368" id="page_368">[368]</a></span> +and assumed all the normal proportions (though not yet the size) of the +adult creature. The organ, therefore, is one of the very latest to +appear in the ontogeny of <span class='sn'>R. radiata</span>; and, moreover, it does not attain +its full <i>development</i> (i. e. not merely <i>growth</i>, but transforming of +muscular fibres into electrical elements) till the fish attains +maturity. Read in the light of embryology, these facts prove, (1) that +the electric organ of <span class='sn'>R. radiata</span> must be one of the very <span class="pagenum"><a +name='page_370' id='page_370'>[370]</a></span>latest products of the +animal’s phylogeny; and, (2) that as yet, at all events, it has +not begun to degenerate. But, if not, it must either be at a +stand-still, or it must be in course of further evolution; and, +whichever of these alternatives we adopt, the difficulty of accounting +for its present condition remains. In this connexion also it is worth +while to remark that the electric <span class="pagenum"><a name="page_371" id="page_371">[371]</a></span> +organ, even after it has attained its full <i>development</i>, continues its +<i>growth</i> with the growth of the fish, and this in a much higher ratio, +either than the tail alone, or the whole animal. Lastly, Prof. Burdon +Sanderson finds that <i>section for section</i> the organ in the skate is as +efficient as it is in <span class='sn'>Torpedo</span>. It is evident that these facts also point +to the skate’s organ being in course of phylogenetic evolution.</p> + +<p class="pagenum"><a name="page_368i" id="page_368i">[368i.]</a></p> +<div class="figcenter" style="width: 445px;"> +<img src="images/i_387_118.jpg" width="445" height="500" alt="Raia radiata, representing the life size of +the youngest individual in which muscle fibres have been found +developing into electric cells." title="" /> +<div class="captionh"><span class="sc">Fig.</span> 118.—<span class='sn'>Raia radiata</span>, representing the life size of +the youngest individual in which muscle fibres have been found +developing into electric cells.</div></div> + +<p class="pagenum"><a name="page_369" id="page_369">[369]</a></p> +<div class="figcenter" style="width: 313px;"> +<img src="images/i_388_119.jpg" width="313" height="500" alt="Electric organ of the Skate." title="" /> +<div class="captionh"><span class="sc">Fig.</span> 119.—Electric organ of the Skate. The left-hand +drawing (I) represents the entire organ (natural size) of a full-grown +<span class='sn'>R. radiata</span>. This is a small skate, which rarely exceeds 50 centms. in +length; but in the large <span class='sn'>R. batis</span>, the organ may exceed two feet in +length. The other drawings represent single muscle-fibres in successive +stages of transition. In the first of the series (II) the motor plate, +and the nerves connected with it, have already been considerably +enlarged. In the other three specimens, the fibre becomes more and more +club-like, and eventually cup-like. These changes of shape are +expressive of great changes of structure, as may be seen in the last of +the series (V), where the shallow cup is seen in partial section. The +electric plate lines the concavity of the cup, and is richly supplied +with nerves (only a few of which are represented in the last drawing); +the thick walls of the cup are composed of muscular fibres, the +striation of which is distinctly visible.</div></div> + +<p class="pagenum"><a name="page_370i" id="page_370i">[370i.]</a></p> +<div class="figcenter" style="width: 376px;"> +<img src="images/i_389_120.jpg" width="376" height="500" alt="Electric cells of Raia radiata." title="" /> +<div class="captionh"><span class="sc">Fig.</span> 120.—Electric cells of <span class='sn'>Raia radiata</span>. The drawing on +the left represents one of the clubs magnified, as in the preceding +wood-cut. The drawing on the right represents a number of these clubs, +less highly magnified, <i>in situ</i>.</div></div> + +<p><span class="pagenum"><a name="page_371c" id="page_371c">[371c.]</a></span> +Again, it cannot be answered that the principle of correlation may be +drawn upon in mitigation of the difficulty. The structure of the +electric organ is far too elaborate, far too specialized, and far too +obviously directed to a particular end, to admit of our conceivably +supposing it due to any accidental correlation with structural changes +going on elsewhere. Even as regards the initial changes of +muscle-elements into electrical-elements, I do not think the principle +of correlation can be reasonably adduced by way of explanation; for, as +shown in the illustrations, even this initial change is most +extraordinarily peculiar, elaborate, and specialized. But, be this as it +may, I am perfectly certain that the principle of correlation cannot +possibly be adduced to explain the subsequent <i>association of these +electrical elements into an electric battery</i>, actuated by a special +nervous mechanism of enormous size and elaboration—unless of course, +the progress of such a structure were assumed to have been throughout of +some utility. Under this supposition, however, the principle of +correlation would be forsaken in favour of that of natural selection; +and we should again be in the presence of the same difficulty as that +with which we started.</p> + +<p>But now, and further, if we do thus abandon correlation <span class="pagenum"><a name="page_372" id="page_372">[372]</a></span> +in favour of natural selection, and therefore if for the sake of saving +an hypothesis we assume that the organ as it now stands <i>must</i> be of +some use to the existing skate, we should still have to face the +question—Of what conceivable use can those initial stages of its +formation have been, when first the muscle-elements began to be changed +into the very different electrical-elements, and when therefore they +became useless as muscles while not yet capable of performing even so +much of the electrical function as they now perform?</p> + +<p>Lastly, we must remember that not only have we here the most highly +specialized, the most complex, and altogether the most elaboratively +adaptive organ in the animal kingdom; but also that in the formation of +this structure there has been needed an altogether unparalleled +expenditure of the most physiologically expensive of all +materials—namely, nervous tissue. Whether estimated by volume or by +weight, the quantity of nervous tissue which is consumed in the electric +organ of the skate is in excess of all the rest of the nervous system +put together. It is needless to say that nowhere else in the animal +kingdom—except, of course, in other electric fishes—is there any +approach to so enormous a development of nervous tissue for the +discharge of a special function. Therefore, as nervous tissue is, +physiologically speaking, the most valuable of all materials, we are +forced to conclude that natural selection ought strongly to have +<i>opposed</i> the evolution of such organs, unless from the first moment of +their inception, and throughout the whole course of their development, +they were of some such paramount importance as biologically to justify +so unexampled an expenditure. Yet this paramount importance <span class="pagenum"><a name="page_373" id="page_373">[373]</a></span> +does not admit of being so much as surmised, even where the organ has +already attained the size and degree of elaboration which it presents in +the skate.</p> + +<p>In view of all these considerations taken together, I freely confess +that the difficulty presented by this case appears to me of a magnitude +and importance altogether unequalled by that of any other single +case—or any series of cases—which has hitherto been encountered by the +theory of natural selection. So that, if there were many other cases of +the like kind to be met with in nature, I should myself at once allow +that the theory of natural selection would have to be discarded. But +inasmuch as this particular case stands so far entirely by itself, and +therefore out of analogy with thousands, or even millions, of other +cases throughout the whole range of organic nature, I am constrained to +feel it more probable that the electric organ of the skate will some day +admit of being marshalled under the general law of natural selection—in +just the same way as proved to be the case with the conspicuous +colouring of those caterpillars, which, as explained in the last +chapter, at one time seemed to constitute a serious difficulty to the +theory, and yet, through a better knowledge of all the relations +involved, has now come to constitute one of the strongest witnesses in +its favour.</p> + +<hr class='minor' /> + +<p>I have now stated all the objections of any importance which have +hitherto been brought against the theory of natural selection, excepting +three, which I left to be dealt with together because they form a +logically connected group. With a brief consideration of these, +therefore, I will bring this chapter to a close.</p> + +<p><span class="pagenum"><a name="page_374" id="page_374">[374]</a></span> +The three objections to which I allude are, (1) that a large +proportional number of specific, as well as of higher taxonomic +characters, are seemingly useless characters, and therefore do not lend +themselves to explanation by the Darwinian theory; (2) that the most +general of all specific characters—viz. cross-infertility between +allied species—cannot possibly be due to natural selection, as is +demonstrated by Darwin himself; (3) that the swamping effects of free +intercrossing must always render impossible by natural selection alone +any evolution of species in divergent (as distinguished from serial) +lines of change.</p> + +<p>These three objections have been urged from time to time by not a few of +the most eminent botanists and zoologists of our century; and from one +point of view I cannot myself have the smallest doubt that the +objections thus advanced are not only valid in themselves, but also by +far the most formidable objections which the theory of natural selection +has encountered. From another point of view, however, I am equally +convinced that they all admit of absolute annihilation. This strong +antithesis arises, as I have said, from differences of standpoint, or +from differences in the view which we take of the theory of natural +selection itself. If we understand this theory to set forth natural +selection as the sole cause of organic evolution, then all the above +objections to the theory are not merely, as already stated, valid and +formidable, but as I will now add, logically insurmountable. On the +other hand, if we take theory to consist merely in setting forth natural +selection as a factor of organic evolution, even although we believe +<span class="pagenum"><a name="page_375" id="page_375">[375]</a></span> +it to have been the chief factor or principal cause, all the three +objections in question necessarily vanish. For in this case, even if it +be satisfactorily proved that the theory of natural selection is unable +to explain the three classes of facts above mentioned, the theory is not +thereby affected: facts of each and all of these classes may be +consistently left by the theory to be explained by causes other than +natural selection—whether these be so far capable or incapable of +hypothetical formulation. Thus it is evident that whether the three +objections above named are to be regarded as logically insurmountable by +the theory, or as logically non-existent in respect to it, depends +simply upon the manner in which the theory itself is stated.</p> + +<p>In the next volume a great deal more will have to be said upon these +matters—especially with regard to the causes other than natural +selection which in my opinion are capable of explaining these so-called +“difficulties.” In the present connexion, however, all I +have attempted to show is, that, whatever may be thought touching the +supplementary theories whereby I shall endeavour to explain the facts of +inutility, cross-sterility, and non-occurrence of free intercrossing, no +one of these facts is entitled to rank as an objection against the +theory of natural selection, unless we understand this theory to claim +an exclusive prerogative in the field of organic evolution. This, as we +have previously seen, is what Mr. Wallace does claim for it; while on +the other hand, Mr. Darwin expressly—and even vehemently—repudiates +the claim: from which it follows that all the three main objections +against the theory of natural selection <span class="pagenum"><a name="page_376" id="page_376">[376]</a></span> +are objections which vitally affect the theory only as it has been +stated and upheld by Wallace. As the theory has been stated and upheld +by Darwin, all these objections are irrelevant. This is a fact which I +had not myself perceived at the time when I mentioned these objections +in a paper entitled <i>Physiological Selection</i>, which was published in +1886. The discussions to which that paper gave rise, however, led me to +consider these matters more closely; and further study of Darwin’s +writings, with these matters specially in view, has led me to see that +none of the objections in question are relevant to his theory, as +distinguished from that of Mr. Wallace. This, I acknowledge, I ought to +have perceived before I published the paper just alluded to; but in +those days I had had no occasion to follow out the differences between +Darwin and Wallace to all their consequences, and therefore adopted the +prevalent view that their theories of evolution were virtually +identical. Now, however, I have endeavoured to make it clear that the +points wherein they differ involve the important consequences above set +forth. All these the most formidable objections against the theory of +natural selection arise simply and solely from what I conceive to be the +erroneous manner in which the theory has been presented by +Darwin’s distinguished colleague.</p> + +<hr class='minor' /> + +<p>I have now considered, as impartially as I can, all the main criticisms +and objections which have been brought against the theory of natural +selection; and the result is to show that, neither singly nor +collectively, are they entitled to much weight. On the <span class="pagenum"><a name="page_377" id="page_377">[377]</a></span> +other hand, as we have seen in the preceding chapter, there is a vast +accumulation of evidence in favour of the theory. Hence, it is no wonder +that the theory has now been accepted by all naturalists, with scarcely +any one notable exception, as at any rate the best working hypothesis +which has ever been propounded whereby to explain the facts of organic +evolution. Moreover, in the opinion of those most competent to judge, +the theory is entitled to be regarded as something very much more than a +working hypothesis: it is held to be virtually a completed induction, +or, in other words, the proved exhibition of a general law, whereby the +causation of organic evolution admits of being in large part—if +not altogether—explained.</p> + +<p>Now, whether or not we subscribe to this latter conclusion ought, I +think, to depend upon what we mean by an explanation in the case which +is before us. If we mean only that, given the large class of known facts +and unknown causes which are conveniently summarized under the terms +Heredity and Variability, then the further facts of Struggle and +Survival serve, in some considerable degree or another, to account for +the phenomena of adaptive evolution, I cannot see any room to question +that the evidence is sufficient to prove the statement. But it is clear +that by taking for granted these great facts of Heredity and +Variability, we have assumed the larger part of the problem as a whole. +Or, more correctly, by thus generalizing, in a merely verbal form, all +the unknown causes which are concerned in these two great factors of the +process in question, we are not so much as attempting to explain the +precedent <span class="pagenum"><a name="page_378" id="page_378">[378]</a></span> +causation which serves as a condition to the process. Much more than +half the battle would already have been won, had Darwin’s +predecessors been able to explain the causes of Heredity and Variation; +hence it is but a very partial victory which we have hitherto gained in +our recent discovery of the effects of Struggle and Survival.</p> + +<p>Yet partial though it be in relation to the whole battle, in itself, or +considered absolutely, there can be no reasonable doubt that it +constitutes the greatest single victory which has ever been gained by +the science of Biology. For this very reason, however, it behoves us to +consider all the more carefully the extent to which it goes. But my +discussion of this matter must be relegated to the next volume, where I +hope to give abundant proof of the soundness of Darwin’s judgment +as conveyed in the words:—"I am convinced that natural selection has +been the main, but not the exclusive, means of modification.”</p> + +<hr /><p class="pagenum"><a name="page_379" id="page_379">[379]</a></p> +<h2><a name="CHAPTER_X" id="CHAPTER_X"></a>CHAPTER X.<br /><br /> +<span class="sc">The Theory of Sexual Selection, and Concluding Remarks.</span></h2> + +<p>Although the explanatory value of the Darwinian theory of natural +selection is, as we have now seen, incalculably great, it nevertheless +does not meet those phenomena of organic nature which perhaps more than +any other attract the general attention, as well as the general +admiration, of mankind: I mean all that class of phenomena which go to +constitute the Beautiful. Whatever value beauty as such may have, it +clearly has not a life-preserving value. The gorgeous plumage of a +peacock, for instance, is of no advantage to the peacock in his struggle +for life, and therefore cannot be attributed to the agency of natural +selection. Now this fact of beauty in organic structures is a fact of +wide generality—almost as wide, indeed, as is the fact of their +utility. Mr. Darwin, therefore, suggested another hypothesis whereby to +render a scientific explanation of this fact. Just as by his theory of +natural selection he sought to explain the major fact of utility, so did +he endeavour to explain the minor fact of beauty by a theory of what he +termed Sexual Selection.</p><p><span class="pagenum"><a name="page_380" id="page_380">[380]</a></span></p> + +<p>It is a matter of observation that the higher animals do not pair +indiscriminately; but that the members of either sex prefer those +individuals of the opposite sex which are to them most attractive. It is +important to understand <i>in limine</i> that nobody has ever attempted to +challenge this statement. In other words, it is an unquestionable fact +that among many of the higher animals there literally and habitually +occurs a <i>sexual selection</i>; and this fact is not a matter of inference, +but, as I have said, a matter of observation. The inference only begins +where, from this observable fact, it is argued,—1st, that the sexual +selection has reference to an æsthetic taste on the part of the animals +themselves; and 2nd, that, supposing the selection to be determined by +such a taste, the cause thus given is adequate to explain the phenomena +of beauty which are presented by these animals. I will consider these +two points separately.</p> + +<p>From the evidence which Darwin has collected, it appears to me +impossible to doubt that an æsthetic sense is displayed by many birds, +and not a few mammals. This of course does not necessarily imply that +the <i>standards</i> of such a sense are the same as our own; nor does it +necessarily imply that there is any constant relation between such a +sense and high levels of intelligence in other respects. In point of +fact, such is certainly not the case, because the best evidence that we +have of an æsthetic sense in animals is derived from birds, and not from +mammals. The most cogent cases to quote in this connexion are those of +the numerous species of birds which habitually adorn their nests with +gaily coloured feathers, wool, cotton, or any other gaudy materials +which they <span class="pagenum"><a name="page_381" id="page_381">[381]</a></span> +may find lying about the woods and fields. In many cases a marked +preference is shown for particular objects—as, for instance, in +the case of the Syrian nut-hatch, which chooses the iridescent wings of +insects, or that of the great crested fly-catcher, which similarly +chooses the cast-off skins of snakes. But no doubt the most remarkable +of these cases is that of the baya-bird of Asia, which after having +completed its bottle-shaped and chambered nest<a name="FNanchor_47_47" id="FNanchor_47_47"></a><a href="#Footnote_47_47" class="fnanchor">[47]</a>, +studs it over with small lumps of clay, both inside and out, upon which +the cock-bird sticks fire-flies, apparently for the sole purpose of +securing a brilliantly decorative effect. Other birds, such as the +hammer-head of Africa, adorn the surroundings of their nests (which are +built upon the ground) with shells, bones, pieces of broken glass and +earthenware, or any objects of a bright and conspicuous character which +they may happen to find. The most consummate artists in this respect +are, however, the bower-birds; for the species of this family construct +elaborate play-houses in the form of arched tunnels, built of twigs upon +the ground. Through and around such a tunnel they chase one another; and +it is always observable that not only is the floor paved with a great +collection of shells, bones, coloured stones, and any other brilliant +objects which they are able to carry in their beaks, but also that the +walls are decorated with the most gaudy articles which the birds can +find. There is one genus, in Papua, which even goes so far as to provide +the theatre with a surrounding garden. A level piece of <span class="pagenum"><a name="page_382" id="page_382">[382]</a></span> +ground is selected as a site for the building. The latter is about two +feet high, and constructed round the growing stalk of a shrub, which +therefore serves as a central pillar to which the frame-work of the roof +is attached. Twigs are woven into this frame-work until the whole is +rendered rain-proof. The tent thus erected is about nine feet in +circumference at its base, and presents a large arch as an entrance. The +central pillar is banked up with moss at its base, and a gallery is +built round the interior of the edifice. This gallery is decorated with +flowers, fruits, fungi, &c. These are also spread over the garden, +which covers <span class="pagenum"><a name="page_383" id="page_383">[383]</a></span> +about the same area as the play-house. The flowers are said to be +removed when they fade, while fresh ones are gathered to supply their +places. Thus the garden is always kept bright with flowers, as well as +with the brilliant green of mosses, which are collected and distributed +in patches, resembling tiny lawns.</p> + +<p class="pagenum"><a name="page_382i" id="page_382i">[382i.]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_401_121.jpg" width="500" height="401" alt="The Garden Bower-bird (Amblyornis inornata)." title="" /> +<span class="caption"><span class="sc">Fig.</span> 121.—The Garden Bower-bird (<span class='sn'>Amblyornis inornata</span>). +Reduced from <i>Gould’s Birds of New Guinea</i> to ¼ nat. size.</span></div> + +<p><span class="pagenum"><a name="page_383c" id="page_383c">[383c.]</a></span> +Now these sundry cases alone seem to prove a high degree of the æsthetic +sense as occurring among birds; for, it is needless to say, none of the +facts just mentioned can be due to natural selection, seeing that they +have no reference to utility, or the preservation of life. But if an +æsthetic sense occurs in birds, we should expect, on <i>a priori</i> grounds, +that it would probably be exercised with reference to the personal +appearance of the sexes. And this expectation is fully realized. For it +is an observable fact that in most species of birds where the males are +remarkable for the brilliancy of their plumage, not only is this +brilliancy most remarkable during the pairing season, but at this season +also the male birds take elaborate pains to display their charms before +the females. Then it is that the peacock erects his tail to strut round +and round the hens, taking care always to present to them a front view, +where the coloration is most gorgeous. And the same is true of all other +gaily coloured male birds. During the pairing season they actively +compete with one another in exhibiting their attractiveness to the +females; and in many cases there are added all sorts of extraordinary +antics in the way of dancings and crowings. Again, in the case of all +song-birds, the object of the singing is to please the females; and for +this purpose the <span class="pagenum"><a name="page_384" id="page_384">[384]</a></span> +males rival one another to the best of their musical ability.</p> + +<p>Thus there can be no question that the courtship of birds is a highly +elaborate business, in which the males do their best to surpass one +another in charming the females. Obviously the inference is that the +males do not take all this trouble for nothing; but that the females +give their consent to pair with the males whose personal appearance, or +whose voice, proves to be the most attractive. But, if so, the young of +the male bird who is thus <i>selected</i> will inherit his superior beauty; +and thus, in successive generations, a continuous advance will be made +in the beauty of plumage or of song, as the case may be,—both the +origin and development of beauty in the animal world being thus supposed +due to the æsthetic taste of animals themselves.</p> + +<p>Such is the theory of sexual selection in its main outlines; and with +regard to it we must begin by noting two things which are of most +importance. In the first place, it is a theory wholly and completely +distinct from the theory of natural selection; so that any truth or +error in the one does not in the least affect the other. The second +point is, that there is not so great a wealth of evidence in favour of +sexual selection as there is in favour of natural selection; and, +therefore, that while all naturalists nowadays accept natural selection +as <i>a</i> (whether or not <i>the</i>) cause of adaptive, useful, or +life-preserving structures, there is no such universal—but only a very +general—agreement with reference to sexual selection as a cause of +decorative, beautiful, or life-embellishing structures. Nevertheless, +the evidence in favour of sexual <span class="pagenum"><a name="page_385" id="page_385">[385]</a></span> +selection is both large in amount and massive in weight.</p> + +<p>Our consideration of this evidence will bring us to the second division +of our subject, as previously marked out for discussion—namely, +granting that an æsthetic sense occurs in certain large divisions of the +animal kingdom, what is the proof that such a sense is a cause of the +beauty which is presented by the animals in question?</p> + +<p>Before proceeding to state this proof, however, it is desirable to +observe that under the theory of sexual selection Darwin has included +two essentially different classes of facts. For besides the large class +of facts to which I have thus far been alluding,—i. e. the cases where +two sexes of the same species differ from one another in respect of +ornamentation,—there is another class of facts equally important, +namely, the cases where the two sexes of the same species differ from +one another in respect of size, strength, and the possession of natural +weapons, such as spurs, horns, &c. In most of these cases it is the +males which are thus superiorly endowed; and it is a matter of +observation that in all cases where they are so endowed they use their +superior strength and natural weapons for fighting together, in order to +secure possession of the females. Hence results what Mr. Darwin has +called the Law of Battle between males of the same species; and this law +of battle he includes under his theory of sexual selection. But it is +evident that the principle which is operative in the law of battle +differs from the principle which is concerned in the form of sexual +selection that has to do with embellishment, and consequent charm. The +law of <span class="pagenum"><a name="page_386" id="page_386">[386]</a></span> +battle, in fact, more nearly approaches the law of natural selection; +seeing that it expresses the natural advantages of brute force in the +struggling of rival animals, and so frequently results in <i>death of the +less fitted</i>, as distinguished from a mere failure to propagate. Now +against this doctrine of the law of battle, and the consequences to +which it leads in the superior fighting powers of male animals, no +objection has been raised in any quarter. It is only with regard to the +other aspect of the theory of sexual selection—or that which is +concerned with the superior embellishment of male animals—that any +difference of opinion obtains. I will now proceed to give the main +arguments on both sides of this question, beginning with a +<i>résumé</i> of the evidences in favour of sexual selection.</p> + +<p>In the first place, the fact that secondary sexual characters of the +embellishing kind are so generally restricted to the male sex in itself +seems to constitute very cogent proof that, in some way or another, such +characters are connected with the part which is played by the male in +the act of propagation. Moreover, secondary sexual characters of this +kind are of quite as general occurrence as are those of the other kind +which have to do with rivalry in battle; and the former are usually of +the more elaborate description. Therefore, as there is no doubt that +secondary sexual characters of the one order have an immediate purpose +to serve in the act of propagation, we are by this close analogy +confirmed in our surmise that secondary sexual characters of the other, +and still more elaborate, order are likewise so concerned. Moreover, +this view of their meaning becomes still further strengthened when we +<span class="pagenum"><a name="page_387" id="page_387">[387]</a></span> +take into consideration the following facts. Namely, (<i>a</i>) secondary +sexual characters of the embellishing kind are, as a rule, developed +only at maturity; and most frequently during only a part of the year, +which is invariably the breeding season: (<i>b</i>) they are always more or +less seriously affected by emasculation: (<i>c</i>) they are always, and +only, displayed in perfection during the act of courtship: (<i>d</i>) then, +however, they are displayed with the most elaborate pains; yet always, +and only, before the females: (<i>e</i>) they appear, at all events in many +cases, to have the effect of charming the females into a performance of +the sexual act; while it is certain that in many cases, both among +quadrupeds and birds, individuals of the one sex are capable of feeling +a strong antipathy against, or a strong preference for, certain +individuals of the opposite sex.</p> + +<p>Such are the main lines of evidence in favour of the theory of sexual +selection. And although it is enough that some of them should be merely +stated as above in order that their immense significance should become +apparent, in the case of others a bare statement is not sufficient for +this purpose. More especially is this the case as regards the enormous +profusion, variety, and elaboration of sexually-embellishing characters +which occur in birds and mammals—not to mention several divisions of +Arthropoda; together with the extraordinary amount of trouble which, in +a no less extraordinary number of different ways, is taken by the male +animals to display their embellishments before the females. And even in +many cases where to our eyes there is no particular embellishment to +display, the process of courtship consists in such an elaborate +performance of dancings, struttings, and <span class="pagenum"><a name="page_388" id="page_388">[388]</a></span> +attitudinizings that it is scarcely possible to doubt their object is +to incite the opposite sex. Here, for instance, is a series of drawings +illustrating the courtship of spiders. I choose this case as an example, +partly <span class="pagenum"><a name="page_390" id="page_390">[390]</a></span>because it is the one which has been published most +recently, and partly because it is of particular interest as occurring +so low down in the zoological scale. I am indebted to the kindness of +Mr. and Mrs. Peckham for permission to reproduce these few selected +drawings from their very admirable work, which is published by the +Natural History Society of Wisconsin, U.S. It is evident at a glance +that all these elaborate, and to our eyes ludicrous, performances are +more suggestive of incitation than of any other imaginable purpose. And +this view of the matter is strongly corroborated by the fact that it is +the most brightly coloured parts of the male spiders which are most +obtruded upon the notice of the female by these peculiar +attitudes—in just the same way as is invariably the case in the +analogous phenomena of courtship among birds, insects, &c.</p> + +<p class="pagenum"><a name="page_388i" id="page_388i">[388i.]</a></p> +<div class="figcenter" style="width: 329px;"> +<img src="images/i_407_122.jpg" width="329" height="500" alt="Courtship of Spiders." title="" /> +<div class="captionh"><span class="sc">Fig.</span> 122.—Courtship of Spiders. A few examples of some +of the attitudes adopted by different species of males when approaching +their females. (After Peckham.)</div></div> + +<p class="pagenum"><a name="page_389" id="page_389">[389]</a></p> +<div class="figcenter" style="width: 335px;"> +<img src="images/i_408_123.jpg" width="335" height="500" alt="Courtship of Spiders (Continued)." title="" /> +<div class="captionh"><span class="sc">Fig.</span> 123.—Courtship of Spiders. Continued from Fig. 122, +similarly showing some of the attitudes of approach adopted by males of +yet other different species. (After Peckham.)</div></div> + +<p><span class="pagenum"><a name="page_390c" id="page_390c">[390c.]</a></span> +But so great is the mass of material which Darwin has collected in proof +of all the points mentioned in the foregoing paragraph, that to attempt +anything in the way of an epitome would really be to damage its +evidential force. Therefore I deem it best simply to refer to it as it +stands in his <i>Descent of Man</i>, concluding, as he concludes,—"This +surprising uniformity in the laws regulating the differences between the +sexes in so many and such widely separated classes is intelligible if we +admit the action throughout all the higher divisions of the animal +kingdom of one common cause, namely, sexual selection"; while, as he +might well have added, it is difficult to imagine that all the large +classes of facts which an admission of this common cause serves to +explain, can ever admit of being rendered intelligible by any other +theory.</p> + +<p><span class="pagenum"><a name="page_391" id="page_391">[391]</a></span> +We may next proceed to consider the objections which have been brought +against the theory of sexual selection. And this is virtually the same +thing as saying that we may now consider Mr. Wallace’s views upon +the subject.</p> + +<p>Reserving for subsequent consideration the most general of these +objections—namely, that at best the theory can only apply to the more +intelligent animals, and so must necessarily fail to explain the +phenomena of beauty in the less intelligent, or in the non-intelligent, +as well as in all species of plants—we may take <i>seriatim</i> the other +objections which, in the opinion of Mr. Wallace, are sufficient to +dispose of the theory even as regards the higher animals.</p> + +<p>In the first place, he argues that the principal cause of the greater +brilliancy of male animals in general, and of male birds in particular, +is that they do not so much stand in need of protection arising from +concealment as is the case with their respective females. Consequently +natural selection is not so active in repressing brilliancy of colour in +the males, or, which amounts to the same thing, is more active in +“repressing in the female those bright colours which are normally +produced in both sexes by general laws.”</p> + +<p>Next, he argues that not only does natural selection thus exercise a +negative influence in passively permitting more heightened colour to +appear in the males, but even exercises a positive influence in actively +promoting its development in the males, while, at the same time, +actively repressing its appearance in the females. For heightened +colour, he says, is correlated with health and vigour; and as there +<span class="pagenum"><a name="page_392" id="page_392">[392]</a></span> +can be no doubt that healthy and vigorous birds best provide for their +young, natural selection, by always placing its premium on health and +vigour in the males, thus also incidentally promotes, through correlated +growth, their superior coloration.</p> + +<p>Again, with regard to the display which is practised by male birds, and +which constitutes the strongest of all Mr. Darwin’s arguments in +favour of sexual selection, Mr. Wallace points out that there is no +evidence of the females being in any way affected thereby. On the other +hand, he argues that this display may be due merely to general +excitement; and he lays stress upon the more special fact that moveable +feathers are habitually erected under the influence of anger and +rivalry, in order to make the bird look more formidable in the eyes of +antagonists.</p> + +<p>Furthermore, he adduces the consideration that, even if the females are +in any way affected by colour and its display on the part of the males, +and if, therefore, sexual selection be conceded a true principle in +theory, still we must remember that, as a matter of fact, it can only +operate in so far as it is allowed to operate by natural selection. Now, +according to Mr. Wallace, natural selection must wholly neutralize any +such supposed influence of sexual selection. For, unless the survivors +in the general struggle for existence happen to be those which are also +the most highly ornamented, natural selection must neutralize and +destroy any influence that may be exerted by female selection. But +obviously the chances against the otherwise best fitted males happening +to be likewise the most highly ornamented must be many to <span class="pagenum"><a name="page_393" id="page_393">[393]</a></span> +one, unless, as Wallace supposes, there is some correlation between +embellishment and general perfection, in which case, as he points out, +the theory of sexual selection lapses altogether, and becomes but a +special case of natural selection.</p> + +<p>Once more, Mr. Wallace argues that the evidence collected by Mr. Darwin +himself proves that each bird finds a mate under any circumstances—a +general fact which in itself must quite neutralize any effect of sexual +selection of colour or ornament, since the less highly coloured birds +would be at no disadvantage as regards the leaving of healthy progeny.</p> + +<p>Lastly, he urges the high improbability that through thousands of +generations all the females of any particular species—possibly spread +over an enormous area—should uniformly and always have displayed +exactly the same taste with respect to every detail of colour to be +presented by the males.</p> + +<p>Now, without any question, we have here a most powerful array of +objections against the theory of sexual selection. Each of them is ably +developed by Mr. Wallace himself in his work on <i>Tropical Nature</i>; and +although I have here space only to state them in the most abbreviated of +possible forms, I think it will be apparent how formidable these +objections appear. Unfortunately the work in which they are mainly +presented was published several years after the second edition of the +<i>Descent of Man</i>, so that Mr. Darwin never had a suitable opportunity of +replying. But, if he had had such an opportunity, as far as I can judge +it seems that his reply would have been more or less as follows.</p> + +<p>In the first place, Mr. Wallace fails to distinguish <span class="pagenum"><a name="page_394" id="page_394">[394]</a></span> +between brilliancy and ornamentation—or between colour as merely +“heightened,” and as distinctively decorative. Yet there is +obviously the greatest possible difference between these two things. We +may readily enough admit that a mere heightening of already existing +coloration is likely enough—at all events in many cases—to +accompany a general increase of vigour, and therefore that natural +selection, by promoting the latter, may also incidentally promote the +former, in cases where brilliancy is not a source of danger. But clearly +this is a widely different thing from showing that not only a <i>general +brilliancy of colour</i>, but also <i>the particular disposition of colours</i>, +in the form of ornamental patterns, can thus be accounted for by natural +selection. Indeed, it is expressly in order to account for the +occurrence of such ornamental patterns that Mr. Darwin constructed his +theory of sexual selection; and therefore, by thus virtually ignoring +the only facts which that theory endeavours to explain, Mr. Wallace is +not really criticizing the theory at all. By representing that the +theory has to do only with brilliancy of colour, as distinguished from +disposition of colours, he is going off upon a false issue which has +never really been raised<a name="FNanchor_48_48" id="FNanchor_48_48"></a><a href="#Footnote_48_48" class="fnanchor">[48]</a>. +Look, for example, at a peacock’s tail. No doubt it is +sufficiently brilliant; but far more remarkable than its brilliancy is +its elaborate pattern on the one hand, and its enormous size on the +other. There is no conceivable reason why mere <i>brilliancy of colour</i>, +as an accidental concomitant of general vigour, should have run into so +extraordinary, so elaborate, and so beautiful <span class="pagenum"><a name="page_395" id="page_395">[395]</a></span> +a <i>design of colours</i>. Moreover, this design is only unfolded when the +tail is erected, and the tail is not erected in battle (as Mr. +Wallace’s theory of the erectile function in feathers would +require), but in courtship; obviously, therefore, the purpose of the +pattern, so to speak, is correlated with the act of courtship—it +being only then, in fact, that the general purpose of the whole +structure, as well as the more special purpose of the pattern, becomes +revealed. Lastly, the fact of this whole structure being so large, +entailing not only a great amount of physiological material in its +production, but also of physiological energy in carrying about such a +weight, as well as of increased danger from impeding locomotion and +inviting capture—all this is obviously incompatible with the +supposition of the peacock’s tail having been produced by natural +selection. And such a case does not stand alone. There are multitudes of +other instances of ornamental structures imposing a drain upon the vital +energies of their possessors, without conferring any compensating +benefit from a utilitarian point of view. Now, in all these cases, +without any exception, such structures are ornamental structures which +present a plain and obvious reference to the relationship of the sexes. +Therefore it becomes almost impossible to doubt—first, that they +exist for the sake of ornament; and next, that the ornament exists on +account of that relationship. If such structures were due merely to a +superabundance of energy, as Mr. Wallace supposes, not only ought they +to have been kept down by the economizing influence of natural +selection; but we can see no reason, either why they should be so highly +ornamental on the one hand, or <span class="pagenum"><a name="page_396" id="page_396">[396]</a></span> +so exclusively related to the sexual relationship on the other.</p> + +<div class="figcenter" style="width: 500px;"> +<img src="images/i_415_124.jpg" width="500" height="405" alt="The Bell-bird." title="" /> +<div class="captionh"><span class="sc">Fig.</span> 124.—The Bell-bird (<span class='sn'>Chasmorhynchus niveus</span>, ¼ +natural size). Drawn from nature (<i>R. Coll. Surg. Mus.</i>). In the drawing +of the adult male the ornamental appendage is represented in its +inflated condition, during courtship; in the drawing of the young male +it is shown in its flaccid condition.</div></div> + +<p>Finally, we must take notice of the fact that where peculiar +<i>structures</i> are concerned for purposes of display in courtship, the +<i>elaboration</i> of these structures is often no less remarkable than that +of patterns where colours are thus concerned. Take, for example, the +case of the Bell-bird, which I select from an innumerable number of +instances that might be mentioned because, while giving a verbal +description of this animal, Darwin does not supply a pictorial +representation <span class="pagenum"><a name="page_397" id="page_397">[397]</a></span> +thereof. The bird, which lives in South America, has a very loud and +peculiar call, that can be heard at a distance of two or three miles. +The female is dusky-green; but the adult male is a beautiful white, +excepting the extraordinary structure with which we are at present +concerned. This is a tube about three inches long, which rises from the +base of the beak. It is jet black, and dotted over with small downy +feathers. The tube is closed at the top, but its cavity communicates +with the palate, and thus the whole admits of being inflated from +within, when, of course, it stands erect as represented in one of the +two drawings.<span class="pagenum"><a name="page_398" id="page_398">[398]</a></span> +When not thus inflated, it hangs down, as shown in the second figure, +which represents the plumage of a young male. (Fig. 124.)</p> + +<p>In another species of the genus there are three of these appendages—the +two additional ones being mounted on the corners of the mouth. (Fig. +125.) In all species of the genus (four in number) the tubes are +inflated during courtship, and therefore perform the function of sexual +embellishments. Now the point to which I wish to draw attention is, that +so specialized and morphologically elaborate a structure cannot be +regarded as merely adventitious. It must have been developed by some +definite cause, acting through a long series of generations. And as no +other function can be assigned to it than that of charming the female +when it is erected in courtship, the peculiarity of form and mechanism +which it presents—like the elaboration of patterns in cases where +colour only is concerned—virtually compels us to recognise in sexual +selection the only conceivable cause of its production.</p> + +<p class="pagenum"><a name="page_397i" id="page_397i">[397i.]</a></p> +<div class="figcenter" style="width: 500px;"> +<img src="images/i_416_125.jpg" width="500" height="417" alt="C. tricarunculatus." title="" /> +<div class="captionh"><span class="sc">Fig.</span> 125.—<span class='sn'>C. tricarunculatus</span>, ¼ natural size. Copied +from the <i>Ibis</i>. The ornamental appendages of the male are represented +in a partly inflated condition.</div></div> + +<p><span class="pagenum"><a name="page_398c" id="page_398c">[398c.]</a></span> +For these reasons I think that Mr. Wallace’s main objection falls +to the ground. Passing on to his subsidiary objections, I do not see +much weight in his merely negative difficulty as to there being an +absence of evidence upon hen birds being charmed by the plumage, or the +voice, of their consorts. For, on the one hand, it is not very safe to +infer what sentiments may be in the mind of a hen; and, on the other +hand, it is impossible to conceive what motive can be in the mind of a +cock, other than that of making himself attractive, when he performs his +various antics, displays his ornamental plumes, or sings his melodious +songs. Considerations somewhat analogous apply to the <span class="pagenum"><a name="page_399" id="page_399">[399]</a></span> +difficulty of supposing so much similarity and constancy of taste on +the part of female animals as Mr. Darwin’s theory undoubtedly +requires. Although we know very little about the psychology of the lower +animals, we do observe in many cases that small details of mental +organization are often wonderfully constant and uniform throughout all +members of a species, even where it is impossible to suggest any utility +as a cause.</p> + +<p>Again, as regards the objection that each bird finds a mate under any +circumstances, we have here an obvious begging of the whole question. +That every feathered Jack should find a feathered Jill is perhaps what +we might have antecedently expected; but when we meet with innumerable +instances of ornamental plumes, melodious songs, and the rest, as so +many witnesses to a process of sexual selection having always been in +operation, it becomes irrational to exclude such evidence on account of +our antecedent prepossessions.</p> + +<p>There remains the objection that the principles of natural selection +must necessarily swallow up those of sexual selection. And this +consideration, I doubt not, lies at the root of all Mr. Wallace’s +opposition to the supplementary theory of sexual selection. He is +self-consistent in refusing to entertain the evidence of sexual +selection, on the ground of his antecedent persuasion that in the great +drama of evolution there is no possible standing-ground for any other +actor than that which appears in the person of natural selection. But +here, again, we must refuse to allow any merely antecedent presumption +to blind our eyes to the actual evidence of other agencies having +co-operated <span class="pagenum"><a name="page_400" id="page_400">[400]</a></span> +with natural selection in producing the observed results. And, as +regards the particular case now before us, I think I have shown, as far +as space will permit, that in the phenomena of decorative colouring (as +distinguished from merely brilliant colouring), of melodious song (as +distinguished from merely tuneless cries), of enormous arborescent +antlers (as distinguished from merely offensive weapons), and so +forth—I say that in all these phenomena we have phenomena which +cannot possibly be explained by the theory of natural selection; and, +further, that if they are to be explained at all, this can only be done, +so far as we can at present see, by Mr. Darwin’s supplementary +theory of sexual selection.</p> + +<p>I have now briefly answered all Mr. Wallace’s objections to this +supplementary theory, and, as previously remarked, I feel pretty +confident that, at all events in the main, the answer is such as Mr. +Darwin would himself have supplied, had there been a third edition of +his work upon the subject. At all events, be this as it may, we are +happily in possession of unquestionable evidence that he believed all +Mr. Wallace’s objections to admit of fully satisfactory answers. +For his very last words to science—read only a few hours before his +death at a meeting of the Zoological Society—were:</p> + +<div class='blockquot'><p>I may perhaps be here permitted to say that, after having carefully +weighed, to the best of my ability, the various arguments which +have been advanced against the principle of sexual selection, I +remain firmly convinced of its truth<a name="FNanchor_49_49" id="FNanchor_49_49"></a><a href="#Footnote_49_49" class="fnanchor">[49]</a>.</p></div> + +<p class="pagenum"><a name="page_401" id="page_401">[401]</a></p> +<h3 class='i'>Concluding Remarks.</h3> + +<p>I will now conclude this chapter, and with it the present volume, by +offering a few general remarks on what may be termed the philosophical +relations of Darwinian doctrine to the facts of adaptation on the one +hand, and to those of beauty on the other. Of course we are all aware +that before the days of this doctrine the facts of adaptation in organic +nature were taken to constitute the clearest possible evidence of +special design, on account of the wonderful mechanisms which they +everywhere displayed; while the facts of beauty were taken as +constituting no less conclusive evidence of the quality of such special +design as beneficent, not to say artistic. But now that the Darwinian +doctrine appears to have explained scientifically the former class of +facts by its theory of natural selection, and the latter class of facts +by its theory of sexual selection, we may fitly conclude this brief +exposition of the doctrine as a whole by considering what influence such +naturalistic explanations may fairly be taken to exercise upon the +older, or super-naturalistic, interpretations.</p> + +<p>To begin with the facts of adaptation, we must first of all observe that +the Darwinian doctrine is immediately concerned with these facts only in +so far <span class="pagenum"><a name="page_402" id="page_402">[402]</a></span> +as they occur in organic nature. With the adaptations—if they can +properly be so called—which occur in all the rest of nature, and +which go to constitute the Cosmos as a whole so wondrous a spectacle of +universal law and perfect order, this doctrine is but indirectly +concerned. Nevertheless, it is of course fundamentally concerned with +them to the extent that it seeks to bring the phenomena of organic +nature into line with those of inorganic; and therefore to show that +whatever view we may severally take as to the kind of causation which is +energizing in the latter we must now extend to the former. This is +usually expressed by saying that the theory of evolution by natural +selection is a mechanical theory. It endeavours to comprise all the +facts of adaptation in organic nature under the same category of +explanation as those which occur in inorganic nature—that is to +say, under the category of physical, or ascertainable, causation. +Indeed, unless the theory has succeeded in doing this, it has not +succeeded in doing anything—beyond making a great noise in the +world. If Mr. Darwin has not discovered a new mechanical cause in the +selection principle, his labour has been worse than in vain.</p> + +<p>Now, without unduly repeating what has already been said in Chapter +VIII, I may remark that, whatever we may each think of the measure of +success which has thus far attended the theory of natural selection in +explaining the facts of adaptation, we ought all to agree that, +considered as a matter of general reasoning, the theory does certainly +refer to a <i>vera causa</i> of a strictly physical kind; and, therefore, +that no exception can be taken to the theory in this respect <span class="pagenum"><a name="page_403" id="page_403">[403]</a></span> +on grounds of <i>logic</i>. If the theory in this respect is to be attacked +at all, it can only be on grounds of <i>fact</i>—namely, by arguing +that the cause does not occur in nature, or that, if it does, its +importance has been exaggerated by the theory. Even, however, if the +latter proposition should ever be proved, we may now be virtually +certain that the only result would be the relegation of all the residual +phenomena of adaptation to other causes of the physical +order—whether known or unknown. Hence, as far as the matter of +<i>principle</i> is concerned, we may definitely conclude that the great +naturalistic movement of our century has already brought all the +phenomena of adaptation in organic nature under precisely the same +category of mechanical causation, as similar movements in previous +centuries have brought all the known phenomena of inorganic nature: the +only question that remains for solution is the strictly <i>scientific</i> +question touching the particular causes of the mechanical order which +have been at work.</p> + +<p>So much, then, for the phenomena of adaptation. Turning next to those of +beauty, we have already seen that the theory of sexual selection stands +to these in precisely the same relation as the theory of natural +selection does to those of adaptation. In other words, it supplies a +physical explanation of them; because, as far as our present purposes +are concerned, it may be taken for granted, or for the sake of argument, +that inasmuch as psychological elements enter into the question the +cerebral basis which they demand involves a physical side.</p> + +<p>There is, moreover, this further point of resemblance between the two +theories: neither of them has any <span class="pagenum"><a name="page_404" id="page_404">[404]</a></span> +reference to inorganic nature. Therefore, with the charm or the +loveliness of landscapes, of earth and sea and sky, of pebbles, +crystals, and so forth, we have at present nothing to do. How it is that +so many inanimate objects are invested with beauty—why it is that +beauty attaches to architecture, music, poetry, and many other +things—these are questions which do not specially concern the +biologist. If they are ever to receive any satisfactory explanation in +terms of natural causation, this must be furnished at the hands of the +psychologist. It may be possible for him to show, more satisfactorily +than hitherto, that all beauty, whenever and wherever it occurs, is +literally “in the eyes of the beholder"; or that objectively +considered, there is no such thing as beauty. It may be—and in my +opinion it probably is—purely an affair of the percipient mind +itself, depending on the association of ideas with pleasure-giving +objects. This association may well lead to a liking for such objects, +and so to the formation of what is known as æsthetic feeling with +regard to them. Moreover, beauty of inanimate nature must be an affair +of the percipient mind itself, unless there be a creating intelligence +with organs of sense and ideals of beauty similar to our own. And, apart +from any deeper considerations, this latter possibility is scarcely +entitled to be regarded as a probability, looking to the immense +diversities in those ideals among different races of mankind. But, be +this as it may, the scientific problem which is presented by the fact of +æsthetic feeling, even if it is ever to be satisfactorily solved, +is a problem which, as already remarked, must be dealt with by +psychologists. As biologists we have simply to accept this feeling as a +fact, and to consider <span class="pagenum"><a name="page_405" id="page_405">[405]</a></span> +how, out of such a feeling as a cause, the beauty of organic nature may +have followed as an effect.</p> + +<p>Now we have already seen how the theory of sexual selection supposes +this to have happened. But against this theory a formidable objection +arises, and one which I have thought it best to reserve for treatment in +this place, because it serves to show the principal difference between +Mr. Darwin’s two great generalizations, considered as +generalizations in the way of mechanical theory. For while the theory of +natural selection extends equally throughout the whole range of organic +nature, the theory of sexual selection has but a comparatively +restricted scope, which, moreover, is but vaguely defined. For it is +obvious that the theory can only apply to living organisms which are +sufficiently intelligent to admit of our reasonably accrediting them +with æsthetic taste—namely, in effect, the higher animals. And just as +this consideration greatly restricts the possible scope of the theory, +as compared with that of natural selection, so does it render undefined +the zoological limits within which it can be reasonably employed. +Lastly, this necessarily undefined, and yet most important limitation +exposes the theory to the objection just alluded to, and which I shall +now mention.</p> + +<p>The theory, as we have just seen, is necessarily restricted in its +application to the higher animals. Yet the facts which it is designed to +explain are not thus restricted. For beauty is by no means restricted to +the higher animals. The whole of the vegetable world, and the whole of +the animal world at least as high up in the scale as the insects, must +be taken as incapable of æsthetic feeling. Therefore, the extreme +<span class="pagenum"><a name="page_406" id="page_406">[406]</a></span> +beauty of flowers, sea-anemones, corals, and so forth, cannot possibly +be ascribed to sexual selection.</p> + +<p>Now, with regard to this difficulty, we must begin by excluding the case +of the vegetable kingdom as irrelevant. For it has been rendered highly +probable—if not actually proved—by Darwin and others, that the beauty +of flowers and of fruits is in large part due to natural selection. It +is to the advantage of flowering plants that their organs of +fructification should be rendered conspicuous—and in many cases also +odoriferous,—in order to attract the insects on which the process of +fertilization depends. Similarly, it is to the advantage of all plants +which have brightly coloured fruits that these should be conspicuous for +the purpose of attracting birds, which eat the fruits and so disseminate +the seed. Hence all the gay colours and varied forms, both of flowers +and fruits, have been thus adequately explained as due to natural +causes, working for the welfare, as distinguished from the beauty, of +the plants. For even the distribution of colours on flowers, or the +beautiful patterns which so many of them present, are found to be useful +in guiding insects to the organs of fructification.</p> + +<p>Again, the green colouring of leaves, which lends so much beauty to the +vegetable world, has likewise been shown to be of vital importance to +the physiology of plant-life; and, therefore, may also be ascribed to +natural selection. Thus, there remains only the forms of plants other +than the flowers. But the forms of leaves have also in many cases been +shown to be governed by principles of utility; and the same is to be +said of the branching structure which is so characteristic of trees and +shrubs, since this is the <span class="pagenum"><a name="page_407" id="page_407">[407]</a></span> +form most effectual for spreading out the leaves to the light and air. +Here, then, we likewise find that the cause determining plant beauty is +natural selection; and so we may conclude that the only reason why the +forms of trees which are thus determined by utility appeal to us as +beautiful, is because we are accustomed to these the most ordinary +forms. Our ideas having been always, as it were, moulded upon these +forms, æsthetic feeling becomes attached to them by the principle +of association. At any rate, it is certain that when we contemplate +almost any forms of plant-structure which, for special reasons of +utility, differ widely from these (to us) more habitual forms, the +result is not suggestive of beauty. Many of the tropical and +un-tree-like plants—such as the cactus tribe—strike us as +odd and quaint, not as beautiful. Be this however as it may, I trust I +have said enough to prove that in the vegetable world, at all events, +the attainment of beauty cannot be held to have been an object aimed at, +so to speak, for its own sake. Even if, for the purposes of argument, we +were to suppose that all the forms and colours in the vegetable world +are due to special design, there could be no doubt that the purpose of +this design has been in chief part a utilitarian purpose; it has not +aimed at beauty exclusively for its own sake. For most of such beauty as +we here perceive is plainly due to the means adopted for the attainment +of life-preserving ends, which, of course, is a metaphorical way of +saying that it is probably due to natural selection<a +name="FNanchor_50_50" id="FNanchor_50_50"></a><a href="#Footnote_50_50" class="fnanchor">[50]</a>.</p> + +<p><span class="pagenum"><a name="page_408" id="page_408">[408]</a></span> +Turning, then, to the animal kingdom below the level of insects, here we +are bound to confess that the beauty which so often meets us cannot +reasonably be ascribed either to natural or to sexual selection. Not to +sexual selection for the reasons already given; the animals in question +are neither sufficiently intelligent to possess any æsthetic +taste, nor, as a matter of fact, do we observe that they exercise any +choice in pairing. Not to natural selection, because we cannot here, as +in the case of vegetables, point to any benefit as generally arising +from bright colours and beautiful forms. On the principles of +naturalism, therefore, we are driven to conclude that the beauty here is +purely adventitious, or accidental. Nor need we be afraid to make this +admission, if only we take a sufficiently wide view of the facts. For, +when we do take such a view, we find that beauty here is by no means of +invariable, or even of general, occurrence. There is no loveliness about +an oyster or a lob-worm; parasites, as a rule, are positively ugly, and +they constitute a good half of all animal species. The truth seems to +be, when we look attentively at the matter, that in all cases where +beauty does occur in these lower forms of animal life, its presence is +owing to one of two things—either to the radiate form, or to the +bright tints. Now, seeing that the radiate form is of such general +occurrence among these lower animals—appearing over and over +again, with the utmost insistence, even among groups widely separated +from one another by <span class="pagenum"><a name="page_409" id="page_409">[409]</a></span> +the latest results of scientific classification—seeing this, it +becomes impossible to doubt that the radiate form is due to some +morphological reasons of wide generality. Whether these reasons be +connected with the internal laws of growth, or to the external +conditions of environment, I do not pretend to suggest. But I feel safe +in saying that it cannot possibly be due to any design to secure beauty +for its own sake. The very generality of the radiate form is in itself +enough to suggest that it must have some physical, as distinguished from +an æsthetic, explanation; for, if the attainment of beauty had +here been the object, surely it might have been even more effectually +accomplished by adopting a greater variety of typical forms—as, +for instance, in the case of flowers.</p> + +<p>Coming then, lastly, to the case of brilliant tints in the lower +animals, Mr. Darwin has soundly argued that there is nothing forced or +improbable in the supposition that organic compounds, presenting as they +do such highly complex and such varied chemical constitutions, should +often present brilliant colouring <i>incidentally</i>. Considered merely as +colouring, there is nothing in the world more magnificent than arterial +blood; yet here the colouring is of purely utilitarian significance. It +is of the first importance in the chemistry of respiration; but is +surely without any meaning from an æsthetic point of view. For the +colour of the cheeks, and of the flesh generally, in the <i>white</i> races +of mankind, could have been produced quite as effectually by the use of +pigment—as in the case of certain monkeys. Now the fact that in the +case of blood, as in that of many other highly coloured fluids and +solids throughout the animal <span class="pagenum"><a name="page_410" id="page_410">[410]</a></span> +kingdom, the colour is <i>concealed</i>, is surely sufficient proof that the +colour, if regarded from an æsthetic point of view, is accidental. +Therefore, when, as in other cases, such colouring occurs upon the +surface, and thus becomes apparent, are we not irresistibly led to +conclude that its <i>exhibition</i> in such cases is likewise accidental, so +far as any question of æsthetic design is concerned?</p> + +<p>I have now briefly glanced at all the main facts of organic nature with +reference to beauty; and, as a result, I think it is impossible to +resist the general conclusion, that in organic nature beauty does not +exist as an end <i>per se</i>. All cases where beauty can be pointed to in +organic nature are seemingly due—either to natural selection, acting +without reference to beauty, but to utility; to sexual selection, acting +with reference to the taste of animals; or else to sheer accident. And +if this general conclusion should be held to need any special +verification, is it not to be found in the numberless cases where +organic nature not only fails to be beautiful, but reveals itself as the +reverse. Not again to refer to the case of parasites, what can be more +unshapely than a hippopotamus, or more generally repulsive than a +crocodile? If it be said that these are exceptions, and that the forms +of animals as a rule are graceful, the answer—even apart from +parasites—is obvious. In all cases where the habits of life are such as +to render rapid locomotion a matter of utilitarian necessity, the +outlines of an animal <i>must</i> be graceful—else, whether the locomotion +be terrestrial, aerial, or aquatic, it must fail to be swift. Hence it +is only in such cases as that of the hippopotamus, <span class="pagenum"><a name="page_411" id="page_411">[411]</a></span> +rhinoceros, elephant, crocodile, and so forth, where natural selection +has had no concern in developing speed, that the accompanying accident +of gracefulness can be allowed to disappear. But if beauty in organic +nature had been in itself what may be termed an artistic object on the +part of a divine Creator, it is absurd to suggest that his design in +this matter should only have been allowed to appear where we are able to +detect other and very good reasons for its appearance.</p> + +<hr class='minor' /> + +<p>Thus, whether we look to the facts of adaptation or to those of beauty, +everywhere throughout organic nature we meet with abundant evidence of +natural causation, while nowhere do we meet with any independent +evidence of supernatural design. But, having led up to this conclusion, +and having thus stated it as honestly as I can, I should like to finish +by further stating what, in my opinion is its logical bearing upon the +more fundamental tenets of religious thought.</p> + +<p>As I have already observed at the commencement of this brief exposition, +prior to the Darwinian theory of organic evolution, the theologian was +prone to point to the realm of organic nature as furnishing a peculiarly +rich and virtually endless store of facts, all combining in their +testimony to the wisdom and the beneficence of the Deity. Innumerable +adaptations of structures to functions appeared to yield convincing +evidence in favour of design; the beauty so profusely shed by living +forms appeared to yield evidence, no less convincing, of that design as +beneficent. But both these sources of evidence have now, as it were, +been <span class="pagenum"><a name="page_412" id="page_412">[412]</a></span> +tapped at their fountain-head: the adaptation and the beauty are alike +receiving their explanation at the hands of a purely mechanical +philosophy. Nay, even the personality of man himself is assailed; and +this not only in the features which he shares with the lower animals, +but also in his god-like attributes of reason, thought, and conscience. +All nature has thus been transformed before the view of the present +generation in a manner and to an extent that has never before been +possible: and inasmuch as the change which has taken place has taken +place in the direction of naturalism, and this to the extent of +rendering the mechanical interpretation of nature universal, it is no +wonder if the religious mind has suddenly awakened to a new and a +terrible force in the words of its traditional enemy—Where is now +thy God?</p> + +<p>This is not the place to discuss the bearings of science on +religion<a name="FNanchor_51_51" id="FNanchor_51_51"></a><a href="#Footnote_51_51" class="fnanchor">[51]</a>; +but I think it is a place where one may properly point out the limits +within which no such bearings obtain. Now, from what has just been said, +it will be apparent that I am not going to minimise the change which has +been wrought. On the contrary, I believe it is only stupidity or +affectation which can deny that the change in question is more deep and +broad than any single previous change in the whole history of human +thought. It is a fundamental, a cosmical, a world-transforming change. +Nevertheless, in my opinion, it is a change of a non-theistic, as +distinguished from an a-theistic, kind. It has rendered impossible the +appearance in literature of any future Paley, Bell, or Chalmers; but it +has <span class="pagenum"><a name="page_413" id="page_413">[413]</a></span> +done nothing in the way of negativing that belief in a Supreme Being +which it was the object of these authors to substantiate. If it has +demonstrated the futility of their proof, it has furnished nothing in +the way of disproof. It has shown, indeed, that their line of argument +was misjudged when they thus sought to separate organic nature from +inorganic as a theatre for the special or peculiar display of +supernatural design; but further than this it has not shown anything. +The change in question therefore, although greater in degree, is the +same in kind as all its predecessors: like all previous advances in +cosmological theory which have been wrought by the advance of science, +this latest and greatest advance has been that of revealing the +constitution of nature, or the method of causation, as everywhere the +same. But it is evident that this change, vast and to all appearance +final though it be, must end within the limits of natural causation +itself. The whole world of life and mind may now have been annexed to +that of matter and energy as together constituting one magnificent +dominion, which is everywhere subject to the same rule, or method of +government. But the ulterior and ultimate question touching the nature +of this government as mental or non-mental, personal or impersonal, +remains exactly where it was. Indeed, this is a question which cannot be +affected by <i>any</i> advance of science, further than science has proved +herself able to dispose of erroneous arguments based upon ignorance of +nature. For while the sphere of science is necessarily restricted to +that of natural causation which it is her office to explore, the +question touching the <i>nature of this natural causation</i> is one which as +necessarily lies without the <span class="pagenum"><a name="page_414" id="page_414">[414]</a></span> +whole sphere of such causation itself: therefore it lies beyond any +possible intrusion by science. And not only so. But if the nature of +natural causation be that of the highest order of known existence, then, +although we must evidently be incapable of conceiving what such a Mind +is, at least we seem capable of judging what in many respects it is not. +It cannot be more than one; it cannot be limited either in space or +time; it cannot be other than at least as self-consistent as its +manifestations in nature are invariable. Now, from the latter deduction +there arises a point of first-rate importance in the present connexion. +For if the so-called First Cause be intelligent, and therefore all +secondary causes but the expression of a supreme Will, in as far as such +a Will is self-consistent, the operation of all natural causes must be +uniform,—with the result that, as seen by us, this operation must +needs appear to be what we call mechanical. The more unvarying the Will, +the more unvarying must be this expression thereof; so that, if the +former be absolutely self-consistent, the latter cannot fail to be as +reasonably interpreted by the theory of mindless necessity, as by that +of ubiquitous intention. Such being, as it appears to me, the pure logic +of the matter, the proof of organic evolution amounts to nothing more +than the proof of a natural process. What mode of being is ultimately +concerned in this process—or in what it is that this process +ultimately consists—is a question upon which science is as +voiceless as speculation is vociferous.</p> + +<p>But, it may still be urged, surely the principle of natural selection +(with its terrible basis in the struggle for existence) and the +principle of sexual selection<span class="pagenum"><a name="page_415" id="page_415">[415]</a></span> +(with its consequence in denying beauty to be an end in itself) +demonstrate that, <i>if</i> there be design in nature, such design at all +events cannot be beneficent. To this, however, I should again reply +that, just as touching the major question of design itself, so as +touching this minor question of the quality of such design as +beneficent, I do not see how the matter has been much affected by a +discovery of the principles before us. For we did not need a Darwin to +tell us that the whole creation groaneth and travaileth together in +pain. The most that in this connexion Darwin can fairly be said to have +done is to have estimated in a more careful and precise manner than any +of his predecessors, the range and the severity of this travail. And if +it be true that the result of what may be called his scientific analysis +of nature in respect of suffering is to have shown the law of suffering +even more severe, more ubiquitous, and more necessary than it had ever +been shown before, we must remember at the same time how he has proved, +more rigidly than was ever proved before, that suffering is a condition +to improvement—struggle for life being the <i>raison +d’être</i> of higher life, and this not only in the physical +sphere, but also in the mental and moral.</p> + +<p>Lastly, if it be said that the <i>choice</i> of such a method, whereby +improvement is only secured at the cost of suffering, indicates a kind +of callousness on the part of an intelligent Being supposed to be +omnipotent, I confess that such does appear to me a legitimate +conclusion—subject, however, to the reservation that higher knowledge +might displace it. For, as far as matters are now actually presented to +the unbiased contemplation of a human mind, this provisional <span class="pagenum"><a name="page_416" id="page_416">[416]</a></span> +inference appears to me unavoidable—namely, that if the world of +sentient life be due to an Omnipotent Designer, the aim or motive of the +design must have been that of securing a continuous advance of animal +improvement, without any regard at all to animal suffering. For I own it +does not seem to me compatible with a fair and honest exercise of our +reason to set the sum of animal happiness over against the sum of animal +misery, and then to allege that, in so far as the former tends to +balance—or to over-balance—the latter, thus far is the moral +character of the design as a whole vindicated. Even if it could be shown +that the sum of happiness in the brute creation considerably +preponderates over that of unhappiness—which is the customary +argument of theistic apologists,—we should still remain without +evidence as to this state of matters having formed any essential part of +the design. On the other hand, we should still be in possession of +seemingly good evidence to the contrary. For it is clearly a condition +to progress by survival of the fittest, that as soon as organisms become +sentient selection must be exercised with reference to sentiency; and +this means that, if further progress is to take place, states of +sentiency <i>must</i> become so organized with reference to habitual +experience of the race, that pleasures and pains shall answer +respectively to states of agreement and disagreement with the sentient +creature’s environment. Those animals which found pleasure in what +was deleterious to life would not survive, while those which found +pleasure in what was beneficial to life would survive; and so +eventually, in every species of animal, states of sentiency as agreeable +or disagreeable must approximately correspond with what is good for +<span class="pagenum"><a name="page_417" id="page_417">[417]</a></span> +the species or bad for the species. Indeed, we may legitimately surmise +that the reason why sentiency (and, <i>a fortiori</i>, conscious volition) +has ever appeared upon the scene at all, has been because it +furnishes—through this continuously selected adjustment of states +of sentiency to states of the sentient organism—so admirable a +means of securing rapid, and often refined, adjustments by the organism +to the habitual conditions of its life<a name="FNanchor_52_52" id="FNanchor_52_52"></a><a href="#Footnote_52_52" class="fnanchor">[52]</a>. +But, if so, not only is this state of matters a <i>condition</i> to progress +in the future; it is further, and equally, a <i>consequence</i> of progress +in the past.</p> + +<p>However, be this as it may, from all that has gone before does it not +become apparent that pleasure or happiness on the one hand, and pain or +misery on the other, must be present in sentient nature? And so long as +they are both seen to be equally necessary under the process of +evolution by natural selection, we have clearly no more reason to regard +the pleasure than the pain as an object of the supposed design. Rather +must we see in both one and the same condition to progress under the +method of natural causation which is before us; and therefore I cannot +perceive that it makes much difference—so far as the argument for +beneficence is concerned—whether the pleasures of animals outweigh +their pains, or <i>vice versâ</i>.</p> + +<p>Upon the whole, then, it seems to me that such evidence as we have is +against rather than in favour of the inference, that if design be +operative in animate nature it has reference to animal enjoyment or +well-being, as distinguished from animal improvement or evolution. And +if this result should be found distasteful <span class="pagenum"><a name="page_418" id="page_418">[418]</a></span> +to the religious mind—if it be felt that there is no desire to +save the evidences of design unless they serve at the same time to +testify to the nature of that design as beneficent,—I must once +more observe that the difficulty thus presented to theism is not a +difficulty of modern creation. On the contrary, it has always +constituted the fundamental difficulty with which natural theologians +have had to contend. The external world appears, in this respect, to be +at variance with our moral sense; and when the antagonism is brought +home to the religious mind, it must ever be with a shock of terrified +surprise. It has been newly brought home to us by the generalizations of +Darwin; and therefore, as I said at the beginning, the religious thought +of our generation has been more than ever staggered by the +question—Where is now thy God? But I have endeavoured to show that +the logical standing of the case has not been materially changed; and +when this cry of Reason pierces the heart of Faith, it remains for Faith +to answer now, as she has always answered before—and answered with +that trust which is at once her beauty and her life—Verily thou +art a God that hidest thyself.</p> + +<hr /><p class="pagenum"><a name="page_419" id="page_419">[419]</a></p> +<h2><a name="APPENDIX_AND_NOTES" id="APPENDIX_AND_NOTES"></a><i>APPENDIX AND NOTES</i></h2> + +<hr class='minor' /><p class="pagenum"><a name="page_421" id="page_421">[421]</a></p> +<h2><a name="APPENDIX_TO_CHAPTER_V" id="APPENDIX_TO_CHAPTER_V"></a>APPENDIX TO CHAPTER V.</h2> + +<h3 class="sc">On Objections which have been brought against the Theory of Organic +Evolution on grounds of Palæontology.</h3> + +<p>While stating in the text, and in a necessarily general way, the +evidence which is yielded by palæontology to the theory of organic +evolution, I have been desirous of not overstating it. Therefore, in the +earlier paragraphs of the chapter, which deal with the most general +heads of such evidence, I introduced certain qualifying phrases; and I +will now give the reasons which led me to do so.</p> + +<p>Of all the five biological sciences which have been called into +evidence—viz. those of Classification, Morphology, Embryology, +Palæontology, and Geographical Distribution—it is in the case of +palæontology alone that any important or professional opinions still +continue to be unsatisfied. Therefore, in order that justice may be done +to this line of dissent, I have thought it better to deal with the +matter in a separate Appendix, rather than to hurry it over in the text. +And, as all the difficulties or objections which have been advanced +against the theory of evolution on grounds of palæontology must vary, as +to their strength, with the estimate which is taken touching the degree +of imperfection of the geological record, I will begin by adding a few +paragraphs to what has already been said in the text upon this subject.</p> + +<p>First, then, as to the difficulties in the way of fossils being +<span class="pagenum"><a name="page_422" id="page_422">[422]</a></span> +formed at all. We have already noticed in the text that it is only the +more or less hard parts of organisms which under any circumstances can +be fossilized; and even the hardest parts quickly disintegrate if not +protected from the weather on land, or from the water on the sea-bottom. +Moreover, as Darwin says, “we probably take a quite erroneous view +when we assume that sediment is being deposited over nearly the whole +bed of the sea, at a rate sufficiently quick to embed and preserve +fossil remains. Throughout an enormously large proportion of the ocean, +the bright blue tint of the water bespeaks its purity. The many cases on +record of a formation conformably covered, after an immense interval of +time, by another and a later formation, without the underlying bed +having suffered in the interval any wear and tear, seem explicable only +on the view of the bottom of the sea not rarely lying for ages in an +unaltered condition.” Next, as regards littoral animals, he shows +the difficulty which they must have in becoming fossils, and gives a +striking example in several of the existing species of a sub-family of +cirripedes (<span class='sn'>Chthamalinæ</span>), “which coat the rocks all over the +world in infinite numbers,” yet, with the exception of one species +which inhabits deep water, no vestige of any of them has been found in +any tertiary formation, although it is known that the genus <span class='sn'>Chthamalus</span> +existed through the Chalk period. Lastly, “with respect to the +terrestrial productions which lived through the secondary and +palæozoic periods, it is superfluous to state our evidence is +fragmentary in an extreme degree. For instance, until recently not a +land shell was known belonging to either of these vast periods,” +with one exception; while, “in regard to mammiferous remains, a +glance at the historical table in Lyell’s Manual will bring home +the truth, how accidental and rare has been their preservation, far +better than pages of detail. Nor is their rarity surprising, when we +remember how large a proportion of the bones of tertiary mammals have +been discovered either <span class="pagenum"><a name="page_423" id="page_423">[423]</a></span> +in caves or in lacustrine deposits; and that not a cave or true +lacustrine bed is known belonging to the age of our secondary or +palæozoic formations.”</p> + +<p>But perhaps of even more importance than all these known causes which +prevent the formation of fossils, is the existence of unknown causes +which make for the same result. For example, the Flysch-formation is a +formation of several thousand feet in thickness (as much as 6000 in some +places), and it extends for at least 300 miles from Vienna to +Switzerland; moreover, it consists of shale and sandstone. Therefore, +alike in respect of time, space, and character, it is just such a +formation as we should expect to find highly rich in fossils; yet, +“although this great mass has been most carefully searched, no +fossils, except a few vegetable remains, have been found.”</p> + +<p>So much then for the difficulty, so to speak, which nature experiences +in the manufacture of fossils. Probably not one per cent. of the species +of animals which have inhabited the earth has left a single individual +as a fossil, whereby to record its past existence.</p> + +<p>But of even more importance than this difficulty of making fossils in +the first instance, is the difficulty of preserving them when they are +made. The vast majority of fossils have been formed under water, and a +large proportional number of these—whether the animals were marine, +terrestrial, or inhabitants of fresh water—have been formed in +sedimentary deposits either of sand, gravel, or other porous material. +Now, where such deposits have been afterwards raised into the air for +any considerable time—and this has been more or less the case with all +deposits which are available for exploration—their fossiliferous +contents will have been, as a general rule, dissolved by the percolation +of rain-water charged with carbonic acid. Similarly, sea-water has +recently been found to be a surprisingly strong solvent of calcareous +material: hence, Saturn-like, the ocean devours her own progeny +<span class="pagenum"><a name="page_424" id="page_424">[424]</a></span> +as far as shells and bones of all kinds are concerned—and this to +an extent of which we have probably no adequate conception.</p> + +<p>Of still greater destructive influence, however, than these solvent +agencies in earth and sea, are the erosive agencies of both. Any one who +watches the pounding of the waves upon the shore; who then observes the +effect of it upon the rocks broken into shingle, and on the shingle +reduced to sand; who, looking behind him at the cliffs, sees there the +evidence of the gradual advance of this all-pulverising power—an +advance so gradual that no yard of it is accomplished until within that +yard the “white teeth” have eaten well into the +“bowels of the earth"; who then reflects that this process is +going on simultaneously over hundreds of thousands of miles of +coast-lines throughout the world; and who finally extends his mental +vision from space to time, by trying dimly to imagine what this +ever-roaring monster must have consumed during the hundreds of millions +of years that slowly rising and slowly sinking continents have exposed +their whole areas to her jaws; whoever thus observes and thus reflects +must be a dull man, if he does not begin to feel that in the presence of +such a destroyer as this we have no reason to wonder at a frequent +silence in the testimony of the rocks.</p> + +<p>But although the erosive agency of the sea is thus so inconceivably +great, it is positively small if compared with erosive agencies on land. +The constant action of rain, wind, and running water, in wearing down +the surfaces of all lands into “the dust of continents to be"; the +disintegrating effects on all but the very hardest rocks of winter +frosts alternating with summer heats; the grinding power of ice in +periods of glaciation; and last, but not least, the wholesale melting up +of sedimentary formations whenever these have sunk for any considerable +distance beneath the earth’s surface:—all these agencies taken +together constitute so prodigious a sum of energies combined through +immeasureable <span class="pagenum"><a name="page_425" id="page_425">[425]</a></span> +ages in their common work of destruction, that when we try to realise +what it must amount to, we can scarcely fail to wonder, not that the +geological record is highly imperfect, but that so much of the record +has survived as we find to have been the case. And, if we add to these +erosive and solvent agencies on land the erosive and solvent agencies of +the sea, we may almost begin to wonder that anything deserving the name +of a geological record is in existence at all.</p> + +<p>That such estimates of the destructive powers of nature are not mere +matters of speculative reasoning may be amply shown by stating one +single fact, which, like so many others where the present subject is +concerned, we owe to the generalizations of Darwin. Plutonic rocks, +being those which have emerged from subterranean heat of melting +intensity, must clearly at some time or another have lain beneath the +whole thickness of sedimentary deposits, which at that time occupied any +part of the earth’s surface where we now find the Plutonic rocks +exposed to view. Or, in other words, wherever we now find Plutonic rocks +at the surface of the earth, we must conclude that all the sedimentary +rocks by which they were covered when in a molten state have since been +entirely destroyed; several vertical miles of the only kinds of rocks in +which fossils can possibly occur must in all such cases have been +abolished <i>in toto</i>. Now, in many parts of the world metamorphic +rocks—which have thus gradually risen from Plutonic depths, while miles +of various other rock-formations have been removed from their now +exposed surfaces—cover immense areas, and therefore testify by their +present horizontal range, no less than by their previously vertical +depth, to the enormous scale on which a total destruction has taken +place of everything that once lay above them. For instance, the granitic +region of Parime is at least nineteen times the size of Switzerland; a +similar region south of the Amazon is probably larger than France, +<span class="pagenum"><a name="page_426" id="page_426">[426]</a></span> +Spain, Italy, and Great Britain all put together; and, more remarkable +still, over the area of the United States and Canada, granitic rocks +exceed in the proportion of 19 to 12½ the whole of the newer +Palæozoic formations. Lastly, after giving these examples, Darwin adds +the important consideration, that “in many regions the metamorphic +and granitic rocks would be found much more widely extended than they +appear to be, if all the sedimentary beds were removed which rest +unconformably on them, and which could not have formed part of the +original mantle under which they were crystallized.”</p> + +<p>The above is a brief condensation of the already condensed statement +which Darwin has given of the imperfection of the geological record; but +I think it is enough to show, in a general way, how precarious must be +the nature of any objections to the theory of evolution which are +founded merely upon the silence of palæontology in cases where, if the +record were anything like complete, we should be entitled to expect from +it some positive information. But, as we have seen in the text, +imperfect though the record be, in as far as it furnishes positive +information at all, this is well-nigh uniformly in favour of the theory; +and therefore, even on grounds of palæontology alone, it appears to me +that Darwin is much too liberal where he concludes his discussion by +saying,—"Those who believe that the geological record is in any degree +perfect, will undoubtedly at once reject the theory.” If in any +measure reasonable, such persons ought rather to examine their title to +such a belief; and even if they disregard the consensus of testimony +which is yielded by all the biological sciences to the theory of +evolution, they ought at least to hold their judgment in suspense until +they shall have not only set against the apparently negative testimony +which is yielded by geology its unquestionably positive testimony, but +also well considered the causes which may—or rather must—have so +gravely impaired the geological record.</p> + +<p><span class="pagenum"><a name="page_427" id="page_427">[427]</a></span> +However, be this as it may, I will now pass on to consider the +difficulties and objections which have been brought against the theory +on grounds of palæontology.</p> + +<p>These may be classified under four heads. First, the absence of varietal +links between allied species; second, the sudden appearance of whole +groups of species—not only as genera and families, but even sometimes +as orders and classes—without any forms leading up to them; third, the +occurrence of highly organized types at much lower levels of geological +strata than an evolutionist would antecedently expect; and, fourth, the +absence of fossils of any kind lower down than the Cambrian strata.</p> + +<p>Now all these objections depend on estimates of the imperfection of the +geological record much lower than that which is formed by Darwin. +Therefore I have arranged the objections in their order of difficulty in +this respect, or in the order that requires successively increasing +estimates of the imperfection of the record, if they are to be +successively answered.</p> + +<p>I think that the first of them has been already answered in the text, by +showing that even a very moderate estimate of the imperfection of the +record is enough to explain why intermediate <i>varieties</i>, connecting +allied <i>species</i>, are but comparatively seldom met with. Moreover it was +shown that in some cases, where shells are concerned, remarkably +well-connected series of such varieties have been met with. And the same +applies to species and genera in certain other cases, as in the equine +family.</p> + +<p>But no doubt a greater difficulty arises where whole groups of species +and genera, or even families and orders, appear to arise suddenly, +without anything leading up to them. Even this the second difficulty, +however, admits of being fully met, when we remember that in very many +cases it has been proved, quite apart from the theory of descent, that +superjacent formations have been separated from one another by +<span class="pagenum"><a name="page_428" id="page_428">[428]</a></span> +wide intervals of time. And even although it often happens that +intermediate deposits which are absent in one part of the world are +present in another, we have no right to assume that such is always the +case. Besides, even if it were, we should have no right further to +assume that the faunas of widely separated geographical areas were +identical during the time represented by the intermediate formation. +Yet, unless they were identical, we should not expect the fossils of the +intermediate formation, where extant, to yield evidence of what the +fossils would have been in this same formation elsewhere, had it not +been there destroyed. Now, as a matter of fact, “geological +formations of each region are almost invariably intermittent"; and +although in many cases a more or less continuous record of past forms of +life can be obtained by comparing the fossils of one region and +formation with those of another region and adjacent formations, it is +evident (from what we know of the present geographical distribution of +plants and animals) that not a few cases there must have been where the +interruption of the record in one region cannot be made good by thus +interpolating the fossils of another region. And we must remember it is +by selecting the cases where this cannot be done that the objection +before us is made to appear formidable. In other words, <i>unless</i> whole +groups of new species which are unknown in formation A appear suddenly +in formation C of one region (X), where the intermediate formation B is +absent; and <i>unless</i> in some other region (Y), where B is present, the +fossiliferous contents of B fail to supply the fossil ancestry of the +new species in A (X); <i>unless</i> such a state of matters is found to +obtain, the objection before us has nothing to say. But at best this is +negative evidence; and, in order to consider it fairly, we ought to set +against it the cases where an interposition of fossils found in B (Y) +<i>does</i> furnish the fossil ancestry of what would <i>otherwise</i> have been +an abrupt appearance of whole groups of new species in A (X). Now such +<span class="pagenum"><a name="page_429" id="page_429">[429]</a></span> +cases are neither few nor unimportant, and therefore they deprive the +objection of the force it would have had if the selected cases to the +contrary were the general rule.</p> + +<p>In addition to these considerations, the following, some of which are of +a more special kind, appear to me so important that I will quote them +almost <i>in extenso</i>.</p> + +<div class='blockquot'><p>We continually forget how large the world is, compared with the +area over which our geological formations have been carefully +examined: we forget that groups of species may elsewhere have long +existed, and have slowly multiplied, before they invaded the +ancient archipelagoes of Europe and the United States. We do not +make due allowance for the intervals of time which have elapsed +between our consecutive formations,—longer perhaps in many cases +than the time required for the accumulation of each formation. +These intervals will have given time for the multiplication of +species from some one parent form; and, in the succeeding +formation, such groups of species will appear as if suddenly +created.</p> + +<p>I may here recall a remark formerly made, namely, that it might +require a long succession of ages, to adapt an organism to some new +and peculiar line of life, for instance, to fly through the air; +and consequently that the transitional form would often long remain +confined to some one region; but that, when this adaptation had +once been effected, and a few species had thus acquired a great +advantage over other organisms, a comparatively short time would be +necessary to produce many divergent forms, which would spread +rapidly and widely throughout the world....</p> + +<p>In geological treatises, published not many years ago, mammals were +always spoken of as having abruptly come in at the commencement of +the tertiary series. And now one of the richest known accumulations +of fossil mammals belongs to the middle of the secondary series; +and true mammals have been discovered in the new red sandstone at +nearly the commencement of this great series. Cuvier used to urge +that no monkey occurred in any tertiary stratum; but now extinct +species have been discovered in India, South America, and in Europe +as far <span class="pagenum"><a name="page_430" id="page_430">[430]</a></span> +back as the miocene stage. Had it not been for the rare accident of the +preservation of footsteps in the new red sandstone of the United States, +who would have ventured to suppose that, no less than at least thirty +kinds of bird-like animals, some of gigantic size, existed during that +period? Not a fragment of bone has been discovered in these beds. Not +long ago palæontologists maintained that the whole class of birds +came suddenly into existence during the eocene period; but now we know, +on the authority of Professor Owen, that a bird certainly lived during +the deposition of the upper green-sand. And still more recently that +strange bird, the Archeopteryx ... has been discovered in the oolitic +slates of Solenhofen. Hardly any recent discovery shows more forcibly +than this, how little we as yet know of the former inhabitants of the +world.</p> + +<p>I may give another instance, which, from having passed under my own +eyes, has much struck me. In a memoir on Fossil Sessile Cirripedes, +I stated that, from the number of existing and extinct tertiary +species; from the extraordinary abundance of the individuals of +many species all over the world from the Arctic regions to the +equator, inhabiting various zones of depths from the upper tidal +limits to 50 fathoms; from the perfect manner in which specimens +are preserved in the oldest tertiary beds; from the ease with which +even a fragment of a valve can be recognized; from all these +circumstances, I inferred that had sessile cirripedes existed +during the secondary periods, they would certainly have been +preserved and discovered; and as not one species had then been +discovered in beds of this age, I concluded that this great group +had been suddenly developed at the commencement of the tertiary +series. This was a sore trouble to me, adding as I thought one more +instance of the abrupt appearance of a great group of species. But +my work had hardly been published, when a skilful palæontologist, +M. Bosquet, sent me a drawing of a perfect specimen of an +unmistakeable sessile cirripede, which he had himself extracted +from the chalk of Belgium. And, as if to make the case as striking +as possible, this sessile cirripede was a Chthamalus, a very +common, large, and ubiquitous genus, of which not one specimen has +as yet been found even in any tertiary stratum. Still more +recently, a Pyrgoma, a member of a distinct sub-family <span class="pagenum"><a name="page_431" id="page_431">[431]</a></span> +of sessile cirripedes, has been discovered by Mr. Woodward in the upper +chalk; so that we now have abundant evidence of the existence of this +group of animals during the secondary period.</p> + +<p>The case most frequently insisted on by palæontologists of the +apparently sudden appearance of a whole group of species, is that +of the teleostean fishes, low down, according to Agassiz, in the +Chalk period. This group includes the large majority of existing +species. But certain Jurassic and Triassic forms are now commonly +admitted to be teleostean; and even some palæozoic forms have been +thus classed by one high authority. If the teleosteans had really +appeared suddenly in the northern hemisphere, the fact would have +been highly remarkable; but it would not have formed an insuperable +difficulty, unless it could likewise have been shown that at the +same period the species were suddenly and simultaneously developed +in other quarters of the world. It is almost superfluous to remark +that hardly any fossil fish are known from south of the equator; +and by running through Pictet’s Palæontology it will be seen +that very few species are known from several formations in Europe. +Some few families of fish now have a confined range; the teleostean +fish might formerly have had a similarly confined range, and after +having been largely developed in some one sea, might have spread +widely. Nor have we any right to suppose that the seas of the world +have always been so freely open from south to north as they are at +present. Even at this day, if the Malay Archipelago were converted +into land, the tropical parts of the Indian Ocean would form a +large and perfectly enclosed basin, in which any great group of +marine animals might be multiplied; and here they would remain +confined, until some of the species became adapted to a cooler +climate, and were enabled to double the southern capes of Africa or +Australia, and thus reach other and distant seas.</p> + +<p>From these considerations, from our ignorance of the geology of +other countries beyond the confines of Europe and the United +States; and from the revolution in our palæontological knowledge +effected by the discoveries of the last dozen years, it seems to me +to be about as rash to dogmatize on the succession of organic +<span class="pagenum"><a name="page_432" id="page_432">[432]</a></span> +forms throughout the world, as it would be for a naturalist to land for +five minutes on some one barren point in Australia, and then to discuss +the number and range of its productions<a name="FNanchor_53_53" id="FNanchor_53_53"></a><a href="#Footnote_53_53" class="fnanchor">[53]</a>.</p></div> + +<p>In view of all the foregoing facts and considerations, it appears to me +that the second difficulty on our list is completely answered. Indeed, +even on a moderate estimate of the imperfection of the geological +record, the wonder would have been if many cases had <i>not</i> occurred +where groups of species present the fictitious appearance of having been +suddenly and simultaneously created in the particular formations where +their remains now happen to be observable.</p> + +<p>Turning next to the third objection, there cannot be any question that +every here and there in the geological series animals occur of a much +higher grade zoologically than the theory of evolution would have +expected to find in the strata where they are found. At any rate, +speaking for myself, I should not have antecedently expected to meet +with such highly differentiated insects as butterflies and dragonflies +in the middle of the Secondaries: still less should I have expected to +encounter beetles, cockroaches, spiders, and May-flies in the upper and +middle Primaries—not to mention an insect and a scorpion even in the +lower. And I think the same remark applies to a whole sub-kingdom in the +case of Vertebrata. For although it is only the lowest class of the +sub-kingdom which, so far as we positively know, was represented in the +Devonian and Silurian formations, we must remember, on the one hand, +that even a cartilaginous or ganoid fish belongs to the highest +sub-kingdom of the animal series; and, on the other hand, that such +animals are thus proved to have abounded in the very lowest strata where +there is good evidence of there having been any forms of life at all. +Lastly, the fact that Marsupials occur in the Trias, <span class="pagenum"><a name="page_433" id="page_433">[433]</a></span> +coupled with the fact that the still existing Monotremata are what may +be termed animated fossils, referring us by their lowly type of +organization to some period enormously more remote,—these facts +render it practically certain that some members of this very highest +class of the highest sub-kingdom must have existed far back in the +Primaries.</p> + +<p>These things, I say, I should not have expected to find, and I think all +other evolutionists ought to be prepared to make the same +acknowledgment. But as these things have been found, the only possible +way of accounting for them on evolutionary principles is by supposing +that the geological record is even more imperfect than we needed to +suppose in order to meet the previous objections. I cannot see, however, +why evolutionists should be afraid to make this acknowledgment. For I do +not know any reason which would lead us to suppose that there is any +common measure between the distances marked on our tables of geological +formations, and the times which those distances severally represent. Let +the reader turn to the table on page <a href="#page_163">163</a>, and then +let him say why the 30,000 feet of so-called Azoic rocks may not +represent a greater duration of time than does the thickness of all the +Primary rocks above them put together. For my own part I believe that +this is probably the case, looking to the enormous ages during which +these very early formations must have been exposed to destructive +agencies of all kinds, now at one time and now at another, in different +parts of the world. And, of course, we are without any means of +surmising what ranges of time are represented by the so-called Primeval +rocks, for the simple reason that they are non-sedimentary, and +non-sedimentary rocks cannot be expected to contain fossils.</p> + +<p>But, it will be answered, the 30,000 feet of Azoic rocks, lying above +the Primeval, <i>are</i> sedimentary to some extent: they are not all +completely metamorphic: yet they are all destitute of fossils. This is +the fourth and last difficulty <span class="pagenum"><a name="page_434" id="page_434">[434]</a></span> +which has to be met, and it can only be met by the considerations which +have been advanced by Lyell and Darwin. The former says:—</p> + +<div class='blockquot'><p>The total absence of any trace of fossils has inclined many +geologists to attribute the origin of the most ancient strata to an +azoic period, or one antecedent to the existence of organic beings. +Admitting, they say, the obliteration, in some cases, of fossils by +plutonic action, we might still expect that traces of them would +oftener be found in certain ancient systems of slate, which can +scarcely be said to have assumed a crystalline structure. But in +urging this argument it seems to be forgotten that there are +stratified formations of enormous thickness, and of various ages, +some of them even of tertiary date, and which we know were formed +after the earth had become the abode of living creatures, which +are, nevertheless, in some districts, entirely destitute of all +vestiges of organic bodies<a name="FNanchor_54_54" id="FNanchor_54_54"></a><a href="#Footnote_54_54" class="fnanchor">[54]</a>.</p></div> + +<p>He then proceeds to mention sundry causes (in addition to plutonic +action) which are adequate to destroy the fossiliferous contents of +stratified rocks, and to show that these may well have produced enormous +destruction of organic remains in these oldest of known formations.</p> + +<p>Darwin’s view is that, during the vast ages of time now under +consideration, it is probable that the distribution of sea and land over +the earth’s surface has not been uniformly the same, even as +regards oceans and continents. Now, if this were the case, “it +might well happen that strata which had subsided some miles nearer to +the centre of the earth, and which had been pressed on by an enormous +weight of superincumbent water, might have undergone far more +metamorphic action than strata which have always remained nearer to the +surface. The immense areas in some parts of the world, for instance in +South America, of naked metamorphic rocks, which must have been heated +under great pressure, have always seemed to me to require <span class="pagenum"><a name="page_435" id="page_435">[435]</a></span> +some special explanation; and we may perhaps believe that we see, in +these large areas, the many formations long anterior to the Cambrian +epoch in a completely metamorphosed and denuded condition<a name="FNanchor_55_55" id="FNanchor_55_55"></a><a href="#Footnote_55_55" class="fnanchor">[55]</a>.” +The probability of this view he sustains by certain general +considerations, as well as particular facts touching the geology of +oceanic islands, &c.</p> + +<p>On the whole, then, it seems to me but reasonable to conclude, with +regard to all four objections in question, as Darwin concludes with +regard to them:—</p> + +<div class='blockquot'><p>For my part, following out Lyell’s metaphor, I look at the +geological record as a history of the world imperfectly kept, +written in a changing dialect; of this history we possess the last +volume alone, relating only to two or three countries. Of this +volume, only here and there a short chapter has been preserved; and +of each page only here and there a few lines. Each word of the +slowly-changing language, more or less different in the successive +chapters, may represent the forms of life, which are entombed in +our consecutive formations, and which falsely appear to us to have +been abruptly introduced. On this view, the difficulties above +discussed are greatly diminished, or even disappear<a name="FNanchor_56_56" id="FNanchor_56_56"></a><a href="#Footnote_56_56" class="fnanchor">[56]</a>.</p></div> + +<p>As far as I can see, the only reasonable exception that can be taken to +this general view of the whole matter, is one which has been taken from +the side of astronomical physics.</p> + +<p>Put briefly, it is alleged by one of the highest authorities in this +branch of science, that there cannot have been any such enormous reaches +of unrecorded time as would be implied by the supposition of there +having been a lost history of organic evolution before the Cambrian +period. The grounds of this allegation I am not qualified to examine; +but in a general way I agree with Prof. Huxley in feeling that, from the +very nature of the case, they are necessarily <span class="pagenum"><a name="page_436" id="page_436">[436]</a></span> +precarious,—and this in so high a degree that any conclusions +raised on such premises are not entitled to be deemed formidable<a name="FNanchor_57_57" id="FNanchor_57_57"></a><a href="#Footnote_57_57" class="fnanchor">[57]</a>.</p> + +<hr class='minor' /> + +<p>Turning now to plants, the principal and the ablest opponent of the +theory of evolution is here unquestionably Mr. Carruthers<a name="FNanchor_58_58" id="FNanchor_58_58"></a><a href="#Footnote_58_58" class="fnanchor">[58]</a>. +The difficulties which he adduces may be classified under three heads, +as follows:—</p> + +<p>1. There is no evidence of change in specific forms of existing plants. +Not only are the numerous species of plants which have been found in +Egyptian mummies indistinguishable from their successors of to-day; but, +what is of far more importance, a large number of our own indigenous +plants grew in Great Britain during the glacial period (including under +this term the warm periods between those of successive glaciations), and +in no one case does it appear that any modification of specific type has +occurred. This fact is particularly remarkable as regards leaves, +because on the one hand they are the organs of plants which are most +prone to vary, while on the other hand they are likewise the organs +which lend themselves most perfectly to the process of fossilization, so +that all details of their structure can be minutely observed in the +fossil state. Yet the interval since the glacial period, although not a +long one geologically speaking, is certainly what may be called an +appreciable portion of time in the history of Dicotyledonous plants +since their first appearance in the Cretaceous epoch. Again, if we +extend this kind of enquiry so as to include the world as a whole, a +number of other species of plants dating from the glacial epoch are +found to tell the same story—notwithstanding that, in the opinion of +Mr. Carruthers, they must all have undergone many changes of environment +<span class="pagenum"><a name="page_437" id="page_437">[437]</a></span> +while advancing before, and retreating after, successive glaciations in +different parts of the globe. Or, to quote his own words:—"The +various physical conditions which of necessity affected these {41} +species in their diffusion over such large areas of the earth’s +surface in the course of, say, 250,000 years, should have led to the +production of many varieties; but the uniform testimony of the remains +of this considerable pre-glacial flora, as far as the materials admit of +a comparison, is that no appreciable change has taken place.”</p> + +<p>2. There is no appearance of generalized forms among the earliest plants +with which we are acquainted. For example, in the first dry land +flora—the Devonian—we have representatives of the <span class='sn'>Filices</span>, +<span class='sn'>Equisetaceæ</span>, and <span class='sn'>Lycopodiaceæ</span>, +all as highly specialized as their living representatives, and +exhibiting the differential characters of these closely related groups. +Moreover, these plants were even more highly organized than their +existing descendants in regard to their vegetative structure, and in +some cases also in regard to their reproductive organs. So likewise the +Gymnosperms of that time show in their fossil state the same highly +organized woody structure as their living representatives.</p> + +<p>3. Similarly, and more generally, the Dicotyledonous plants, which first +appear in the Cretaceous rocks, appear there suddenly, without any forms +leading up to them—notwithstanding that “we know very well the +extensive flora of the underlying Wealden.” Moreover, we have all +the three great divisions of the Dicotyledons appearing together, and so +highly differentiated that all the species are referred to existing +genera, with the exception of a very few imperfectly preserved, and +therefore uncertain fragments.</p> + +<p>Such being the facts, we may begin by noticing that, even at first +sight, they present different degrees of difficulty. Thus, I cannot see +that there is much difficulty with regard to those in class 2. Only if +we were to take the <span class="pagenum"><a name="page_438" id="page_438">[438]</a></span> +popular (and very erroneous) view of organic evolution as a process +which is always and everywhere bound to promote the specialization of +organic types—only then ought we to see any real difficulty in the +absence of generalized types preceding these existing types. Of course +we may wonder why still lower down in the geological series we do not +meet with more generalized (or ancestral) types; but this is the +difficulty number 3, which we now proceed to examine.</p> + +<p>Concerning the other two difficulties, then, the only possible way of +meeting that as to the absence of any parent forms lower down in the +geological series is by falling back—as in the analogous case of +animals—upon the imperfection of the geological record. Although it is +certainly remarkable that we should not encounter any forms serving to +connect the Dicotyledonous plants of the Chalk with the lower forms of +the underlying Wealden, we must again remember that difficulties thus +depending on the absence of any corroborative record, are by no means +equivalent to what would have arisen in the presence of an adverse +record—such, for instance, as would have been exhibited had the floras +of the Wealden and the Chalk been inverted. But, as the case actually +stands, the mere fact that Dicotyledonous plants, where they first +occur, are found to have been already differentiated into their three +main divisions, is in itself sufficient evidence, on the general theory +of evolution, that there must be a break in the record as hitherto known +between the Wealden and the Chalk. Nor is it easy to see how the +opponents of this theory can prove their negative by furnishing evidence +to the contrary. And although such might justly be deemed an unfair way +of putting the matter, were this the only case where the geological +record is in evidence, it is not so when we remember that there are +numberless other cases where the geological record does testify to +connecting links in a most satisfactory manner. For in view of this +consideration <span class="pagenum"><a name="page_439" id="page_439">[439]</a></span> +the burden of proof is thrown upon those who point to particular cases +where there is thus a conspicuous absence of transitional +forms—the burden, namely, of proving that such cases are not due +merely to a break in the record. Besides, the break in the record as +regards this particular case may be apparent rather than real. For I +suppose there is no greater authority on the pure geology of the subject +than Sir Charles Lyell, and this is what he says of the particular case +in question. “If the passage seem at present to be somewhat sudden +from the flora of the Lower or Neocomian to that of the Upper Cretaceous +period, the abruptness of the change will probably disappear when we are +better acquainted with the fossil vegetation of the uppermost tracts of +the Neocomian and that of the lowest strata of the Gault, or true +Cretaceous series<a name="FNanchor_59_59" id="FNanchor_59_59"></a><a href="#Footnote_59_59" class="fnanchor">[59]</a>.”</p> + +<p>Lastly, the fact of the flora of the glacial epoch not having exhibited +any modifications during the long residence of some of its specific +types in Great Britain and elsewhere, is a fact of some importance to +the general theory of evolution, since it shows a higher degree of +stability on the part of these specific types than might perhaps have +been expected, supposing the theory to be true. But I do not see that +this constitutes a difficulty against the theory, when we have so many +other cases of proved transmutation to set against it. For instance, not +to go further afield than this very glacial flora itself, it will be +remembered that in an earlier chapter I selected it as furnishing +specially cogent proof of the transmutation of species. What, then, is +the explanation of so extraordinary a difference between Mr. +Carruthers’ views and my own upon this point? I believe the +explanation to be that he does not take a sufficiently wide survey of +the facts.</p> + +<p>To begin with, it seems to me that he exaggerates the vicissitudes to +which the species of plants that he calls into <span class="pagenum"><a name="page_440" id="page_440">[440]</a></span> +evidence have been exposed while advancing before, and retreating +after, the ice. Rather do I agree with Darwin that “they would not +have been exposed during their long migrations to any great diversity of +temperature; and as they all migrated in a body together, their mutual +relations will not have been much disturbed; hence, in accordance with +the principles indicated in this volume, these forms will not have been +liable to much modification<a name="FNanchor_60_60" id="FNanchor_60_60"></a><a href="#Footnote_60_60" class="fnanchor">[60]</a>.” +But, be this matter of opinion as it may, a much better test is afforded +by those numerous cases all the world over, where arctic species have +been left stranded on alpine areas by the retreat of glaciation; because +here there is no room for differences of opinion as to a “change +of environment” having taken place. Not to speak of climatic +differences between arctic and alpine stations, consider merely the +changes which must have taken place in the relations of the thus +isolated species to each other, as well as to those of all the foreign +plants, insects, &c., with which they have long been thrown into +close association. If in <i>such</i> cases no variation or transmutation had +taken place since the glacial epoch, then indeed there would have been a +difficulty of some magnitude. But, by parity of reasoning, whatever +degree of difficulty would have been thus presented is not merely +discharged, but converted into at least an equal degree of +corroboration, when it is found that under such circumstances, in +whatever part of the world they have occurred, some considerable amount +of variation and transmutation has always taken place,—and this in +the animals as well as in the plants. For instance, again to quote +Darwin, “If we compare the present Alpine plants and animals of +the several great European mountain-ranges one with another, though many +of the species remain identically the same, some exist as varieties, +some as doubtful forms or sub-species, and some as distinct yet closely +allied species representing each other on <span class="pagenum"><a name="page_441" id="page_441">[441]</a></span> +the several ranges<a name="FNanchor_61_61" id="FNanchor_61_61"></a><a href="#Footnote_61_61" class="fnanchor">[61]</a>.” +Lastly, if instead of considering the case of alpine floras, we take the +much larger case of the Old and New World as a whole, we meet with much +larger proofs of the same general facts. For, “during the slowly +decreasing warmth of the Pliocene period, as soon as the species in +common, which inhabited the New and Old Worlds, migrated south of the +Polar Circle, they will have been completely cut off from each other. +This separation, as far as the more temperate productions are concerned, +must have taken place long ages ago. As the plants and animals migrated +southward, they will have become mingled in one great region with the +native American productions, and would have had to compete with them; +and, in the other great region, with those of the Old World. +Consequently we have here everything favourable for much +modification,—for far more modification than with the Alpine +productions left isolated, within a much more recent period, on the +several mountain ranges and on the arctic lands of Europe and N. +America. Hence it has come, that when we compare the now living +productions of the temperate regions of the New and Old Worlds, we find +very few identical species; but we find in every class many forms, which +some naturalists rank as geographical races, and others as distinct +species; and a host of closely allied or representative forms which are +ranked by all naturalists as specifically distinct<a name="FNanchor_62_62" id="FNanchor_62_62"></a><a href="#Footnote_62_62" class="fnanchor">[62]</a>.”</p> + +<p>In view then of all the above considerations—and especially those +quoted from Darwin—it appears to me that far from raising any +difficulty against the theory of evolution, the facts adduced by Mr. +Carruthers make in favour of it. For when once these facts are taken in +connection with the others above mentioned, they serve to complete the +correspondence between degrees of modification with degrees <span class="pagenum"><a name="page_442" id="page_442">[442]</a></span> +of time on the one hand, and with degrees of evolution, of change of +environment, &c., on the other. Or, in the words of Le Conte, when +dealing with this very subject, “It is impossible to conceive a +more beautiful illustration of the principles we have been trying to +enforce<a name="FNanchor_63_63" id="FNanchor_63_63"></a><a href="#Footnote_63_63" class="fnanchor">[63]</a>.”</p> + +<hr /><p class="pagenum"><a name="page_443" id="page_443">[443]</a></p> + +<h3 class="sc"><a name="Note_A_to_page_257" id="Note_A_to_page_257"></a>Note A to Page <a href="#page_257">257</a>.</h3> + +<p>The passages in Dr. Whewell’s writings, to which allusion is here +made, are somewhat too long to be quoted in the text. But as I think +they deserved to be given, I will here reprint a letter which I wrote to +<i>Nature</i> in March, 1888.</p> + +<div class='blockquot'><p>In his essay on the <i>Reception of the Origin of Species</i>, Prof. +Huxley writes:—</p> + +<p>“It is interesting to observe that the possibility of a fifth +alternative, in addition to the four he has stated, has not dawned +upon Dr. Whewell’s mind” (<i>Life and Lectures of Charles +Darwin</i>, vol. ii, p. 195).</p> + +<p>And again, in the article <i>Science</i>, supplied to <i>The Reign of +Queen Victoria</i>, he says:—</p> + +<p>“Whewell had not the slightest suspicion of Darwin’s +main theorem, even as a logical possibility” (p 365).</p> + +<p>Now, although it is true that no indication of such a logical +possibility is to be met with in the <i>History of the Inductive +Sciences</i>, there are several passages in the <i>Bridgewater Treatise</i> +which show a glimmering idea of such a possibility. Of these the +following are, perhaps, worth quoting. Speaking of the adaptation +of the period of flowering to the length of a year, he says:—</p> + +<p>“Now such an adjustment must surely be accepted as a proof of +design, exercised in the formation of the world. Why should the +solar year be so long and no longer? or, this being such a length, +why should the vegetable cycle be exactly of the same length? Can +this be chance?... And, if not by chance, how otherwise could such +a coincidence occur than by an intentional adjustment of these two +things to one another; by a selection of such an organization in +plants as would fit them to the earth on which they were to grow; +by an adaptation of construction to conditions; of the scale of +construction to the scale of conditions? It cannot be accepted as +an explanation of this fact in the economy of plants, that it is +necessary to their existence; that no plants could possibly have +subsisted, <span class="pagenum"><a name="page_444" id="page_444">[444]</a></span> +and come down to us, except those which were thus suited to their place +on the earth. This is true; but it does not at all remove the necessity +of recurring to design as the origin of the construction by which the +existence and continuance of plants is made possible. A watch could not +go unless there were the most exact adjustment in the forms and +positions of its wheels; yet no one would accept it as an explanation of +the origin of such forms and positions that the watch would not go if +these were other than they were. If the objector were to suppose that +plants were originally fitted to years of various lengths, and that such +only have survived to the present time as had a cycle of a length equal +to our present year, or one which could be accommodated to it, we should +reply that the assumption is too gratuitous and extravagant to require +much consideration.”</p> + +<p>Again, with regard to “the diurnal period,” he adds:—</p> + +<p>“Any supposition that the astronomical cycle has occasioned +the physiological one, that the structure of plants has been +brought to be what it is by the action of external causes, or that +such plants as could not accommodate themselves to the existing day +have perished, would be not only an arbitrary and baseless +assumption, but, moreover, useless for the purposes of explanation +which it professes, as we have noticed of a similar supposition +with respect to the annual cycle.”</p> + +<p>Of course these passages in no way make against Mr. Huxley’s +allusions to Dr. Whewell’s writings in proof that, until the +publication of the <i>Origin of Species</i>, the “main +theorem” of this work had not dawned on any other mind, save +that of Mr. Wallace. But these passages show, even more +emphatically than total silence with regard to the principle of +survival could have done, the real distance which at that time +separated the minds of thinking men from all that was wrapped up in +this principle. For they show that Dr. Whewell, even after he had +obtained a glimpse of the principle “as a logical +possibility,” only saw in it an “arbitrary and baseless +assumption.” Moreover, the passages show a remarkable +juxtaposition of the very terms in which the theory of natural +selection was afterwards formulated. Indeed, if we strike out the +one word “intentional” (which conveys the preconceived +idea of the writer, and thus prevented him from doing justice to +any naturalistic view), all the following parts of the above +quotations might be supposed to have been written by a Darwinian. +“If not by chance, how otherwise could such a coincidence +occur, than by an <i>adjustment</i> of these two things to one another; +by a <i>selection</i> of such an organization in plants as would <i>fit</i> +them to the earth on which they were to grow; by an adaptation of +<i>construction</i> to <i>conditions</i>; of the <i>scale</i> of construction to +the <i>scale</i> of conditions?” Yet he immediately goes on to +<span class="pagenum"><a name="page_445" id="page_445">[445]</a></span> +say: “If the objector were to suppose that plants were originally +<i>fitted</i> to years of various lengths, and that such only have <i>survived</i> +to the present time ... <i>as could be accommodated to it</i> (i. e. the +actual cycle), we should reply that the assumption is too gratuitous and +extravagant to require much consideration.” Was there ever a more +curious exhibition of failure to perceive the importance of a +“logical possibility"? And this at the very time when another mind +was bestowing twenty years of labour on its “consideration.”</p></div> + +<hr /> +<h3 class='sc'><a name="Note_B_to_page_295" id="Note_B_to_page_295"></a>Note B to Page <a href="#page_295">295</a>.</h3> + +<p>Since these remarks were delivered in my lectures as here printed, Mr. +Mivart has alluded to the subject in the following and precisely +opposite sense:—</p> + +<div class='blockquot'><p>Many of the more noteworthy instincts lead us from manifestations +of purpose directed to the maintenance of the individual, to no +less plain manifestations of a purpose directed to the preservation +of the race. But a careful study of the interrelations and +interdependencies which exist between the various orders of +creatures inhabiting this planet shows us yet a more noteworthy +teleology—the existence of whole orders of such creatures being +directed to the service of other orders in various degrees of +subordination and augmentation respectively. This study reveals to +us, as a fact, the enchainment of all the various orders of +creatures in a hierarchy of activities, in harmony with what we +might expect to find in a world the outcome of a First Cause +possessed of intelligence and will<a name="FNanchor_64_64" id="FNanchor_64_64"></a><a href="#Footnote_64_64" class="fnanchor">[64]</a>.</p></div> + +<p>Having read this much, a Darwinian is naturally led to expect that Mr. +Mivart is about to offer some examples of instincts or structures +exemplifying what in the margin he calls the “Hierarchy of +Ministrations.” Yet the only facts he proceeds to adduce are the +sufficiently obvious facts, that the inorganic world existed before the +organic, plants before herbivorous animals, these before carnivorous, +and so on: that is to say, everywhere the conditions to the occurrence +of any given stage of evolution preceded such occurrence, as it is +obvious that they must, if, as of course it is not denied, the +possibility of such occurrence depended on the precedence of such +conditions.<span class="pagenum"><a name="page_446" id="page_446">[446]</a></span> +Now, it is surely obvious that such a “hierarchy of +ministrations” as this, far from telling against the theory of +natural selection, is the very thing which tells most in its favour. The +fact that animals, for instance, only appeared upon the earth after +there were plants for them to feed upon, is clearly a necessity of the +case, whether or not there was any design in the matter. Such +“ministrations,” therefore, as plant-organisms yield to +animal-organisms is just the kind of ministration that the theory of +natural selection requires. Thus far, then, both the +theories—natural selection and super-natural design—have an +equal right to appropriate the facts. But now, if in no one instance can +it be shown that the ministration of plant-life to animal-life is of +such a kind as to subserve the interests of animal-life without at the +same time subserving those of the plant-life itself, then the fact makes +wholly in favour of the naturalistic explanation of such ministration as +appears. If any plants had presented any characters pointing +prospectively to needs of animals without primarily ministering to their +own, then, indeed, there would have been no room for the theory of +natural selection. But as this can nowhere be alleged, the theory of +natural selection finds all the facts to be exactly as it requires them +to be: such ministration as plants yield to animals becomes so much +evidence of natural selection having slowly formed the animals to +appropriate the nutrition which the plants had previously +gathered—and gathered under the previous influence of natural +selection acting on themselves entirely for their own sakes. Therefore I +say it is painfully manifest that “the enchainment of all the +various orders of creatures in a hierarchy of activities,” is +<i>not</i> “in harmony with what we might expect to find in a world the +outcome of a First Cause possessed of intelligence and [beneficent] +will.” So far as any argument from such “enchainment” +reaches, it makes entirely against the view which Mr. Mivart is +advocating. In point of fact, there is a total absence of any such +“ministration” by one “order of creatures” to +the needs of any other order, as the beneficent design theory would +necessarily expect; while such ministration as actually does obtain is +exactly and universally the kind which the naturalistic theory requires.</p> + +<p>Again, quite independently, and still more recently, Mr. Mivart +<span class="pagenum"><a name="page_447" id="page_447">[447]</a></span> +alluded in <i>Nature</i> (vol. xli, p. 41) to the difficulty which the +apparently exceptional case of gall-formation presents to the theory of +natural selection. Therefore I supplied (vol. xli, p. 80) the suggestion +given in the text, viz. that although it appears impossible that the +sometimes remarkably elaborate and adaptive structures of galls can be +due to natural selection acting directly on the plants +themselves—seeing that the adaptation has reference to the needs +of their parasites—it is quite possible that the phenomena may be +due to natural selection acting indirectly on the plants, by always +preserving those individual insects (and larvae) the character of whose +secretions is such as will best induce the particular shapes of galls +that are required. Several other correspondents took part in the +discussion, and most of them accepted the above explanation. Mr. T. D. +A. Cockerell, however, advanced another and very ingenious hypothesis, +showing that there is certainly one conceivable way in which natural +selection might have produced all the phenomena of gall-formation by +acting directly on the plants themselves<a name="FNanchor_65_65" id="FNanchor_65_65"></a><a href="#Footnote_65_65" class="fnanchor">[65]</a>. +Subsequently Mr. Cockerell published another paper upon the subject, +stating his views at greater length. The following is the substance of +his theory as there presented:—</p> + +<div class='blockquot'><p>Doubtless there were internal plant-feeding larvae before there +were galls: and, indeed, we have geological evidence that boring +insects date very far back indeed. The primitive internal feeders, +then, were miners in the roots, stems, twigs, or leaves, such as +occur very commonly at the present day. These miners are +excessively harmful to plant-life, and form a class of the most +destructive insect-pests known to the farmer: they frequently cause +the death of the whole or part of the plant attacked. Now, we may +suppose that the secretions of certain of these insects caused a +swelling to appear where the larvae lived, and on this excrescence +the larvae fed. It is easy to see that the greater the excrescence, +and the greater the tendency of the larvae to feed upon it, instead +of destroying the vital tissues, the smaller is the amount of harm +to the plant. Now the continued life and vitality of the plant is +beneficial to the larvae, and the larger or more perfect the gall, +the greater the amount of available food. Hence natural selection +will have preserved and accumulated the gall-forming tendencies, as +not only beneficial to the larvae, but as a means whereby the +larvae can feed with least harm to the plant. So <span class="pagenum"><a name="page_448" id="page_448">[448]</a></span> +far from being developed for the exclusive benefit of the larvae, it is +easy to see that, allowing a tendency to gall-formation, natural +selection would have developed galls exclusively for the benefit of the +plants, so that they might suffer a minimum of harm from the unavoidable +attacks of insects.</p> + +<p>But here it may be questioned—have we proof that internal feeders +tend to form galls? In answer to this I would point out that +gall-formation is a peculiar feature, and cannot be expected to +arise in every group of internal feeders. But I think we can afford +sufficient proof that wherever it has arisen it has been preserved; +and further, that even the highly complex forms of galls are +evolved from forms so simple that we hesitate to call them galls at +all<a name="FNanchor_66_66" id="FNanchor_66_66"></a><a href="#Footnote_66_66" class="fnanchor">[66]</a>.</p></div> + +<p>The paper then proceeds to give a number of individual cases. No doubt +the principal objection to which Mr. Cockerell’s hypothesis is +open is one that was pointed out by Herr Wetterhan, viz. “the much +greater facility afforded to the indirect action through insects, by the +enormously more rapid succession of generations with the latter than +with many of their vegetable hosts—oaks above all<a name="FNanchor_67_67" id="FNanchor_67_67"></a><a href="#Footnote_67_67" class="fnanchor">[67]</a>.” This +difficulty, however, Mr. Cockerell believes maybe surmounted by the +consideration that a growing plant need not be regarded as a single +individual, but rather as an assemblage of such<a name="FNanchor_68_68" id="FNanchor_68_68"></a><a href="#Footnote_68_68" class="fnanchor">[68]</a>.</p> + +<hr /> +<h3 class='sc'><a name="Note_C_to_page_394" id="Note_C_to_page_394"></a>Note C to Page <a href="#page_394">394</a>.</h3> + +<p>The only remarks that Mr. Wallace has to offer on the <i>pattern of +colours</i>, as distinguished from a mere <i>brilliancy of colour</i>, are added +as an afterthought suggested to him by the late Mr. Alfred Tylor’s +book on <i>Colouration of Animals and Plants</i> (1886). But, in the first +place, it appears to me that Mr. Wallace has formed an altogether +extravagant estimate of the value of this work. For the object of the +work is to show, “that diversified colouration follows the chief +lines of structure, and changes at points, such as the joints, where +function changes.” Now, in publishing this generalization, Mr. +Tylor—who was not a naturalist—took only a very limited view of the +<span class="pagenum"><a name="page_449" id="page_449">[449]</a></span> +facts. When applied to the animal kingdom as a whole, the theory is +worthless; and even within the limits of mammals, birds, and +insects—which are the classes to which Mr. Tylor mainly applies +it—there are vastly more facts to negative than to support it. +This may be at once made apparent by the following brief quotation from +Prof. Lloyd Morgan:—</p> + +<div class='blockquot'><p>It can hardly be maintained that the theory affords us any adequate +explanation of the <i>specific</i> colour-tints of the humming-birds, or +the pheasants, or the Papilionidae among butterflies. If, as Mr. +Wallace argues, the immense tufts of golden plumage in the bird of +paradise owe their origin to the fact that they are attached just +above the point where the arteries and nerves for the supply of the +pectoral muscles leave the interior of the body—and the +physiological rationale is not altogether obvious,—are there no +other birds in which similar arteries and nerves are found in a +similar position? Why have these no similar tufts? And why, in the +birds of paradise themselves, does it require four years ere these +nervous and arterial influences take effect upon the plumage? +Finally, one would inquire how the colour is determined and held +constant in each species. The difficulty of the Tylor-Wallace view, +even as a matter of origin, is especially great in those numerous +cases in which the colour is determined by delicate lines, thin +plates, or thin films of air or fluid. Mr. Poulton, who takes a +similar line of argument in his <i>Colours of Animals</i> (p. 326), lays +special stress on the production of <i>white</i> (pp. 201-202).</p></div> + +<p>As regards the latter point, it may be noticed that not in any part of +his writings, so far as I can find, does Mr. Wallace allude to the +highly important fact of colours in animals being so largely due to +these purely physical causes. Everywhere he argues as if colours were +universally due to pigments; and in my opinion this unaccountable +oversight is the gravest defect in Mr. Wallace’s treatment both of +the facts and the philosophy of colouration in the animal kingdom. For +instance, as regards the particular case of sexual colouration, the +oversight has prevented him from perceiving that his theory of +“brilliancy” as due to “a surplus of vital +energy,” is not so much as logically possible in what must +constitute at least one good half of the facts to which he applies +it—unless he shows that there is some connection between vital energy +and the development of striations, imprisonment of air-bubbles, &c. But +any such connection—so <span class="pagenum"><a name="page_450" id="page_450">[450]</a></span> +essentially important for his theory—he does not even attempt to +show. Lastly, and quite apart from these remarkable oversights, even if +Mr. Tylor’s hypothesis were as reasonable and well-sustained as it +is fanciful and inadequate, still it could not apply to <i>sexual</i> +colouration: it could apply only to colouration as affected by +physiological functions common to both sexes. Yet it is in order to +furnish a “preferable substitute” for Mr. Darwin’s +theory of <i>sexual</i> colouration, that Mr. Wallace adduces the hypothesis +in question as one of “great weight"! In this matter, therefore, I +entirely agree with Poulton and Lloyd Morgan.</p> + +<hr /><p class="pagenum"><a name="page_451" id="page_451">[451]</a></p> +<h2><a name="INDEX" id="INDEX"></a>INDEX</h2> + +<ul class='off'> +<li class='b c mt2'>A.</li> + +<li>Accident, Darwin’s use of the word, <a href="#page_334">334</a>-<a href="#page_340">340</a>; +<ul class='off'><li>beauty due to, <a href="#page_408">408</a>, <a href="#page_409">409</a>.</li></ul></li> + +<li>Achromatin, <a href="#page_126">126</a>-<a href="#page_134">134</a>.</li> + +<li>Acquired characters, <i>see</i> Characters.</li> + +<li><span class='sn'>Acræa eurita</span>, <a href="#page_328">328</a>.</li> + +<li>Adaptation, facts of, in relation to theory of natural selection, <a href="#page_401">401</a>-<a href="#page_403">403</a>, <a href="#page_411">411</a>.</li> + +<li>Adaptive characters, <i>see</i> Characters.</li> + +<li>Æsthetic sense in animals, <a href="#page_380">380</a>-<a href="#page_385">385</a>; +<ul class='off'><li><i>see</i> Beautiful.</li></ul></li> + +<li>Agassiz, Prof. A., on fauna of the Mammoth cave, <a href="#page_70">70</a>.</li> + +<li>Alpine plants, <a href="#page_209">209</a>, <a href="#page_210">210</a>, <a href="#page_440">440</a>-<a href="#page_442">442</a>.</li> + +<li><span class='sn'>Amauris niavius</span>, <a href="#page_328">328</a>.</li> + +<li><span class='sn'>Amblyornis inornata</span>, <a href="#page_381">381</a>-<a href="#page_383">383</a>.</li> + +<li><span class='sn'>Amphioxus</span>, <a href="#page_137">137</a>, <a href="#page_138">138</a>, <a href="#page_145">145</a>, <a href="#page_146">146</a>.</li> + +<li>Analogy, <a href="#page_38">38</a>, <a href="#page_50">50</a>-<a href="#page_65">65</a>, <a href="#page_176">176</a>, <a href="#page_177">177</a>, <a href="#page_347">347</a>-<a href="#page_350">350</a>.</li> + +<li>Anthropoid, <i>see</i> Apes.</li> + +<li>Antlers, <a href="#page_98">98</a>-<a href="#page_100">100</a>, <a href="#page_167">167</a>-<a href="#page_169">169</a>.</li> + +<li>Ants, co-operative instincts of, <a href="#page_268">268</a>; +<ul class='off'><li>leaf-cutting, <a href="#page_332">332</a>;</li> +<li>keeping aphides, <a href="#page_292">292</a>.</li></ul></li> + +<li>Ape, eye of, <a href="#page_75">75</a>; +<ul class='off'><li><span class='sn'>appendix vermiformis</span> of, <a href="#page_84">84</a>-<a href="#page_86">86</a>.</li></ul></li> + +<li>Apes, ears of, compared with those of man, <a href="#page_88">88</a>; +<ul class='off'><li>muscles of, <a href="#page_77">77</a>, <a href="#page_82">82</a>, <a href="#page_83">83</a>;</li> +<li>feet of, <a href="#page_77">77</a>, <a href="#page_78">78</a>;</li> +<li>tail of, compared with that of man, <a href="#page_82">82</a>-<a href="#page_84">84</a>;</li> +<li>hair of, compared with that of man, <a href="#page_89">89</a>-<a href="#page_91">91</a>;</li> +<li>teeth of, compared with those of man, <a href="#page_92">92</a>-<a href="#page_94">94</a>;</li> +<li>flattening of tibiæ of, <a href="#page_95">95</a>, <a href="#page_96">96</a>.</li></ul></li> + +<li>Aphides, <a href="#page_292">292</a>.</li> + +<li><span class='sn'>Appendix vermiformis</span> of man compared with that of orang, <a href="#page_84">84</a>-<a href="#page_86">86</a>.</li> + +<li><span class='sn'>Apteryx</span>, <a href="#page_68">68</a>, <a href="#page_69">69</a>.</li> + +<li><span class='sn'>Archæopteryx</span>, <a href="#page_171">171</a>-<a href="#page_173">173</a>.</li> + +<li>Arctic plants, <a href="#page_209">209</a>, <a href="#page_210">210</a> , <a href="#page_440">440</a>-<a href="#page_442">442</a>.</li> + +<li>Argyll, Duke of, on natural selection, <a href="#page_334">334</a>-<a href="#page_362">362</a> .</li> + +<li>Aristotle, his idea of scientific method, <a href="#page_1">1</a>; +<ul class='off'><li>on classification, <a href="#page_23">23</a>, <a href="#page_24">24</a>.</li></ul></li> + +<li>Arm, distribution of hair on, in man and apes, <a href="#page_89">89</a>-<a href="#page_92">92</a>.</li> + +<li><span class='sn'>Arthropoda</span>, embryology of, <a href="#page_155">155</a>.</li> + +<li>Artificial selection, analogy of, to natural selection, <a href="#page_295">295</a>-<a href="#page_314">314</a>;</li> +<li> pictorial representations of products of, <a href="#page_298">298</a>-<a href="#page_312">312</a>.</li> + +<li>Artiodactyls, <a href="#page_182">182</a>-<a href="#page_191">191</a>.</li> + +<li>Association, principle of, in æsthetics, <a href="#page_404">404</a>-<a href="#page_407">407</a>.</li> + +<li>Aster, <a href="#page_129">129</a>-<a href="#page_133">133</a>.</li> + +<li>Attraction-spheres, <a href="#page_128">128</a>, <a href="#page_132">132</a>, <a href="#page_133">133</a>.</li> + +<li>Australia, fauna of, <a href="#page_204">204</a>, <a href="#page_205">205</a>; +<ul class='off'><li>thriving of exotic species in, <a href="#page_286">286</a>;</li> +<li>portrait of wild dog of, <a href="#page_304">304</a>.</li></ul></li> + +<li>Azores, <a href="#page_224">224</a>, <a href="#page_225">225</a>.</li> + +<li class='b c mt2'>B.</li> + +<li>Bacon, Lord, on scientific method, <a href="#page_2">2</a>.</li> + +<li><span class='sn'>Balanoglossus</span>, <a href="#page_147">147</a>, <a href="#page_148">148</a>.</li> + +<li><span class='sn'>Baptanodon discus</span>, posterior limb of, <a href="#page_179">179</a>-<a href="#page_181">181</a>.</li> + +<li>Barriers, in relation to geographical distribution, <a href="#page_216">216</a>-<a href="#page_224">224</a>.</li> + +<li>Bats, <a href="#page_56">56</a>, <a href="#page_224">224</a>, <a href="#page_226">226</a>, <a href="#page_240">240</a>.</li> + +<li>Battle, law of, <a href="#page_385">385</a>, <a href="#page_386">386</a>.</li> + +<li>Baya-bird, <a href="#page_381">381</a>.<span class="pagenum"><a name="page_452" id="page_452">[452]</a></span></li> + +<li>Bear, skeleton of, <a href="#page_174">174</a>; +<ul class='off'><li>feet of, <a href="#page_178">178</a>.</li></ul></li> + +<li>Beautiful, the, sense of, in animals, <a href="#page_380">380</a>-<a href="#page_385">385</a>; +<ul class='off'><li>standards of, <a href="#page_380">380</a>-<a href="#page_404">404</a>;</li> +<li>Darwin’s explanation of, in organic nature, <a href="#page_379">379</a>-<a href="#page_411">411</a>;</li> +<li>facts of, in inorganic nature in relation to Darwin’s theory of, in organic, <a href="#page_404">404</a>;</li> +<li>often determined by natural selection, <a href="#page_406">406</a>, <a href="#page_407">407</a>;</li> +<li>absent in many plants and animals, <a href="#page_408">408</a>;</li> +<li>in nature often accidental, <a href="#page_409">409</a>-<a href="#page_411">411</a>;</li> +<li>does not exist in organic nature as an end <i>per se</i>, <a href="#page_410">410</a>, <a href="#page_411">411</a>.</li></ul></li> + +<li>Bees, co-operative instincts of, <a href="#page_268">268</a>.</li> + +<li>Beetles, wingless, <a href="#page_68">68</a>-<a href="#page_70">70</a>; +<ul class='off'><li>on oceanic islands, <a href="#page_224">224</a>, <a href="#page_226">226</a>, <a href="#page_229">229</a>, <a href="#page_232">232</a>.</li></ul></li> + +<li>Bell, Dr., on natural theology, <a href="#page_412">412</a>.</li> + +<li>Bell-bird, <a href="#page_396">396</a>-<a href="#page_398">398</a>.</li> + +<li><span class='sn'>Bembidium</span>, <a href="#page_233">233</a>.</li> + +<li>Bermudas, <a href="#page_225">225</a>-<a href="#page_227">227</a>.</li> + +<li>Biology, ideas of method in, <a href="#page_1">1</a>-<a href="#page_9">9</a>.</li> + +<li>Birds, ovum of, <a href="#page_124">124</a>; +<ul class='off'><li>embryology of, <a href="#page_151">151</a>-<a href="#page_155">155</a>;</li> +<li>paleontology of, <a href="#page_163">163</a>-<a href="#page_165">165</a>, <a href="#page_172">172</a>, <a href="#page_173">173</a>;</li> +<li>brain of, <a href="#page_194">194</a>-<a href="#page_197">197</a>;</li> +<li>as carriers of seed, eggs, and small organisms, <a href="#page_217">217</a>, <a href="#page_218">218</a>;</li> +<li>distribution of, <a href="#page_224">224</a>-<a href="#page_240">240</a>;</li> +<li>æsthetic sense of, <a href="#page_380">380</a>-<a href="#page_385">385</a>;</li> +<li>courtship of, <a href="#page_380">380</a>-<a href="#page_385">385</a>.</li></ul></li> + +<li><span class='sn'>Birgus latro</span>, <a href="#page_62">62</a>-<a href="#page_65">65</a>.</li> + +<li>Blood, colour of arterial, <a href="#page_409">409</a>.</li> + +<li>Boar, <i>see</i> Pig.</li> + +<li><span class='sn'>Bombus lapidarius</span>, <a href="#page_331">331</a>.</li> + +<li>Bower-birds, play-houses of, <a href="#page_381">381</a>-<a href="#page_383">383</a>.</li> + +<li>Boyd-Dawkins, on flattening of early human tibiæ, <a href="#page_96">96</a>.</li> + +<li>Brain, palæontology of, <a href="#page_194">194</a>-<a href="#page_197">197</a>.</li> + +<li>British Isles, <i>see</i> Islands.</li> + +<li>Broca, <a href="#page_363">363</a>.</li> + +<li>Bronn, <a href="#page_363">363</a>.</li> + +<li>Budding, <i>see</i> Germination.</li> + +<li>Burdon-Sanderson, Prof., on electric organ of skate, <a href="#page_366">366</a>.</li> + +<li>Butler, Bishop, on argument from ignorance, <a href="#page_41">41</a>.</li> + +<li>Butterflies, defensive colouring of, <a href="#page_321">321</a>-<a href="#page_329">329</a>.</li> + +<li class='b c mt2'>C.</li> + +<li>Cæsalpino, on classification, <a href="#page_24">24</a>.</li> + +<li>Calf, embryology of, <a href="#page_153">153</a>.</li> + +<li>Camel, foot of, <a href="#page_187">187</a>-<a href="#page_191">191</a>.</li> + +<li>Canadian stag, <a href="#page_196">196</a>, <a href="#page_198">198</a>, <a href="#page_199">199</a>.</li> + +<li>Canaries, portraits of, <a href="#page_303">303</a>; +<ul class='off'><li>first mentioned by Gesner, <a href="#page_312">312</a>, <a href="#page_313">313</a>.</li></ul></li> + +<li>Cape de Verde Archipelagoes, fauna of, <a href="#page_228">228</a>.</li> + +<li><span class='sn'>Carcharias melanopterus</span>, <a href="#page_149">149</a>.</li> + +<li>Carruthers, on evolution, <a href="#page_436">436</a>-<a href="#page_442">442</a>.</li> + +<li>Caterpillars, colours and forms of, <a href="#page_319">319</a>, <a href="#page_322">322</a>-<a href="#page_326">326</a>.</li> + +<li>Cattle, portraits of, <a href="#page_311">311</a>.</li> + +<li>Causation, natural, <a href="#page_402">402</a>, <a href="#page_413">413</a>, <a href="#page_414">414</a>.</li> + +<li>Caves, faunas of dark, <a href="#page_70">70</a>-<a href="#page_72">72</a>.</li> + +<li>Cell, physiological, and properties of the, <a href="#page_104">104</a>-<a href="#page_134">134</a>.</li> + +<li><span class='sn'>Cerura vinula</span>, <a href="#page_325">325</a>, <a href="#page_326">326</a>.</li> + +<li><span class='sn'>Cervalces Americanus</span>, <a href="#page_196">196</a>, <a href="#page_198">198</a>, <a href="#page_199">199</a>.</li> + +<li><span class='sn'>Cervus dicrocerus</span>, <span class='sn'>issiodorensis</span>, <span class='sn'>matheronis</span>, <span class='sn'>pardinensis</span>, <span class='sn'>Sedgwickii</span>, <span class='sn'>tetraceros</span>, <a href="#page_168">168</a>.</li> + +<li>Chalmers, Dr., on natural theology, <a href="#page_412">412</a>.</li> + +<li>Chameleons, <a href="#page_317">317</a>.</li> + +<li>Characters, as adaptive, <a href="#page_273">273</a>-<a href="#page_276">276</a>, <a href="#page_286">286</a>-<a href="#page_293">293</a>, <a href="#page_349">349</a>; +<ul class='off'><li>as specific, <a href="#page_274">274</a>-<a href="#page_276">276</a>, <a href="#page_286">286</a>-<a href="#page_295">295</a>;</li> +<li>as congenital and acquired, <a href="#page_274">274</a>-<a href="#page_276">276</a>.</li></ul></li> + +<li><span class='sn'>Chasmorhynchus niveus</span>, and <span class='sn'>C. tricarunculatus</span>, <a href="#page_396">396</a>-<a href="#page_398">398</a>.</li> + +<li><span class='sn'>Chelydra serpentina</span>, anterior limb of, <a href="#page_179">179</a>-<a href="#page_181">181</a>.</li> + +<li>Chick, embryology of, <a href="#page_153">153</a>.</li> + +<li>Chimpanzee, <i>see</i> Apes.</li> + +<li>Chlorophyll, <a href="#page_408">408</a>.</li> + +<li><span class='sn'>Chondracanthus cornutus</span>, <a href="#page_122">122</a>.</li> + +<li>Cirripedes, <a href="#page_430">430</a>.</li> + +<li>Classification, <a href="#page_23">23</a>-<a href="#page_49">49</a>; +<ul class='off'><li>of organic nature by Genesis and Leviticus, <a href="#page_23">23</a>;</li> +<li>artificial and natural, <a href="#page_24">24</a>-<a href="#page_26">26</a>;</li> +<li>empirical rules of, <a href="#page_33">33</a>-<a href="#page_40">40</a>;</li> +<li>Darwin on, <a href="#page_35">35</a>, <a href="#page_36">36</a>, <a href="#page_39">39</a>,<a href="#page_40">40</a>;</li> +<li>form of, a nexus or tree, <a href="#page_29">29</a>-<a href="#page_32">32</a>;</li> +<li>of organic forms like that of languages, <a href="#page_32">32</a>;</li> +<li>single characters in relation to, <a href="#page_37">37</a>;</li> +<li>aggregates of characters in relation to, <a href="#page_35">35</a>-<a href="#page_37">37</a>;</li> +<li>adaptive and non-adaptive characters in relation to, <a href="#page_34">34</a>, <a href="#page_35">35</a>, <a href="#page_38">38</a>, <a href="#page_39">39</a>;<span class="pagenum"><a name="page_453" id="page_453">[453]</a></span></li> +<li>chains of affinities in relation to, <a href="#page_39">39</a>-<a href="#page_40">40</a>;</li> +<li>biological differs from astronomical, <a href="#page_43">43</a>.</li></ul></li> + +<li>Cockerell, on vegetable galls, <a href="#page_447">447</a>, <a href="#page_448">448</a>.</li> + +<li>Colours, of plants and animals in relation to the theory of natural selection, <a href="#page_317">317</a>-<a href="#page_332">332</a>; +<ul class='off'><li>in relation to the theory of sexual selection, <a href="#page_391">391</a>, <a href="#page_392">392</a>, <a href="#page_394">394</a>-<a href="#page_396">396</a>, <a href="#page_408">408</a>-<a href="#page_410">410</a>, <a href="#page_448">448</a>-<a href="#page_450">450</a>.</li></ul></li> + +<li>Colouring, <i>see</i> Recognition marks, Protective, Seasonal, Warning, and Mimicry.</li> + +<li>Congenital characters, <i>see</i> Characters.</li> + +<li>Conjugation, of Protozoa, <a href="#page_115">115</a>-<a href="#page_117">117</a>.</li> + +<li>Continuity, principle of, in nature, <a href="#page_15">15</a>-<a href="#page_21">21</a>.</li> + +<li>Contrivance, Darwin’s use of the word, <a href="#page_281">281</a>.</li> + +<li>Co-operation, mutual, of species alleged, <a href="#page_445">445</a>-<a href="#page_448">448</a>.</li> + +<li>Co-operative instincts, due to natural selection, <a href="#page_267">267</a>, <a href="#page_269">269</a>.</li> + +<li>Cope, Professor, his table of geological formations, <a href="#page_163">163</a>, <a href="#page_164">164</a>; +<ul class='off'><li>his table of palæontological development of feet, vertebral column, and brain, <a href="#page_197">197</a>.</li></ul></li> + +<li>Correlation of growth, <a href="#page_357">357</a>-<a href="#page_362">362</a>.</li> + +<li><span class='sn'>Cossonidæ</span>, <a href="#page_233">233</a>.</li> + +<li>Courtship, <i>see</i> Sexual Selection.</li> + +<li>Crabs, <a href="#page_62">62</a>-<a href="#page_65">65</a>, <a href="#page_139">139</a>.</li> + +<li>Cuttle-fish, <a href="#page_317">317</a>.</li> + +<li>Cuvier, on method in natural history, <a href="#page_3">3</a>-<a href="#page_4">4</a>; +<ul class='off'><li>on monkeys, <a href="#page_429">429</a>.</li></ul></li> + +<li>Cyst, <i>see</i> Encystation.</li> + +<li class='b c mt2'>D.</li> + +<li>Darwin, Charles, his influence on ideas of method, <a href="#page_1">1</a>-<a href="#page_9">9</a>; +<ul class='off'><li>on classification, <a href="#page_35">35</a>, <a href="#page_36">36</a>, <a href="#page_39">39</a>, <a href="#page_40">40</a>;</li> +<li>on vestigial characters in man, <a href="#page_77">77</a>, <a href="#page_86">86</a>, <a href="#page_87">87</a>, <a href="#page_92">92</a>;</li> +<li>on imperfection of geological record, <a href="#page_165">165</a>, and <a href="#APPENDIX_AND_NOTES">Appendix</a>;</li> +<li>on means of dispersal, <a href="#page_216">216</a>, <a href="#page_218">218</a>;</li> +<li>on geographical distribution, <a href="#page_218">218</a>, <a href="#page_219">219</a>;</li> +<li>on fauna of the Galapagos Archipelago, <a href="#page_227">227</a>, <a href="#page_228">228</a>;</li> +<li>on natural selection, <a href="#page_252">252</a>, <a href="#page_253">253</a>, <a href='#page_255'>255</a>, <a href='#page_256'>256</a>, <a href='#page_286'>286</a>, <a href='#page_375'>375</a>, <a href='#page_376'>376</a>;</li> +<li>his use of such words as ‘accident,’ ‘fortuitous,’ ‘purpose,’ ‘contrivance,’ &c., <a href='#page_281'>281</a>, <a href='#page_334'>334</a>-<a href='#page_340'>340</a></li> +<li>on sexual selection, <a href='#page_379'>379</a>-<a href='#page_400'>400</a>.</li></ul></li> + +<li>Darwin, Erasmus, his theory of evolution, <a href='#page_253'>253</a>.</li> + +<li>De Blainville, on the theory of descent, <a href='#page_258'>258</a>.</li> + +<li>De Candolle, on classification, <a href='#page_34'>34</a>.</li> + +<li>Deer, <a href='#page_98'>98</a>, <a href='#page_99'>99</a>, <a href='#page_167'>167</a>-<a href='#page_169'>169</a>, <a href='#page_187'>187</a>, <a href='#page_191'>191</a>, <a href='#page_196'>196</a>, <a href='#page_198'>198</a>, <a href='#page_199'>199</a>.</li> + +<li>Degeneration, <a href='#page_269'>269</a>, <a href='#page_270'>270</a>, <a href='#page_342'>342</a>.</li> + +<li>Delamination, <a href='#page_139'>139</a>.</li> + +<li><span class='sn'>Diadema euryta</span>, <a href='#page_330'>330</a>.</li> + +<li>Diaster, <a href='#page_129'>129</a>-<a href='#page_133'>133</a>.</li> + +<li>Dingo, <i>see</i> Dog.</li> + +<li><span class='sn'>Dinornis</span>, <a href='#page_60'>60</a>, <a href='#page_61'>61</a>.</li> + +<li><span class='sn'>Diptera</span> mimicking <span class='sn'>Hymenoptera</span>, <a href='#page_329'>329</a>.</li> + +<li>Dog, dentition of, <a href='#page_39'>39</a>; +<ul class='off'><li>Dingo, <a href='#page_304'>304</a>;</li> +<li>domesticated varieties of, <a href='#page_305'>305</a>, <a href='#page_307'>307</a>;</li> +<li>hairless, <a href='#page_307'>307</a>;</li> +<li>skulls of, <a href='#page_307'>307</a>.</li></ul></li> + +<li>Duck, logger-headed, <a href='#page_68'>68</a>.</li> + +<li>Dugong, eye of, <a href='#page_75'>75</a>.</li> + +<li class='b c mt2'>E.</li> + +<li>Eagle, eye of, <a href='#page_75'>75</a>.</li> + +<li>Ear, of whales, <a href='#page_65'>65</a>; +<ul class='off'><li>vestigial features of human, <a href='#page_76'>76</a>, <a href='#page_86'>86</a>-<a href='#page_89'>89</a>;</li> +<li>of man and apes compared, <a href='#page_88'>88</a>.</li></ul></li> + +<li>Eaton, Rev. A. E., on wingless insects, <a href='#page_70'>70</a>.</li> + +<li><span class='sn'>Echinodermata</span>, <a href='#page_125'>125</a>-<a href='#page_127'>127</a>, <a href='#page_138'>138</a>, <a href='#page_155'>155</a>.</li> + +<li>Ectoderm, <a href='#page_137'>137</a>-<a href='#page_142'>142</a>.</li> + +<li>Egg, <i>see</i> Ovum.</li> + +<li>Eimer, <a href='#page_363'>363</a>.</li> + +<li><span class='sn'>Elaps fulvius</span> imitated by non-venomous snakes, <a href='#page_330'>330</a>.</li> + +<li>Electric organs, <a href='#page_365'>365</a>-<a href='#page_373'>373</a>.</li> + +<li>Elephant, foot of, <a href='#page_185'>185</a>, <a href='#page_186'>186</a>; +<ul class='off'><li>rate of propagation of, <a href='#page_261'>261</a>, <a href='#page_262'>262</a>.</li></ul></li> + +<li>Elk, <a href='#page_196'>196</a>-<a href='#page_198'>198</a>, <a href='#page_199'>199</a>.</li> + +<li>Embryo, human, <i>see</i> Man.</li> + +<li>Embryogeny, <i>see</i> Ontogeny.</li> + +<li>Embryology, <a href='#page_98'>98</a>-<a href='#page_155'>155</a>.</li> + +<li>Embryos, comparative series of, <a href='#page_152'>152</a>, <a href='#page_153'>153</a>.<span class="pagenum"><a name="page_454" id="page_454">[454]</a></span></li> + +<li><i>Encyclopædia Britannica</i>, eighth ed., on instinct, <a href='#page_289'>289</a>-<a href='#page_291'>291</a>.</li> + +<li>Encystation of Protozoa, <a href='#page_115'>115</a>.</li> + +<li>Endoderm, <a href='#page_137'>137</a>-<a href='#page_142'>142</a>.</li> + +<li>Equatorial plate, <a href='#page_129'>129</a>.</li> + +<li><span class='sn'>Equus</span>, <i>see</i> Horse.</li> + +<li><span class='sn'>Erythrolamprus venustissimus</span>, <a href='#page_330'>330</a>.</li> + +<li>Evolution, organic, fact of, Section I; +<ul class='off'><li>Method of, Section II;</li> +<li>ideas upon, prior to Darwin, <a href='#page_253'>253</a>-<a href='#page_258'>258</a>;</li> +<li>divergent, <a href='#page_266'>266</a>, <a href='#page_267'>267</a>.</li></ul></li> + +<li>Ewart, Professor Cossar, on electric organ of skate, <a href='#page_364'>364</a>, <a href='#page_367'>367</a>.</li> + +<li>Existence, <i>see</i> Struggle for.</li> + +<li>Eye, of octopus, <a href='#page_57'>57</a>, <a href='#page_58'>58</a>, <a href='#page_347'>347</a>-<a href='#page_350'>350</a>; +<ul class='off'><li>absence of, in dark cave animals, <a href='#page_70'>70</a>-<a href='#page_72'>72</a>;</li> +<li>nictitating membrane of, <a href='#page_74'>74</a>, <a href='#page_75'>75</a>;</li> +<li>development of, from cutaneous nerve-ending, <a href='#page_352'>352</a>-<a href='#page_354'>354</a>.</li></ul></li> + +<li class='b c mt2'>F.</li> + +<li>Feet, <a href='#page_51'>51</a>-<a href='#page_59'>59</a>, <a href='#page_66'>66</a>, <a href='#page_77'>77</a>-<a href='#page_80'>80</a>, <a href='#page_174'>174</a>-<a href='#page_192'>192</a>, <a href='#page_197'>197</a>.</li> + +<li>Fertilization of ova, <a href='#page_127'>127</a>, <a href='#page_128'>128</a>; +<ul class='off'><li>of flowers by insects, <a href='#page_406'>406</a>.</li></ul></li> + +<li>Fish, embryology of, <a href='#page_143'>143</a>-<a href='#page_155'>155</a>; +<ul class='off'><li>palæontology of, <a href='#page_163'>163</a>, <a href='#page_165'>165</a>, <a href='#page_169'>169</a>-<a href='#page_171'>171</a>;</li> +<li>brain of, <a href='#page_194'>194</a>-<a href='#page_197'>197</a>;</li> +<li>distribution of, <a href='#page_224'>224</a>-<a href='#page_246'>246</a>;</li> +<li>flying, <a href='#page_355'>355</a>.</li></ul></li> + +<li>Fission, reproduction by, <a href='#page_106'>106</a>, <a href='#page_107'>107</a>.</li> + +<li>Flat fish, <a href='#page_317'>317</a>.</li> + +<li>Float, <i>see</i> Swim-bladder.</li> + +<li>Flowers, fertilization of, by insects, <a href='#page_406'>406</a>.</li> + +<li>Fly, imitating a wasp, <a href='#page_329'>329</a>.</li> + +<li>Flying-fish, and squirrels, <a href='#page_355'>355</a>.</li> + +<li><span class='sn'>Foraminifera</span>, <a href='#page_346'>346</a>.</li> + +<li>Forbes, H. O., on scapulo-coracoid bones of <span class='sn'>Dinornis</span>, <a href='#page_60'>60</a>.</li> + +<li>Fortuitous, Darwin’s use of the word, <a href='#page_340'>340</a>.</li> + +<li>Fossils, <i>see</i> Palæontology.</li> + +<li>Frogs, <a href='#page_317'>317</a>.</li> + +<li class='b c mt2'>G.</li> + +<li>Galapagos Islands, <a href='#page_227'>227</a>-<a href='#page_231'>231</a>, <a href='#page_236'>236</a>, <a href='#page_237'>237</a>.</li> + +<li><span class='sn'>Galeus</span>, eye of, <a href='#page_75'>75</a>.</li> + +<li>Galls, vegetable, <a href='#page_293'>293</a>-<a href='#page_295'>295</a>, <a href='#page_446'>446</a>-<a href='#page_448'>448</a>.</li> + +<li>Gastræa, <a href='#page_137'>137</a>-<a href='#page_140'>140</a>.</li> + +<li><span class='sn'>Gastrophysema</span>, <a href='#page_138'>138</a>.</li> + +<li>Gastrulation, <a href='#page_137'>137</a>, <a href='#page_140'>140</a>.</li> + +<li>Gegenbaur, <a href='#page_147'>147</a>, <a href='#page_181'>181</a>.</li> + +<li>Gemmation, reproduction by, <a href='#page_106'>106</a>, <a href='#page_107'>107</a>, <a href='#page_110'>110</a>, <a href='#page_111'>111</a>.</li> + +<li>Generalization, <a href='#page_5'>5</a>.</li> + +<li>Generalized types, <a href='#page_33'>33</a>.</li> + +<li>Genesis, classification of organic nature in, <a href='#page_23'>23</a>.</li> + +<li>Genial tubercle, <a href='#page_96'>96</a>.</li> + +<li>Geographical distribution, <a href='#page_204'>204</a>-<a href='#page_248'>248</a>; +<ul class='off'><li><i>see</i> Glacial period, Barriers Transport of organisms, Oceanic islands, &c.</li></ul></li> + +<li>Geology, record of imperfect, <a href='#page_156'>156</a>-<a href='#page_160'>160</a>, and <a href="#APPENDIX_AND_NOTES">Appendix</a>; +<ul class='off'><li><i>see</i> Palæontology.</li></ul></li> + +<li>Germs, prophetic, <a href='#page_272'>272</a>, <a href='#page_351'>351</a>-<a href='#page_362'>362</a>.</li> + +<li>Gesner, on classification, <a href='#page_24'>24</a>; +<ul class='off'><li>on canaries, <a href='#page_313'>313</a>.</li></ul></li> + +<li>Gill-arches, <a href='#page_146'>146</a>, <a href='#page_147'>147</a>, <a href='#page_150'>150</a>, <a href='#page_151'>151</a>.</li> + +<li>Gill-slits, <a href='#page_146'>146</a>, <a href='#page_147'>147</a>, <a href='#page_150'>150</a>-<a href='#page_153'>153</a>.</li> + +<li>Gills, of young salamanders, <a href='#page_102'>102</a>; +<ul class='off'><li>origin of, in embryo, <a href='#page_144'>144</a>;</li> +<li>of fish, <a href='#page_150'>150</a>, <a href='#page_152'>152</a>.</li></ul></li> + +<li>Giraffe, neck of, in relation to Lamarck’s theory, <a href='#page_254'>254</a>.</li> + +<li>Glacial periods, effects of, on distribution of plants and animals, <a href='#page_209'>209</a>, <a href='#page_210'>210</a>, and <a href="#APPENDIX_AND_NOTES">Appendix</a>.</li> + +<li>Goose, Frizzled, portrait of, <a href='#page_304'>304</a>.</li> + +<li>Gorilla, <i>see</i> Apes.</li> + +<li>Gray, Professor Asa, <a href='#page_337'>337</a></li> + +<li>Great-toe, in man and apes, <a href='#page_79'>79</a>-<a href='#page_81'>81</a>.</li> + +<li>Grouse, <a href='#page_317'>317</a>-<a href='#page_319'>319</a></li> + +<li>Growth, correlation of, <a href='#page_357'>357</a>, <a href='#page_362'>362</a>.</li> + +<li><span class='sn'>Gymnotus</span>, <a href='#page_365'>365</a>, <a href='#page_367'>367</a>.</li> + +<li class='b c mt2'>H.</li> + +<li>Häckel, on analogy between species and languages, <a href='#page_32'>32</a>; +<ul class='off'><li>on reproduction as discontinuous growth, <a href='#page_105'>105</a>, <a href='#page_106'>106</a>;</li> +<li>his ideal primitive vertebrate, <a href='#page_143'>143</a>, <a href='#page_144'>144</a>.</li></ul></li> + +<li>Hair, vestigial characters of, in man, <a href='#page_89'>89</a>-<a href='#page_92'>92</a>.</li> + +<li>Hales, <a href='#page_3'>3</a>.</li> + +<li>Haller, <a href='#page_3'>3</a>.</li> + +<li>Hamilton, Sir William, <a href='#page_272'>272</a>.</li> + +<li>Hands, <a href='#page_51'>51</a>-<a href='#page_55'>55</a>, <a href='#page_66'>66</a>, <a href='#page_80'>80</a>-<a href='#page_82'>82</a>, <a href='#page_174'>174</a>-<a href='#page_192'>192</a>.</li> + +<li>Hare, <a href='#page_318'>318</a>, <a href='#page_319'>319</a>.<span class="pagenum"><a name="page_455" id="page_455">[455]</a></span></li> + +<li>Hartmann, on flattening of early human tibiæ, <a href='#page_96'>96</a>.</li> + +<li>Harvey, on Lord Bacon’s writings, <a href='#page_2'>2</a>.</li> + +<li>Heart, development of, <a href='#page_154'>154</a>.</li> + +<li>Heilprin, on skulls of deer, <a href='#page_198'>198</a>, <a href='#page_199'>199</a>; +<ul class='off'><li>on fossil shells, <a href='#page_201'>201</a>, <a href='#page_202'>202</a>.</li></ul></li> + +<li>Hen, ovum of, <a href='#page_122'>122</a>.</li> + +<li>Heredity, in relation to classification, <a href='#page_28'>28</a>-<a href='#page_31'>31</a>; +<ul class='off'><li>in relation to embryology, <a href='#page_98'>98</a>-<a href='#page_102'>102</a>;</li> +<li>chromatin-fibres in relation to, <a href='#page_134'>134</a>;</li> +<li>in relation to theories of organic evolution, <a href='#page_253'>253</a>-<a href='#page_255'>255</a>, <a href='#page_260'>260</a>-<a href='#page_264'>264</a>, <a href='#page_377'>377</a>.</li></ul></li> + +<li>Hermit-crabs, <a href='#page_62'>62</a>-<a href='#page_65'>65</a>, <a href='#page_288'>288</a>, <a href='#page_289'>289</a>.</li> + +<li><span class='sn'>Heteromera</span>, <a href='#page_233'>233</a>.</li> + +<li>Hilgendorf, on shells of <span class='sn'>Planorbis</span>, <a href='#page_201'>201</a>.</li> + +<li><span class='sn'>Hipparion</span>, <a href='#page_191'>191</a>, <a href='#page_192'>192</a>.</li> + +<li>Hippopotamus, foot of, <a href='#page_187'>187</a>.</li> + +<li>Hog, <i>see</i> Pig.</li> + +<li>Homology, <a href='#page_38'>38</a>, <a href='#page_50'>50</a>-<a href='#page_65'>65</a>, <a href='#page_176'>176</a>, <a href='#page_177'>177</a>, <a href='#page_347'>347</a>-<a href='#page_350'>350</a>, <a href='#page_357'>357</a>-<a href='#page_359'>359</a>.</li> + +<li>Homopterous insect, imitating leaf-cutting ants, <a href='#page_331'>331</a>, <a href='#page_332'>332</a>.</li> + +<li>Hooker, Sir Joseph, on flora of St. Helena, <a href='#page_234'>234</a>.</li> + +<li>Horns, <a href='#page_98'>98</a>-<a href='#page_100'>100</a>, <a href='#page_167'>167</a>. -<a href='#page_169'>169</a>.</li> + +<li>Horse, eye of, <a href='#page_75'>75</a>; +<ul class='off'><li>limb-bones of, <a href='#page_176'>176</a>, <a href='#page_177'>177</a>, <a href='#page_186'>186</a>, <a href='#page_188'>188</a>-<a href='#page_192'>192</a>;</li> +<li>teeth of, <a href='#page_189'>189</a>-<a href='#page_191'>191</a>;</li> +<li>portraits of domesticated breeds of, <a href='#page_309'>309</a>.</li></ul></li> + +<li>Human, <i>see</i> Man.</li> + +<li><span class='sn'>Humerus</span>, perforations of, in quadrumana and man, <a href='#page_94'>94</a>, <a href='#page_95'>95</a>.</li> + +<li>Humming-birds, restricted to the New World, <a href='#page_211'>211</a>.</li> + +<li>Hunter, <a href='#page_3'>3</a>; +<ul class='off'><li>on ear of whale, <a href='#page_65'>65</a>.</li></ul></li> + +<li>Huxley, Prof., on mechanical selection, <a href='#page_283'>283</a>; +<ul class='off'><li>on age of the earth, <a href='#page_435'>435</a>, <a href='#page_436'>436</a>;</li> +<li>on Dr. Whewell, <a href='#page_243'>243</a>.</li></ul></li> + +<li>Hyatt, on shells of <span class='sn'>Planorbis</span>, <a href='#page_201'>201</a>.</li> + +<li><span class='sn'>Hydra</span>, <a href='#page_111'>111</a>, <a href='#page_122'>122</a>.</li> + +<li><span class='sn'>Hyrax</span>, foot of, <a href='#page_185'>185</a>, <a href='#page_186'>186</a>.</li> + +<li class='b c mt2'>I.</li> + +<li>Ignorance, argument from, <a href='#page_41'>41</a>, <a href='#page_42'>42</a>, <a href='#page_49'>49</a>.</li> + +<li>Illative Sense, <a href='#page_6'>6</a>.</li> + +<li>Imitative colours, <a href='#page_317'>317</a>-<a href='#page_323'>323</a>, <a href='#page_326'>326</a>-<a href='#page_332'>332</a>.</li> + +<li>Infant, feet of, <a href='#page_78'>78</a>, <a href='#page_79'>79</a>; +<ul class='off'><li>grasping power of, <a href='#page_81'>81</a>.</li></ul></li> + +<li>Infertility, inter-specific, in relation to natural selection, <a href='#page_374'>374</a>-<a href='#page_376'>376</a>.</li> + +<li>Insects, wingless, <a href='#page_68'>68</a>-<a href='#page_70'>70</a>; +<ul class='off'><li>in primary formations, <a href='#page_163'>163</a>, <a href="#APPENDIX_AND_NOTES">Appendix</a>;</li> +<li>on oceanic islands, <a href='#page_224'>224</a>-<a href='#page_238'>238</a>;</li> +<li>in relation to galls, <a href='#page_293'>293</a>-<a href='#page_295'>295</a>, <a href='#page_446'>446</a>-<a href='#page_448'>448</a>;</li> +<li>defensive colouring of, <a href='#page_321'>321</a>-<a href='#page_332'>332</a>;</li> +<li>fertilizing flowers, <a href='#page_406'>406</a>.</li></ul></li> + +<li>Instincts, always of primary use to species presenting them, <a href='#page_286'>286</a>-<a href='#page_293'>293</a>.</li> + +<li>Intercrossing, in relation to natural selection, <a href='#page_374'>374</a>-<a href='#page_376'>376</a>.</li> + +<li>Inutility of specific characters, in relation to natural selection, <a href='#page_374'>374</a>-<a href='#page_376'>376</a>.</li> + +<li>Islands, oceanic, <a href='#page_224'>224</a>-<a href='#page_237'>237</a>; +<ul class='off'><li>British, <a href='#page_238'>238</a>-<a href='#page_241'>241</a>.</li></ul></li> + +<li class='b c mt2'>J.</li> + +<li>Japan, hairless dog of, <a href='#page_101'>101</a>.</li> + +<li>Jelly-fish, <a href='#page_119'>119</a>, <a href='#page_120'>120</a>.</li> + +<li class='b c mt2'>K.</li> + +<li><span class='sn'>Kallima</span>, <a href='#page_323'>323</a>.</li> + +<li>Karyokinesis, <a href='#page_112'>112</a>-<a href='#page_114'>114</a>, <a href='#page_128'>128</a>-<a href='#page_134'>134</a>.</li> + +<li>Kepler, <a href='#page_272'>272</a>.</li> + +<li>Kerguelen Island, flightless insects of, <a href='#page_70'>70</a>.</li> + +<li>Kropotkin, Prince, on co-operative instincts, <a href='#page_269'>269</a>.</li> + +<li class='b c mt2'>L.</li> + +<li><span class='sn'>Lagopus mutus</span>, <a href='#page_317'>317</a>, <a href='#page_318'>318</a>.</li> + +<li>Lamarck, his method in natural history, <a href='#page_4'>4</a>; +<ul class='off'><li>his theory of evolution, <a href='#page_253'>253</a>-<a href='#page_256'>256</a>.</li></ul></li> + +<li>Lamprey, <a href='#page_148'>148</a>.</li> + +<li>Languages, classification of, resembles that of organic forms, <a href='#page_32'>32</a>.</li> + +<li>Lankester, E. Ray, on karyokinesis, <a href='#page_129'>129</a>, <a href='#page_130'>130</a>.</li> + +<li>Leaf insect, <a href='#page_322'>322</a>.</li> + +<li>Le Conte, on geological succession of animal classes, 164, 165; +<ul class='off'><li>on types of tails, <a href='#page_169'>169</a>-<a href='#page_173'>173</a>;</li> +<li>on fossil shells of <span class='sn'>Planorbis</span>, <a href='#page_201'>201</a>;</li> +<li>his work on the relation of the theory of evolution to religious thought, <a href='#page_412'>412</a>.<span class="pagenum"><a name="page_456" id="page_456">[456]</a></span></li></ul></li> + +<li><span class='sn'>Leptalis</span>, <a href='#page_328'>328</a>.</li> + +<li><span class='sn'>Leuculmis echinus</span>, <a href='#page_122'>122</a>.</li> + +<li>Leviticus, classification of organic nature in, <a href='#page_23'>23</a>.</li> + +<li>Life, origin of, <a href='#page_15'>15</a>.</li> + +<li>Linnæus, on method in natural history, <a href='#page_3'>3</a>; +<ul class='off'><li>on classification, <a href='#page_26'>26</a>, <a href='#page_35'>35</a>-<a href='#page_40'>40</a>.</li></ul></li> + +<li>Lion, skeleton of, <a href='#page_175'>175</a>; +<ul class='off'><li>feet of, <a href='#page_178'>178</a>.</li></ul></li> + +<li>Lizard, heart and gill-arches of, <a href='#page_150'>150</a>.</li> + +<li>Lloyd Morgan, <a href='#page_273'>273</a>, <a href='#page_449'>449</a>, <a href='#page_450'>450</a>.</li> + +<li>Lungs, development of, <a href='#page_154'>154</a>, <a href='#page_354'>354</a>.</li> + +<li>Lyell, Sir Charles, on classification, <a href='#page_32'>32</a>; +<ul class='off'><li>on uniformitarianism, <a href='#page_258'>258</a>;</li> +<li>on rational species, <a href='#page_344'>344</a>;</li> +<li>on geological record, <a href='#page_422'>422</a>, <a href='#page_435'>435</a>, <a href='#page_439'>439</a>.</li></ul></li> + +<li class='b c mt2'>M.</li> + +<li>Madeira, wingless beetles of, <a href='#page_68'>68</a>-<a href='#page_70'>70</a>; +<ul class='off'><li>peculiar beetles of, <a href='#page_226'>226</a>, <a href='#page_227'>227</a>.</li></ul></li> + +<li>Mammals, ovum of, <a href='#page_120'>120</a>-<a href='#page_124'>124</a>; +<ul class='off'><li>embryology of, <a href='#page_151'>151</a>-<a href='#page_155'>155</a>;</li> +<li>palæontology of, <a href='#page_163'>163</a>, <a href='#page_165'>165</a>, <a href='#page_167'>167</a>, <a href='#page_180'>180</a>-<a href='#page_199'>199</a>;</li> +<li>limbs of, <a href='#page_174'>174</a>-<a href='#page_178'>178</a>, <a href='#page_182'>182</a>-<a href='#page_199'>199</a>;</li> +<li>brain of, <a href='#page_194'>194</a>-<a href='#page_199'>199</a>;</li> +<li>of Australia and New Zealand, <a href='#page_204'>204</a>, <a href='#page_205'>205</a>;</li> +<li>distribution of, on islands, <a href='#page_224'>224</a>-<a href='#page_240'>240</a>.</li></ul></li> + +<li>Mammoth cave, fauna of, <a href='#page_70'>70</a>-<a href='#page_72'>72</a>.</li> + +<li>Man, nictitating membrane of, <a href='#page_75'>75</a>; +<ul class='off'><li> vestigial muscles of, <a href='#page_76'>76</a>, <a href='#page_77'>77</a>, <a href='#page_82'>82</a>, <a href='#page_83'>83</a>;</li> +<li>tail of, compared with that of apes, <a href='#page_82'>82</a>-<a href='#page_84'>84</a>;</li> +<li>hair of, compared with that of apes, <a href='#page_89'>89</a>-<a href='#page_92'>92</a>;</li> +<li>teeth of, compared with those of apes, <a href='#page_92'>92</a>-<a href='#page_94'>94</a>;</li> +<li>perforation of humerus of, <a href='#page_94'>94</a>, <a href='#page_95'>95</a>;</li> +<li>flattening of ancient tibiæ of, <a href='#page_95'>95</a>, <a href='#page_96'>96</a>;</li> +<li>embryology of, <a href='#page_119'>119</a>, <a href='#page_153'>153</a>;</li> +<li>hand of, <a href='#page_54'>54</a>;</li> +<li>arm of, <a href='#page_90'>90</a>, <a href='#page_91'>91</a>;</li> +<li>limb-bones of, <a href='#page_176'>176</a>, <a href='#page_177'>177</a>;</li> +<li>palæontology of, <a href='#page_163'>163</a>, <a href='#page_165'>165</a>;</li> +<li>brain of, <a href='#page_194'>194</a>, <a href='#page_195'>195</a>;</li> +<li>Mr. Syme on, <a href='#page_346'>346</a>, <a href='#page_347'>347</a>.</li></ul></li> + +<li>Marsh, on palæontology of the horse, <a href='#page_188'>188</a>-<a href='#page_190'>190</a>.</li> + +<li>Matthew, Patrick, on natural selection, <a href='#page_257'>257</a>.</li> + +<li>Mesoderm, <a href='#page_142'>142</a>.</li> + +<li><span class='sn'>Mesohippus</span>, <a href='#page_189'>189</a>, <a href='#page_192'>192</a>.</li> + +<li>Metaphyta, <a href='#page_104'>104</a>, <a href='#page_105'>105</a>.</li> + +<li>Metazoa, <a href='#page_104'>104</a>.</li> + +<li>Method, ideas of, in natural history, <a href='#page_1'>1</a>-<a href='#page_9'>9</a>; +<ul class='off'><li>of organic evolution, <a href='#page_252'>252</a>-<a href='#page_261'>261</a>.</li></ul></li> + +<li>Meyer, Professor Ludwig, on helix of the human ear, <a href='#page_86'>86</a>.</li> + +<li>Mimicry, <a href='#page_320'>320</a>-<a href='#page_322'>322</a>.</li> + +<li>Ministration, mutual, of species alleged, <a href='#page_445'>445</a>, <a href='#page_446'>446</a>.</li> + +<li><span class='sn'>Miohippus</span>, <a href='#page_189'>189</a>.</li> + +<li>Mivart, St. George, on eye of octopus, <a href='#page_57'>57</a>, <a href='#page_58'>58</a>, <a href='#page_348'>348</a>, <a href='#page_349'>349</a>; +<ul class='off'><li>on incipient organs, <a href='#page_362'>362</a>;</li> +<li>on mutual ministration of species, <a href='#page_445'>445</a>, <a href='#page_446'>446</a>.</li></ul></li> + +<li>Mollusca, shells of, <a href='#page_19'>19</a>, <a href='#page_199'>199</a>-<a href='#page_203'>203</a>; +<ul class='off'><li>eye of, <a href='#page_57'>57</a>, <a href='#page_58'>58</a>;</li> +<li>embryology of, <a href='#page_155'>155</a>;</li> +<li>palæontology of, <a href='#page_163'>163</a>, <a href='#page_165'>165</a>.</li></ul></li> + +<li>Monkeys, why all, do not become men, <a href='#page_342'>342</a>-<a href='#page_344'>344</a>.</li> + +<li><span class='sn'>Monotremata</span>, <a href='#page_205'>205</a>.</li> + +<li>Morgan, <i>see</i> Lloyd Morgan.</li> + +<li>Morphology, <a href='#page_50'>50</a>-<a href='#page_97'>97</a>.</li> + +<li>Mule, portrait of, <a href='#page_309'>309</a>.</li> + +<li>Multicellular organisms, <a href='#page_104'>104</a>.</li> + +<li>Multiplication, <i>see</i> Reproduction.</li> + +<li class='b c mt2'>N.</li> + +<li>Nägeli, Prof., <a href='#page_337'>337</a>, <a href='#page_367'>367</a>.</li> + +<li>Natural History, ideas of method in, <a href='#page_1'>1</a>-<a href='#page_9'>9</a>.</li> + +<li>Natural, interpretations as opposed to super-natural, <a href='#page_13'>13</a>-<a href='#page_15'>15</a>; +<ul class='off'><li>causation, <a href='#page_13'>13</a>-<a href='#page_15'>15</a>.</li></ul></li> + +<li>Natural selection, <a href='#page_252'>252</a>-<a href='#page_378'>378</a>, <a href='#page_401'>401</a>-<a href='#page_410'>410</a>; +<ul class='off'><li>Wells, Matthew, and Whewell on, <a href='#page_257'>257</a>, <a href='#page_258'>258</a>, <a href='#page_443'>443</a>-<a href='#page_445'>445</a>;</li> +<li>statement of theory of, <a href='#page_256'>256</a>-<a href='#page_284'>284</a>; +<ul class='off'><li>of evidences of, <a href='#page_285'>285</a>-<a href='#page_332'>332</a>;</li> +<li>of criticisms of, <a href='#page_333'>333</a>-<a href='#page_378'>378</a>;</li></ul></li> +<li>relation of theory of, to religious thought, <a href='#page_401'>401</a>-<a href='#page_410'>410</a>;</li> +<li>preserves types, <a href='#page_264'>264</a>-<a href='#page_267'>267</a>;</li> +<li>cessation and reversal of, <a href='#page_270'>270</a>, <a href='#page_342'>342</a>;</li> +<li>errors touching theory of, <a href='#page_270'>270</a>-<a href='#page_284'>284</a>, <a href='#page_332'>332</a>-<a href='#page_364'>364</a>;</li> +<li>definition of, <a href='#page_275'>275</a>-<a href='#page_276'>276</a>;[**]</li> +<li>antecedent standing of theory of, <a href='#page_277'>277</a>-<a href='#page_284'>284</a>;</li> +<li>Prof. Owen on, <a href='#page_333'>333</a>, <a href='#page_334'>334</a>;</li> +<li>Duke of Argyll on, <a href='#page_334'>334</a>-<a href='#page_362'>362</a>;</li> +<li>Mr. Syme on, <a href='#page_340'>340</a>, <a href='#page_341'>341</a>, <a href='#page_345'>345</a>;</li> +<li>need not always make for improvement, <a href='#page_341'>341</a>-<a href='#page_347'>347</a>;</li> +<li>homology and analogy in relation to, <a href='#page_347'>347</a>-<a href='#page_350'>350</a>;</li> +<li>often determines beauty, <a href='#page_406'>406</a>, <a href='#page_407'>407</a>;</li> +<li>in relation to the formation of galls, <a href='#page_293'>293</a>-<a href='#page_295'>295</a>. <a href='#page_446'>446</a>-<a href='#page_448'>448</a>.<span class="pagenum"><a name="page_457" id="page_457">[457]</a></span></li></ul></li> + +<li>Nature, organic, <a href='#page_17'>17</a>; +<ul class='off'><li>inorganic, <a href='#page_1'>1</a>, <a href='#page_17'>17</a>, <a href='#page_18'>18</a>.</li></ul></li> + +<li><span class='sn'>Nauplius</span>, <a href='#page_138'>138</a>.</li> + +<li>Neumayr, <a href='#page_19'>19</a>.</li> + +<li>New Zealand, fauna of, <a href='#page_68'>68</a>, <a href='#page_204'>204</a>, <a href='#page_205'>205</a>; +<ul class='off'><li>thriving of exotic species in, <a href='#page_286'>286</a>.</li></ul></li> + +<li>Newman, on the Illative Sense, <a href='#page_6'>6</a>.</li> + +<li>Newton, his idea of scientific method, <a href='#page_6'>6</a>.</li> + +<li>Nictitating membrane, <a href='#page_74'>74</a>, <a href='#page_75'>75</a>.</li> + +<li>Notochord, <a href='#page_146'>146</a>.</li> + +<li>Novum Organon, the, on scientific method, <a href='#page_2'>2</a>.</li> + +<li>Nucleus, <a href='#page_105'>105</a>, <a href='#page_112'>112</a>-<a href='#page_134'>134</a>.</li> + +<li>Nucleus-spindle, <a href='#page_129'>129</a>.</li> + +<li>Nut-hatch, Syrian, ornamented nests of, <a href='#page_381'>381</a>.</li> + +<li class='b c mt2'>O.</li> + +<li>Objective methods, <a href='#page_6'>6</a>.</li> + +<li>Oceanic islands, <i>see</i> Islands.</li> + +<li>Octopus, eye of, <a href='#page_57'>57</a>, <a href='#page_58'>58</a>, <a href='#page_348'>348</a>-<a href='#page_350'>350</a>.</li> + +<li><span class='sn'>Œdicnemus crepitans</span>, <a href='#page_320'>320</a>.</li> + +<li>Ontogeny, as recapitulation of phylogeny, <a href='#page_98'>98</a>-<a href='#page_104'>104</a>.</li> + +<li>Orang Outang, <i>see</i> Apes.</li> + +<li><span class='sn'>Oredon Culbertsoni</span>, <a href='#page_167'>167</a>.</li> + +<li>Origin of Species, the, influence exercised by, on ideas of method, <a href='#page_1'>1</a>-<a href='#page_9'>9</a></li> + +<li><span class='sn'>Orohippus</span>, <a href='#page_189'>189</a>.</li> + +<li>Otaria, eye of, <a href='#page_75'>75</a>.</li> + +<li>Ovum, <a href='#page_113'>113</a>-<a href='#page_142'>142</a>; +<ul class='off'><li>human, <a href='#page_120'>120</a>-<a href='#page_133'>133</a>;</li> +<li>amœboid movements of young, <a href='#page_121'>121</a>-<a href='#page_123'>123</a>;</li> +<li>segmentation of, <a href='#page_134'>134</a>, <a href='#page_135'>135</a>.</li></ul></li> + +<li>Owen, on ear of whale, <a href='#page_65'>65</a>; +<ul class='off'><li>on natural selection, <a href='#page_333'>333</a>, <a href='#page_334'>334</a>.</li></ul></li> + +<li>Owl, eye of, <a href='#page_75'>75</a>.</li> + +<li class='b c mt2'>P.</li> + +<li>Paddle, <i>see</i> Whale, and <span class='sn'>Baptanodon discus</span>.</li> + +<li><span class='sn'>Pagurus bernhardus</span>, <a href='#page_64'>64</a>.</li> + +<li>Pain, in relation to the theory of evolution, <a href='#page_417'>417</a>.</li> + +<li>Palæontology, <a href='#page_159'>159</a>-<a href='#page_203'>203</a>; +<ul class='off'><li>general testimony of, <a href='#page_156'>156</a>-<a href='#page_165'>165</a>;</li> +<li>testimony of, in particular cases, <a href='#page_165'>165</a>-<a href='#page_203'>203</a>;</li> +<li>consideration of objections to theory of evolution founded on grounds of, <a href='#page_156'>156</a>-<a href='#page_165'>165</a>, and <a href="#APPENDIX_AND_NOTES">Appendix</a>.</li></ul></li> + +<li><span class='sn'>Palæotherium</span>, <a href='#page_190'>190</a>, <a href='#page_191'>191</a>.</li> + +<li>Paley, on natural theology, <a href='#page_98'>98</a>, <a href='#page_412'>412</a>.</li> + +<li><span class='sn'>Paludina</span>, successive forms of, <a href='#page_19'>19</a>.</li> + +<li>Panama, Isthmus of, <a href='#page_219'>219</a>.</li> + +<li><span class='sn'>Panniculus carnosis</span>, <a href='#page_77'>77</a>.</li> + +<li><span class='sn'>Papilio merope</span>, <a href='#page_330'>330</a>.</li> + +<li>Parasites, of animals, devoid of beauty, <a href='#page_408'>408</a>.</li> + +<li>Parsimony, law of, <a href='#page_272'>272</a>.</li> + +<li>Parthenogenesis, <a href='#page_119'>119</a>.</li> + +<li>Partridges, <a href='#page_319'>319</a>.</li> + +<li>Peacock, tail of, <a href='#page_378'>378</a>; +<ul class='off'><li>courtship of, <a href='#page_383'>383</a>.</li></ul></li> + +<li>Peckham, Mr. and Mrs., on courtship of spiders, <a href='#page_388'>388</a>-<a href='#page_390'>390</a>.</li> + +<li>Perissodactyls, <a href='#page_182'>182</a>-<a href='#page_192'>192</a>.</li> + +<li><span class='sn'>Petromyzon marinus</span>, <a href='#page_148'>148</a>.</li> + +<li><span class='sn'>Phenacodus primævus</span>, <a href='#page_184'>184</a>, <a href='#page_185'>185</a>.</li> + +<li>Phylogeny, <i>see</i> Ontogeny.</li> + +<li>Physiological selection, <a href='#page_376'>376</a>.</li> + +<li>Pig, embryology of, <a href='#page_153'>153</a>; +<ul class='off'><li>feet of, <a href='#page_176'>176</a>, <a href='#page_187'>187</a>;</li> +<li>portraits of wild and domesticated, <a href='#page_312'>312</a>.</li></ul></li> + +<li>Pigeons, portraits of, <a href='#page_298'>298</a>, <a href='#page_299'>299</a>; +<ul class='off'><li>feather-footed, <a href='#page_359'>359</a>.</li></ul></li> + +<li>Pilot fish, <a href='#page_289'>289</a>.</li> + +<li><span class='sn'>Planorbis</span>, transmutations of, <a href='#page_200'>200</a>, <a href='#page_201'>201</a>.</li> + +<li>Pleasure and pain, in relation to the theory of evolution, <a href='#page_417'>417</a>.</li> + +<li><span class='sn'>Plica semilunaris</span>, <a href='#page_75'>75</a>.</li> + +<li><span class='sn'>Pliohippus</span>, <a href='#page_189'>189</a>.</li> + +<li>Polar bear, skeleton of, <a href='#page_174'>174</a>; +<ul class='off'><li>feet of, <a href='#page_178'>178</a>.</li></ul></li> + +<li>Polar bodies, <a href='#page_125'>125</a>, <a href='#page_126'>126</a>.</li> + +<li>Polar star, <a href='#page_129'>129</a>.</li> + +<li>Polyps, <a href='#page_114'>114</a>.</li> + +<li>Porpoises, <a href='#page_24'>24</a>, <a href='#page_25'>25</a>, <a href='#page_50'>50</a>.</li> + +<li>Poulton, E. B., on warning colours, <a href='#page_325'>325</a>, <a href='#page_326'>326</a>; +<ul class='off'><li>on mimicry, <a href='#page_331'>331</a>, <a href='#page_332'>332</a>;</li> +<li>sexual selection, <a href='#page_400'>400</a>, <a href='#page_401'>401</a>, <a href='#page_449'>449</a>, <a href='#page_450'>450</a>.</li></ul></li> + +<li>Poultry, portraits of, <a href='#page_300'>300</a>-<a href='#page_302'>302</a>.</li> + +<li>Pronucleus, <a href='#page_126'>126</a>-<a href='#page_128'>128</a>.</li> + +<li>Prophetic types, <a href='#page_272'>272</a>, <a href='#page_351'>351</a>-<a href='#page_362'>362</a>.</li> + +<li><span class='sn'>Prophysema primordiale</span>, <a href='#page_140'>140</a>.</li> + +<li>Protective colouring, <a href='#page_317'>317</a>-<a href='#page_323'>323</a>.</li> + +<li><span class='sn'>Protohippus</span>, <a href='#page_189'>189</a>.<span class="pagenum"><a name="page_458" id="page_458">[458]</a></span></li> + +<li>Protozoa, <a href='#page_104'>104</a>.</li> + +<li>Ptarmigan, <a href='#page_317'>317</a>, <a href='#page_318'>318</a>.</li> + +<li><span class='sn'>Pterodactyl</span>, wing of, <a href='#page_56'>56</a>.</li> + +<li>Purpose, Darwin’s use of the word, <a href='#page_281'>281</a>, <a href='#page_340'>340</a>.</li> + +<li>Puss moth, larva of, <a href='#page_325'>325</a>, <a href='#page_326'>326</a>.</li> + +<li>Python, <a href='#page_66'>66</a>, <a href='#page_67'>67</a>.</li> + +<li class='b c mt2'>Q.</li> + +<li>Quadrumana, muscles of, <a href='#page_76'>76</a>, <a href='#page_82'>82</a>, <a href='#page_83'>83</a>; +<ul class='off'><li>perforations of humeri of, <a href='#page_94'>94</a>, <a href='#page_95'>95</a>;</li> +<li>hair on phalanges of, <a href='#page_91'>91</a>.</li></ul></li> + +<li class='b c mt2'>R.</li> + +<li>Rabbit, embryology of, <a href='#page_153'>153</a>; +<ul class='off'><li>multiplication of, in Australia, <a href='#page_286'>286</a>;</li> +<li>portraits of wild and domesticated breeds of, <a href='#page_308'>308</a>;</li> +<li>protective colouring of, <a href='#page_319'>319</a>, <a href='#page_320'>320</a>.</li></ul></li> + +<li>Radiate form, beauty of, <a href='#page_408'>408</a>, <a href='#page_409'>409</a>.</li> + +<li><span class='sn'>Raia radiata</span>, and <span class='sn'>batis</span>, <a href='#page_367'>367</a>-<a href='#page_371'>371</a>.</li> + +<li>Rats, species of, restricted to Old and New Worlds, <a href='#page_212'>212</a>; +<ul class='off'><li>British and Norwegian, <a href='#page_285'>285</a>, <a href='#page_286'>286</a>.</li></ul></li> + +<li>Rattle-snake, tail of, <a href='#page_289'>289</a>.</li> + +<li>Recognition marks, <a href='#page_271'>271</a>-<a href='#page_273'>273</a>.</li> + +<li>Religion, in relation to Darwinism, <a href='#page_401'>401</a>-<a href='#page_418'>418</a>.</li> + +<li>Reproduction, different methods of, <a href='#page_106'>106</a>-<a href='#page_117'>117</a>; +<ul class='off'><li>essence of sexual, <a href='#page_110'>110</a>;</li> +<li>foreshadowing of sexual in unicellular organisms, <a href='#page_115'>115</a>-<a href='#page_117'>117</a>.</li></ul></li> + +<li>Reptiles, wing of flying, <a href='#page_56'>56</a>; +<ul class='off'><li>rudimentary limbs of, <a href='#page_67'>67</a>;</li> +<li>nictitating membrane of, <a href='#page_75'>75</a>;</li> +<li>branchial arches of, <a href='#page_150'>150</a>;</li> +<li>embryology of, <a href='#page_152'>152</a>;</li> +<li>palæontology of, <a href='#page_163'>163</a>, <a href='#page_165'>165</a>, <a href='#page_178'>178</a>-<a href='#page_180'>180</a>;</li> +<li>brain of, <a href='#page_194'>194</a>-<a href='#page_197'>197</a>;</li> +<li>distribution of, <a href='#page_224'>224</a>-<a href='#page_240'>240</a>.</li></ul></li> + +<li><span class='sn'>Rhinoceros</span>, foot of, <a href='#page_186'>186</a>.</li> + +<li>Robinson, Dr L., on grasping power of an infant’s hands, <a href='#page_80'>80</a>-<a href='#page_82'>82</a>.</li> + +<li>Rudimentary organs, <a href='#page_65'>65</a>-<a href='#page_97'>97</a>.</li> + +<li>Ruminants, palæontology of, <a href='#page_167'>167</a>, <a href='#page_168'>168</a>.</li> + +<li class='b c mt2'>S.</li> + +<li>Sacrum of man, compared with that of apes, <a href='#page_82'>82-</a><a href='#page_84'>84</a>.</li> + +<li><span class='sn'>Sagitta</span>, <a href='#page_138'>138</a>.</li> + +<li>Salamander, young of terrestrial, living in water, <a href='#page_102'>102</a>; +<ul class='off'><li>embryology of, <a href='#page_152'>152</a>.</li></ul></li> + +<li>Sandwich Islands, <a href='#page_234'>234</a>-<a href='#page_237'>237</a>.</li> + +<li>Science, method of, <a href='#page_1'>1</a>-<a href='#page_9'>9</a>.</li> + +<li>Sclater, W. L., on a case of mimicry, <a href='#page_331'>331</a>, <a href='#page_332'>332</a>.</li> + +<li>Scorpion in Silurian formation, <a href='#page_163'>163</a>.</li> + +<li>Sea, lamprey, <a href='#page_148'>148</a>; +<ul class='off'><li>destructive agency of the, <a href='#page_423'>423</a>, <a href='#page_424'>424</a>.</li></ul></li> + +<li>Seal, <a href='#page_51'>51</a>, <a href='#page_52'>52</a>, <a href='#page_75'>75</a>.</li> + +<li>Seasonal changes of colour, <a href='#page_317'>317</a>-<a href='#page_319'>319</a>.</li> + +<li>Selection, value, <a href='#page_275'>275</a>; +<ul class='off'><li>by physical processes, <a href='#page_282'>282</a>, <a href='#page_283'>283</a>, <a href='#page_335'>335</a>. <i>See also</i> Natural selection, Artificial selection, Sexual selection, Physiological selection.</li></ul></li> + +<li>Sentiency, in relation to the theory of evolution, <a href='#page_417'>417</a>.</li> + +<li>Sex, difference of, restricted to Metazoa and Metaphyta, <a href='#page_105'>105</a>.</li> + +<li>Sexual reproduction, <i>see</i> Reproduction.</li> + +<li>Sexual selection, theory of, <a href='#page_277'>277</a>, <a href='#page_378'>378</a>-<a href='#page_410'>410</a>; +<ul class='off'><li>statement and evidences of, <a href='#page_379'>379</a>-<a href='#page_391'>391</a>;</li> +<li>criticisms of, <a href='#page_391'>391</a>-<a href='#page_400'>400</a>;</li> +<li>includes law of battle with that of charming, <a href='#page_385'>385</a>, <a href='#page_386'>386</a>;</li> +<li>in relation to religious thought, <a href='#page_411'>411</a>-<a href='#page_418'>418</a>;</li> +<li>Tylor’s theory substituted for, by Wallace, <a href='#page_449'>449</a>, <a href='#page_450'>450</a>.</li></ul></li> + +<li>Shark, eye of, <a href='#page_75'>75</a>; +<ul class='off'><li>man-eating, <a href='#page_149'>149</a>;</li> +<li>and pilot-fish, <a href='#page_289'>289</a>.</li></ul></li> + +<li>Sheep, limb-bones of, <a href='#page_176'>176</a>, <a href='#page_177'>177</a>; +<ul class='off'><li>portraits of, <a href='#page_310'>310</a>.</li></ul></li> + +<li>Shells, of crabs, <a href='#page_62'>62</a>-<a href='#page_64'>64</a>; +<ul class='off'><li>palæontology of mollusks, <a href='#page_199'>199</a>-<a href='#page_203'>203</a>;</li> +<li>land on oceanic islands, <a href='#page_224'>224</a>-<a href='#page_240'>240</a>.</li></ul></li> + +<li>Silliman’s Journal, on fauna of the Mammoth Cave, <a href='#page_70'>70</a>.</li> + +<li>Skate, electric organ of, <a href='#page_364'>364</a>-<a href='#page_373'>373</a>.</li> + +<li>Skull, palæontology of, <a href='#page_194'>194</a>-<a href='#page_199'>199</a>; +<ul class='off'><li>of bull-dog compared with that of deer-hound, <a href='#page_307'>307</a>.</li></ul></li> + +<li>Slavonia, Tertiary deposits of, <a href='#page_18'>18</a>, <a href='#page_19'>19</a>.</li> + +<li>Species, not eternal, but either created or evolved, <a href='#page_13'>13</a>; +<ul class='off'><li>named as such through absence of intermediate forms, <a href='#page_18'>18</a>-<a href='#page_20'>20</a>;<span class="pagenum"><a name="page_459" id="page_459">[459]</a></span></li> +<li>groups of, in classification, <a href='#page_20'>20</a>, +<ul class='off'><li>and appearing suddenly in geological formations, <a href='#page_427'>427</a>-<a href='#page_432'>432</a>, <a href='#page_437'>437</a>-<a href='#page_440'>440</a>;</li></ul></li> +<li>origin of, coincide in space and time with pre-existing and allied species, <a href='#page_22'>22</a>;</li> +<li>geographical distribution of, <a href='#page_204'>204</a>-<a href='#page_248'>248</a>;</li> +<li>extinct and living allied on same areas, <a href='#page_213'>213</a>;</li> +<li>life of, preserved by natural selection, <a href='#page_264'>264</a>-<a href='#page_270'>270</a>;</li> +<li>not room for more than one rational, <a href='#page_344'>344</a>;</li> +<li>characters of, <a href='#page_274'>274</a>-<a href='#page_276'>276</a>, <a href='#page_286'>286</a>-<a href='#page_295'>295</a>, <a href='#page_374'>374</a>-<a href='#page_376'>376</a>;</li> +<li>inter-sterility of allied, <a href='#page_374'>374</a>-<a href='#page_376'>376</a>;</li> +<li>mutual ministration of alleged, <a href='#page_445'>445</a>, <a href='#page_446'>446</a>.</li></ul></li> + +<li>Specific characters, <i>see</i> Characters.</li> + +<li>Speculation, method of, <a href='#page_3'>3</a>-<a href='#page_9'>9</a>.</li> + +<li>Spencer, Herbert, on reproduction as discontinuous growth, <a href='#page_105'>105</a>, <a href='#page_106'>106</a>; +<ul class='off'><li>on use-inheritance, <a href='#page_253'>253</a>-<a href='#page_256'>256</a>;</li> +<li>his failure to conceive the idea of natural selection, <a href='#page_257'>257</a>.</li></ul></li> + +<li>Spermatozoa, <a href='#page_123'>123</a>, <a href='#page_126'>126</a>, <a href='#page_128'>128</a>.</li> + +<li>Spiders, in primary formations, <a href='#page_163'>163</a>; +<ul class='off'><li>courtship of, <a href='#page_388'>388</a>, <a href='#page_389'>389</a>.</li></ul></li> + +<li>Sponges, <a href='#page_122'>122</a>, <a href='#page_139'>139</a>, <a href='#page_140'>140</a>.</li> + +<li>Spontaneous, Darwin’s use of the term, <a href='#page_340'>340</a>.</li> + +<li>Spores, <a href='#page_115'>115</a>.</li> + +<li>Squirrels, flying, <a href='#page_355'>355</a>.</li> + +<li>Sterility, <i>see</i> Infertility.</li> + +<li>St. Helena, <a href='#page_231'>231</a>-<a href='#page_234'>234</a>, <a href='#page_236'>236</a>-<a href='#page_237'>237</a>.</li> + +<li>St. Hilaire, <a href='#page_4'>4</a>.</li> + +<li>Stick-insect, <a href='#page_322'>322</a>.</li> + +<li>Stoat, <a href='#page_318'>318</a>.</li> + +<li><span class='sn'>Strombus accipilrinus</span>, <a href='#page_201'>201</a>.</li> + +<li><span class='sn'>Strombus Leidy</span>, <a href='#page_201'>201</a>.</li> + +<li>Struggle for existence, <a href='#page_259'>259</a>-<a href='#page_270'>270</a>.</li> + +<li>Subjective, methods, <a href='#page_6'>6</a>.</li> + +<li>Survival of the fittest, <a href='#page_335'>335</a>. <i>See also</i> Natural selection.</li> + +<li>Swim-bladder of fish, <a href='#page_154'>154</a>, <a href='#page_354'>354</a>.</li> + +<li>Symbiosis, <a href='#page_269'>269</a>.</li> + +<li>Syme, David, on the theory of natural selection, <a href='#page_340'>340</a>, <a href='#page_341'>341</a>.</li> + +<li class='b c mt2'>T.</li> + +<li>Tail, types of, in fish and birds, <a href='#page_169'>169</a>-<a href='#page_173'>173</a>.</li> + +<li>Tasmanian wolf, dentition of, <a href='#page_39'>39</a>.</li> + +<li>Teeth, of Tasmanian wolf, <a href='#page_39'>39</a>; +<ul class='off'><li>molar, of man, compared with those of apes, <a href='#page_92'>92</a>-<a href='#page_94'>94</a>;</li> +<li>palæontology of horses’, <a href='#page_189'>189</a>-<a href='#page_191'>191</a>.</li></ul></li> + +<li>Temperature, sense of, probable origin of that of sight, <a href='#page_353'>353</a>, <a href='#page_354'>354</a>.</li> + +<li>Tennyson, <a href='#page_266'>266</a>.</li> + +<li>Tibiæ, flattening of, <a href='#page_95'>95</a>, <a href='#page_96'>96</a>.</li> + +<li>Tissue-cells, <i>see</i> Cell.</li> + +<li>Toes, <a href='#page_79'>79</a>, <a href='#page_80'>80</a>; <i>see also</i> Feet.</li> + +<li>Tomes, C. S., on molar teeth of man and apes, <a href='#page_94'>94</a>.</li> + +<li><span class='sn'>Torpedo</span>, <a href='#page_365'>365</a>, <a href='#page_367'>367</a>.</li> + +<li>Tortoise, embryology of, <a href='#page_152'>152</a>, <a href='#page_154'>154</a>.</li> + +<li><span class='sn'>Toxopneustes variegatus</span>, and <span class='sn'>T. lividus</span>, <a href='#page_122'>122</a>.</li> + +<li>Transport of organisms, means of, <a href='#page_207'>207</a>, <a href='#page_216'>216</a>-<a href='#page_218'>218</a>.</li> + +<li>Tribal fitness, as distinguished from individual, <a href='#page_267'>267</a>-<a href='#page_269'>269</a>.</li> + +<li>Trout, ovum of, <a href='#page_122'>122</a>.</li> + +<li>Turtle, eye of, <a href='#page_75'>75</a>.</li> + +<li>Tylor, Alfred, on colouration of animals, <a href='#page_448'>448</a>-<a href='#page_450'>450</a>.</li> + +<li>Type, preserved by natural selection, <a href='#page_264'>264</a>-<a href='#page_269'>269</a>; +<ul class='off'><li>improvement of, by natural selection, <a href='#page_269'>269</a>, <a href='#page_270'>270</a>;</li> +<li>prophetic, <a href='#page_272'>272</a>, <a href='#page_351'>351</a>-<a href='#page_362'>362</a>.</li></ul></li> + +<li>Types, as simple and generalized, <a href='#page_33'>33</a>.</li> + +<li class='b c mt2'>U.</li> + +<li>Unicellular organisms, <a href='#page_104'>104</a>.</li> + +<li><span class='sn'>Uraster</span>, <a href='#page_138'>138</a>.</li> + +<li>Utility, of specific characters, <a href='#page_274'>274</a>, <a href='#page_275'>275</a>; +<ul class='off'><li>of incipient characters, <a href='#page_351'>351</a>-<a href='#page_363'>363</a>;</li> +<li>of electric organs, <a href='#page_365'>365</a>-<a href='#page_373'>373</a>.</li></ul></li> + +<li class='b c mt2'>V.</li> + +<li>Variation, in relation to natural selection, <a href='#page_263'>263</a>, <a href='#page_335'>335</a>-<a href='#page_340'>340</a>, <a href='#page_377'>377</a>.</li> + +<li>Verification, <a href='#page_6'>6</a>-<a href='#page_9'>9</a>.</li> + +<li>Vertebral column, embryology of <a href='#page_145'>145</a>, <a href='#page_146'>146</a>; +<ul class='off'><li>palæontology of, <a href='#page_192'>192</a>, <a href='#page_193'>193</a>.</li></ul></li> + +<li>Vertebrated animal, ideal primitive, <a href='#page_143'>143</a>, <a href='#page_144'>144</a>; +<ul class='off'><li>embryology of, <a href='#page_143'>143</a>, <a href='#page_155'>155</a>.</li></ul></li> + +<li><span class='sn'>Vespa vulgaris</span>, <a href='#page_331'>331</a>.</li> + +<li>Vestigial organs, <a href='#page_65'>65</a>-<a href='#page_97'>97</a>.</li> + +<li><span class='sn'>Volucella inans</span>, and <span class='sn'>V. bombylans</span>, <a href='#page_329'>329</a>.<span class="pagenum"><a name="page_460" id="page_460">[460]</a></span></li> + +<li class='b c mt2'>W.</li> + +<li>Wagner, Moritz, on geographical distribution, <a href='#page_216'>216</a>.</li> + +<li>Wallace, A. R., on origin of species as coincident in time and space with pre-existing and allied species, <a href='#page_22'>22</a>; +<ul class='off'><li>on wingless insects, <a href='#page_70'>70</a>;</li> +<li>on absence of hair from human back, and function of on arms of orang, <a href='#page_89'>89</a>;</li> +<li>on geographical distribution, <a href='#page_207'>207</a>, <a href='#page_231'>231</a>, <a href='#page_232'>232</a>, <a href='#page_233'>233</a>, <a href='#page_243'>243</a>;</li> +<li>on natural selection, <a href='#page_256'>256</a>;</li> +<li>on recognition marks, <a href='#page_271'>271</a>-<a href='#page_273'>273</a>;</li> +<li>on alleged deductive consequences of the natural selection theory, <a href='#page_273'>273</a>-<a href='#page_276'>276</a>;</li> +<li>his theory of warning colours, <a href='#page_323'>323</a>, <a href='#page_324'>324</a>;</li> +<li>on sexual selection, <a href='#page_391'>391</a>-<a href='#page_400'>400</a>, <a href='#page_450'>450</a>;</li> +<li>his principal defect in treating of animal colouration, <a href='#page_449'>449</a>, <a href='#page_450'>450</a>.</li></ul></li> + +<li>Warning colours, <a href='#page_323'>323</a>-<a href='#page_326'>326</a>.</li> + +<li>Wasp, imitated by a fly, <a href='#page_329'>329</a>.</li> + +<li>Water-cress, multiplication of, in New Zealand, <a href='#page_286'>286</a>.</li> + +<li>Weevils, on St. Helena, <a href='#page_232'>232</a>.</li> + +<li>Weismann, his theory of heredity, <a href='#page_130'>130</a>, <a href='#page_134'>134</a>.</li> + +<li>Wells, Dr., on natural selection, <a href='#page_257'>257</a>.</li> + +<li>Wetterhan, Prof., on vegetable galls, <a href='#page_448'>448</a>.</li> + +<li>Whales, <a href='#page_38'>38</a>, <a href='#page_50'>50</a>, <a href='#page_53'>53</a>, <a href='#page_54'>54</a>, <a href='#page_65'>65</a>, <a href='#page_180'>180</a>.</li> + +<li>Whewell, on natural selection, <a href='#page_257'>257</a>, <a href='#page_258'>258</a>, <a href='#page_443'>443</a>-<a href='#page_445'>445</a>.</li> + +<li>Wings, <a href='#page_54'>54</a>-<a href='#page_56'>56</a>, <a href='#page_60'>60</a>, <a href='#page_61'>61</a>, <a href='#page_68'>68</a>-<a href='#page_70'>70</a>, <a href='#page_355'>355</a>.</li> + +<li>Wolf, Tasmanian, dentition of, <a href='#page_34'>34</a>.</li> + +<li>Wood, John, on vestigial muscles in man, <a href='#page_77'>77</a>.</li> + +<li>Woodward, on fossil cirripedes, <a href='#page_431'>431</a>.</li> + +<li>Woolner, on the human ear, <a href='#page_86'>86</a>.</li> + +<li>Worms, embryology of, <a href='#page_155'>155</a>.</li> + +<li>Wyman, Prof., on the great toe of human embryo, <a href='#page_79'>79</a>, <a href='#page_80'>80</a>.</li> + +<li class='b c mt2'>Z.</li> + +<li><span class='sn'>Zona pellucida</span>, <a href='#page_121'>121</a>.</li></ul> + +<hr /><div class='footnotes'><h2>FOOTNOTES:</h2> + +<div class="footnote"><p><a name="Footnote_1_1" id="Footnote_1_1"></a><a href="#FNanchor_1_1"><span class="label">[1]</span></a> +<i>Origin of Species</i>, p. 367.</p></div> + +<div class="footnote"><p><a name="Footnote_2_2" id="Footnote_2_2"></a><a href="#FNanchor_2_2"><span class="label">[2]</span></a> +<i>Origin of Species</i>, p. 372.</p></div> + +<div class="footnote"><p><a name="Footnote_3_3" id="Footnote_3_3"></a><a href="#FNanchor_3_3"><span class="label">[3]</span></a> +<i>Origin of Species</i>, pp. 368-9.</p></div> + +<div class="footnote"><p><a name="Footnote_4_4" id="Footnote_4_4"></a><a href="#FNanchor_4_4"><span class="label">[4]</span></a> +It is, however, probable that all species of the genus retained a tiny +rudiment of wings in greatly dwindled scapulo-coracoid bones. And Mr. H. +O. Forbes has detected, in a recently exhumed specimen of the latter, an +indication of the glenoid cavity, for the articulation of an extremely +aborted humerus. (See <i>Nature</i>, Jan. 14th, 1892.)</p></div> + +<div class="footnote"><p><a name="Footnote_5_5" id="Footnote_5_5"></a><a href="#FNanchor_5_5"><span class="label">[5]</span></a> +See especially Mr. John Wood’s papers, <i>Proc. R. S.</i>, xiii to xvi, +and xviii; also <i>Journ. Anat.</i>, i and iii. In this connexion Darwin +refers to M. Richard, <i>Annls. d. Sc. Nat. Zoolg.</i>, tom. xviii, p. 13, +1852.</p></div> + +<div class="footnote"><p><a name="Footnote_6_6" id="Footnote_6_6"></a><a href="#FNanchor_6_6"><span class="label">[6]</span></a> +<i>Proc. Nat. Hist. Soc.</i>, Boston, 1863.</p></div> + +<div class="footnote"><p><a name="Footnote_7_7" id="Footnote_7_7"></a><a href="#FNanchor_7_7"><span class="label">[7]</span></a> +<i>Nineteenth Century</i>, November, 1891.</p></div> + +<div class="footnote"><p><a name="Footnote_8_8" id="Footnote_8_8"></a><a href="#FNanchor_8_8"><span class="label">[8]</span></a> +<i>Descent of Man</i>, 2nd ed., pp. 15-16.</p></div> + +<div class="footnote"><p><a name="Footnote_9_9" id="Footnote_9_9"></a><a href="#FNanchor_9_9"><span class="label">[9]</span></a> +I say “probably,” because analogy points in this direction. +As a matter of fact, in many cases of tissue-formation karyokinesis has +not hitherto been detected. But even if in such cases it does not +occur—i. e. if failure to detect its occurrence be not due merely +to still remaining imperfections of our histological methods,—the +large number of cases in which it has been seen to occur in the +formation of sundry tissues are of themselves sufficient to indicate +some important difference between cells derived from ova (metazoal), and +cells which have not been so derived (protozoal). Which is the point now +under discussion.</p></div> + +<div class="footnote"><p><a name="Footnote_10_10" id="Footnote_10_10"></a><a href="#FNanchor_10_10"><span class="label">[10]</span></a> +Even when propagated by budding, a multicellular organism has been +ultimately derived from a germ-cell.</p></div> + +<div class="footnote"><p><a name="Footnote_11_11" id="Footnote_11_11"></a><a href="#FNanchor_11_11"><span class="label">[11]</span></a> +It has already been stated that both parthenogenesis and gemmation are +ultimately derived from sexual reproduction. It may now be added, on the +other hand, that the earlier stages of parthenogenesis have been +observed to occur sporadically in all sub-kingdoms of the Metaxoa, +including the Vertebrata, and even the highest class, Mammalia. These +earlier stages consist in <i>spontaneous</i> segmentations of the ovum; so +that even if a virgin has ever conceived and borne a son, and even if +such a fact in the human species has been unique, still it would not +betoken any breach of physiological continuity. Indeed, according to +Weismann’s not improbable hypothesis touching the physiological +meaning of polar bodies, such a fact need betoken nothing more than a +slight disturbance of the complex machinery of ovulation, on account of +which the ovum failed to eliminate from its substance an almost +inconceivably minute portion of its nucleus.</p></div> + +<div class="footnote"><p><a name="Footnote_12_12" id="Footnote_12_12"></a><a href="#FNanchor_12_12"><span class="label">[12]</span></a> +The spermatozooids of certain plants can be strongly attracted towards a +pipette which is filled with malic acid—crowding around and into +it with avidity.</p></div> + +<div class="footnote"><p><a name="Footnote_13_13" id="Footnote_13_13"></a><a href="#FNanchor_13_13"><span class="label">[13]</span></a> +Ray Lankester, <i>Encyclop. Brit.</i>, 9th ed., Vol. XIX, pp. 832-3.</p></div> + +<div class="footnote"><p><a name="Footnote_14_14" id="Footnote_14_14"></a><a href="#FNanchor_14_14"><span class="label">[14]</span></a> +In most vertebrated animals this process of gastrulation has been more +or less superseded by another, which is called delamination; but it +scarcely seems necessary for our present purposes to describe the +latter. For not only does it eventually lead to the same result as +gastrulation—i. e. the converting of the ovum into a double-walled +sac,—but there is good evidence among the lower Vertebrata of its +being preceded by gastrulation; so that, even as to the higher +Vertebrata, embryologists are pretty well agreed that delamination has +been but a later development of, or possibly improvement upon, +gastrulation.</p></div> + +<div class="footnote"><p><a name="Footnote_15_15" id="Footnote_15_15"></a><a href="#FNanchor_15_15"><span class="label">[15]</span></a> +The most extreme of them is that which is mentioned in the last +foot-note.</p></div> + +<div class="footnote"><p><a name="Footnote_16_16" id="Footnote_16_16"></a><a href="#FNanchor_16_16"><span class="label">[16]</span></a> +For objections which may be brought against this and similar statements, +see the <a href="#APPENDIX_AND_NOTES">Appendix</a>.</p></div> + +<div class="footnote"><p><a name="Footnote_17_17" id="Footnote_17_17"></a><a href="#FNanchor_17_17"><span class="label">[17]</span></a> +For difficulties and objections, see Appendix.</p></div> + +<div class="footnote"><p><a name="Footnote_18_18" id="Footnote_18_18"></a><a href="#FNanchor_18_18"><span class="label">[18]</span></a> +Heilprin, <i>Geological Evidences of Evolution</i>, pp. 73-4 (1888).</p></div> + +<div class="footnote"><p><a name="Footnote_19_19" id="Footnote_19_19"></a><a href="#FNanchor_19_19"><span class="label">[19]</span></a> +Le Conte, <span class='sn'>loc. cit.</span>, pp. 236-7.</p></div> + +<div class="footnote"><p><a name="Footnote_20_20" id="Footnote_20_20"></a><a href="#FNanchor_20_20"><span class="label">[20]</span></a> +I say “<i>large</i> areas” for the sake of argument; but the same +correlation between distribution and affinity extends likewise to +<i>small</i> areas where only <i>small</i> differences of affinity are concerned. +Thus, for instance, speaking of smaller areas, Moritz Wagner +says:—"The broader and more rapid the river, the higher and more +regular the mountain-chain, the calmer and more extensive the sea, the +more considerable, as a general rule, will be the taxonomic separation +between the populations"; and he shows that, in correlation with such +differences in the <i>degrees</i> of separation, are the <i>degrees</i> of +diversification—i. e., the <i>numbers</i> of species, and even of +varieties, which these topographical barriers determine.</p></div> + +<div class="footnote"><p><a name="Footnote_21_21" id="Footnote_21_21"></a><a href="#FNanchor_21_21"><span class="label">[21]</span></a> +The only exception is in the case of the fish on each side of the +Isthmus of Panama, where about 30 per cent, of the species are +identical. But it is possible enough that at some previous time this +narrow Isthmus may have been even narrower than at present, if not +actually open. At all events, the fact that this partial exception +occurs just where the land-barrier is so narrow, is more suggestive of +migration than of independent creation.</p></div> + +<div class="footnote"><p><a name="Footnote_22_22" id="Footnote_22_22"></a><a href="#FNanchor_22_22"><span class="label">[22]</span></a> +<i>Origin of Species</i>, pp. 353-4.</p></div> + +<div class="footnote"><p><a name="Footnote_23_23" id="Footnote_23_23"></a><a href="#FNanchor_23_23"><span class="label">[23]</span></a> +Wallace, <i>Island Life</i>, pp. 271-2.</p></div> + +<div class="footnote"><p><a name="Footnote_24_24" id="Footnote_24_24"></a><a href="#FNanchor_24_24"><span class="label">[24]</span></a> +Wallace, <i>Island Life</i>, p. 287.</p></div> + +<div class="footnote"><p><a name="Footnote_25_25" id="Footnote_25_25"></a><a href="#FNanchor_25_25"><span class="label">[25]</span></a> +Wallace, <i>Island Life</i>, p. 287.</p></div> + +<div class="footnote"><p><a name="Footnote_26_26" id="Footnote_26_26"></a><a href="#FNanchor_26_26"><span class="label">[26]</span></a> +For quotations, see Note A.</p></div> + +<div class="footnote"><p><a name="Footnote_27_27" id="Footnote_27_27"></a><a href="#FNanchor_27_27"><span class="label">[27]</span></a> +Whewell, <i>Indications of the Creator</i>, 2nd ed., 1846.</p></div> + +<div class="footnote"><p><a name="Footnote_28_28" id="Footnote_28_28"></a><a href="#FNanchor_28_28"><span class="label">[28]</span></a> +De Blainville, <i>Compte Rendu</i>, 1837.</p></div> + +<div class="footnote"><p><a name="Footnote_29_29" id="Footnote_29_29"></a><a href="#FNanchor_29_29"><span class="label">[29]</span></a> +Whewell, <i>ibid.</i>, p. 162.</p></div> + +<div class="footnote"><p><a name="Footnote_30_30" id="Footnote_30_30"></a><a href="#FNanchor_30_30"><span class="label">[30]</span></a> +For cases, see <i>Animal Intelligence</i>, in the chapters on Ants and Bees; +and, for discussion of principles, <i>Mental Evolution in Animals</i>, in the +chapters on Instinct.</p></div> + +<div class="footnote"><p><a name="Footnote_31_31" id="Footnote_31_31"></a><a href="#FNanchor_31_31"><span class="label">[31]</span></a> +Prince Kropotkin in the <i>Nineteenth Century</i> (Feb. 1888, Apr. 1891) has +adduced a large and interesting body of facts, showing the great +prevalence of the principle of co-operation in organic nature.</p></div> + +<div class="footnote"><p><a name="Footnote_32_32" id="Footnote_32_32"></a><a href="#FNanchor_32_32"><span class="label">[32]</span></a> +<i>Darwinism</i>, pp. 218 and 227.</p></div> + +<div class="footnote"><p><a name="Footnote_33_33" id="Footnote_33_33"></a><a href="#FNanchor_33_33"><span class="label">[33]</span></a> +Since the above was written Prof. Lloyd Morgan has published a closely +similar notice of the passage in question. “This language,” +he says, “seems to savour of teleology (that pitfall of the +evolutionist). The cart is put before the horse. The recognition-marks +were, I believe, not produced to prevent intercrossing, but +intercrossing has been prevented because of preferential mating between +individuals possessing special recognition-marks. To miss this point is +to miss an important segregation-factor."—(<i>Animal Life and +Intelligence</i>, p. 103.) Again, on pp. 184-9, he furnishes an excellent +discussion on the whole subject of the fallacy alluded to in the text, +and gives illustrative quotations from other prominent Darwinians. I +should like to add that Darwin himself has nowhere fallen into this, or +any of the other fallacies, which are mentioned in the text.</p></div> + +<div class="footnote"><p><a name="Footnote_34_34" id="Footnote_34_34"></a><a href="#FNanchor_34_34"><span class="label">[34]</span></a> +Of course adaptive modifications produced in the individual lifetime, +and not <i>inherited</i>, do not concern the question at all. In this and the +following paragraphs, therefore, “adaptations,” +“adaptive modifications,” &c., refer exclusively to such +as are hereditary, i. e. phyletic.</p></div> + +<div class="footnote"><p><a name="Footnote_35_35" id="Footnote_35_35"></a><a href="#FNanchor_35_35"><span class="label">[35]</span></a> +The question as to whether natural selection has been the only principle +concerned in the origination of species, is quite distinct from that as +to the accuracy of the above definition.</p></div> + +<div class="footnote"><p><a name="Footnote_36_36" id="Footnote_36_36"></a><a href="#FNanchor_36_36"><span class="label">[36]</span></a> +It is often objected to Darwin’s terminology, that it embraces +such words as “contrivance,” “purpose,” &c., +which are strictly applicable only to the processes or the products of +thought. But when it is understood that they are used in a neutral or +metaphorical sense, I cannot see that any harm arises from their use.</p></div> + +<div class="footnote"><p><a name="Footnote_37_37" id="Footnote_37_37"></a><a href="#FNanchor_37_37"><span class="label">[37]</span></a> +Note B.</p></div> + +<div class="footnote"><p><a name="Footnote_38_38" id="Footnote_38_38"></a><a href="#FNanchor_38_38"><span class="label">[38]</span></a> +Were it not that some of Darwin’s critics have overlooked the very +point wherein the great value of protective colouring as evidence of +natural selection consists, it would be needless to observe that it does +so in the <i>minuteness</i> of the protective resemblance which in so many +cases is presented. Of course where the resemblance is only very +general, the phenomena might be ascribed to mere coincidence, of which +the instincts of the animal have taken advantage. But in the measure +that the resemblance becomes minutely detailed, the supposition of mere +coincidence is excluded, and the agency of some specially adaptive cause +demonstrated. Again, it is almost needless to say, no real difficulty is +presented (as has been alleged) by the cases above quoted of seasonal +imitations, on the ground that natural selection could not act +alternately on the same individual. Natural selection is not supposed to +act alternately on the same individual. It is supposed to act always in +the same manner, and if, as in the case of a regularly recurring change +in the colours of the environment, correspondingly recurrent changes are +required to appear in the colours of the animals, natural selection sets +its premium upon those individuals the constitutions of which best lend +themselves to seasonal changes of the needful kind—probably under +the influence of stimuli supplied by the changes of external conditions +(temperature, moisture, &c.).</p></div> + +<div class="footnote"><p><a name="Footnote_39_39" id="Footnote_39_39"></a><a href="#FNanchor_39_39"><span class="label">[39]</span></a> +For a full account of this instinct and its probable purpose, see +<i>Animal Intelligence</i>, pp. 93-6.</p></div> + +<div class="footnote"><p><a name="Footnote_40_40" id="Footnote_40_40"></a><a href="#FNanchor_40_40"><span class="label">[40]</span></a> +Both drawings are reproduced from Mr. Poulton’s paper upon the +subject (<i>Proc. Zool. Soc.</i>, June 16, 1891).</p></div> + +<div class="footnote"><p><a name="Footnote_41_41" id="Footnote_41_41"></a><a href="#FNanchor_41_41"><span class="label">[41]</span></a> +<i>Anatomy of Vertebrates</i>, vol. iii. p. 794.</p></div> + +<div class="footnote"><p><a name="Footnote_42_42" id="Footnote_42_42"></a><a href="#FNanchor_42_42"><span class="label">[42]</span></a> +The degree in which variability is indefinite, or, on the contrary, +determinate, is a question which is not yet ripe for decision—nor +even, in my opinion, for <ins class="corr" title="Transcriber’s +note: The original showed ‘dicussion’.">discussion</ins>. But I +may here state the following general principles with regard to it.</p> + +<p>(1) It is evident that up to some point or another variations <i>must</i> be +pre-determined in definite lines. Men do not gather grapes from thorns, +figs from thistles, nor even moss-roses from sweet-briars. In other +words, “the nature of the organism” in all cases +necessitates the limiting of variations within certain bounds.</p> + +<p>(2) But when the question is as to what these bounds may be, we can only +answer in a general way that, according to the general theory of +evolution, they must be such as are imposed by heredity, coupled with +the degree to which external conditions of life (and possibly also +use-inheritance) are capable, in given cases, of modifying congenital +characters. These are the only causes which the theory of descent can +consistently recognise as producing variations in determinate +directions.</p> + +<p>(3) Inasmuch as variation presupposes the existence of parts that vary, +and inasmuch as the variation of parts can only be in the alternative +directions of increase or decrease around an average, it follows that, +in the first instance at all events, every variation, if determinate, +must be so only in one or other of these two opposite directions.</p> + +<p>(4) In as far as variations are summated in successive generations, so +as eventually to give rise to new structures, organs, mechanisms, +&c., natural selection is theoretically competent to explain the +facts, without our having to postulate the operation of unknown causes +producing variations in determinate lines,—or not further than is +stated in paragraphs 1 and 2.</p> + +<p>(5) Nevertheless, it does not follow that there are not such other +unknown causes; and, if there are, of course the importance of natural +selection as a cause of adaptive modification would be limited in +proportion to their number and the extent of their operation. But it is +for those who, like the late Professors Asa Gray and Nägeli, +maintain the existence of such causes, to substantiate their belief by +indicating them.</p></div> + +<div class="footnote"><p><a name="Footnote_43_43" id="Footnote_43_43"></a><a href="#FNanchor_43_43"><span class="label">[43]</span></a> +Within the last few months this objection has been presented anew by Mr. +D. Syme, whose book <i>On the Modification of Organisms</i> exhibits a +curious combination of shrewd criticisms with almost ludicrous +misunderstandings. One of the latter it is necessary to state, because +it pervades the quotation which I am about to supply. He everywhere +compares “natural selection” with “the struggle for +existence,” uses them as convertible terms, and while absurdly +stating that “Darwin defines natural selection as the struggle for +existence,” complains of “the liability of error, both on +his own part and on the part of his readers,” which arises from +his not having everywhere adhered to this definition! (p. 8).</p> + +<p>“Darwin has put forth two distinct and contradictory theories of +the functions of natural selection. According to the one theory natural +selection is selective or preservative, and nothing more. According to +the other theory natural selection creates the variations(!) ... It +certainly seems absurd to speak of natural selection, or the struggle +for existence, as selective or preservative, for the struggle for +existence does not preserve at all, not even the fit variations, as both +the fit and the unfit struggle for existence, the unfit naturally more +than the fit, and the fit are preserved, not in consequence of the +struggle, but in consequence of their fitness. Suppose two varieties of +the same species are driven, by an increase of their numbers, to seek +for subsistence in a colder region than they have been accustomed to, +and that one of these varieties had a hardier constitution than the +other; and let us suppose that the former withstood the severe climate +better than the latter, and consequently survived, while the other +perished. In this case the hardier survived, not because of the +struggle, but because it had a constitution better adapted to the +climate. I wish to ascertain if a certain metal in my possession is gold +or some baser metal, and I apply the usual test; but the mere fact of my +testing this metal would not make it gold or any other kind of +metal.”</p> + +<p> I have thought it worth while to quote this passage for the sake of +showing the extraordinary confusion of mind which still prevails on the +part of Darwin’s critics, even with reference to the very +fundamental parts of his theory. For, as I have said, the writer of this +passage shows himself a shrewd critic in some other parts of his essay, +where he is not engaged especially on the theory of natural selection.</p></div> + +<div class="footnote"><p><a name="Footnote_44_44" id="Footnote_44_44"></a><a href="#FNanchor_44_44"><span class="label">[44]</span></a> +<i>Principles of Geology</i>, vol. ii. p. 487 (11th ed.).</p></div> + +<div class="footnote"><p><a name="Footnote_45_45" id="Footnote_45_45"></a><a href="#FNanchor_45_45"><span class="label">[45]</span></a> +Syme, on the <i>Modification of Organisms</i>, p. 46.</p></div> + +<div class="footnote"><p><a name="Footnote_46_46" id="Footnote_46_46"></a><a href="#FNanchor_46_46"><span class="label">[46]</span></a> +<i>Variation of Plants and Animals</i>, vol. ii. p. 315.</p></div> + +<div class="footnote"><p><a name="Footnote_47_47" id="Footnote_47_47"></a><a href="#FNanchor_47_47"><span class="label">[47]</span></a> +The chambers are three in number. The two upper ones are occupied +respectively by the male and the sitting female. The lower one serves as +a general living room when the young are hatched.</p></div> + +<div class="footnote"><p><a name="Footnote_48_48" id="Footnote_48_48"></a><a href="#FNanchor_48_48"><span class="label">[48]</span></a> +Note C.</p></div> + +<div class="footnote"><p><a name="Footnote_49_49" id="Footnote_49_49"></a><a href="#FNanchor_49_49"><span class="label">[49]</span></a> +Since the above exposition of the theory of sexual selection was +written, Mr. Poulton has published his work on the <i>Colours of Animals</i>. +He there reproduces some of the illustrations which occur in Mr. and +Mrs. Peckham’s work on <i>Sexual Selection in Spiders</i>, and +furnishes appropriate descriptions. Therefore, while retaining the +illustrations, I have withdrawn my own descriptions.</p> + +<p> Mr. Poulton has also in his book supplied a <i>résumé</i> of +the arguments for and against the theory of sexual selection in general. +Of course in nearly all respects this corresponds with the +<i>résumé</i> which is given in the foregoing pages; but I have +left the latter as it was originally written, because all the critical +part is reproduced <i>verbatim</i> from a review of Mr. Wallace’s +<i>Darwinism</i>, of a date still earlier than that of Mr. Poulton’s +book—viz. <i>Contemporary Review</i>, August, 1889.</p></div> + +<div class="footnote"><p><a name="Footnote_50_50" id="Footnote_50_50"></a><a href="#FNanchor_50_50"><span class="label">[50]</span></a> +The beauty of autumnal tints in fading leaves may possibly be adduced +<i>per contra</i>. But here we have to remember that it is only some kinds of +leaves which thus become beautiful when fading, while, even as regards +those that do, it is not remarkable that their chlorophyll should, as it +were, accidentally assume brilliant tints while breaking down into lower +grades of chemical constitution. The case, in fact, is exactly parallel +to those in the animal kingdom which are considered in the ensuing +paragraphs.</p></div> + +<div class="footnote"><p><a name="Footnote_51_51" id="Footnote_51_51"></a><a href="#FNanchor_51_51"><span class="label">[51]</span></a> +The best treatise on this subject is Prof. Le Conte’s <i>Evolution +and its Relation to Religious Thought</i> (Appleton & Co. 1888).</p></div> + +<div class="footnote"><p><a name="Footnote_52_52" id="Footnote_52_52"></a><a href="#FNanchor_52_52"><span class="label">[52]</span></a> +See <i>Mental Evolution in Animals</i>, pp. 110-111.</p></div> + +<div class="footnote"><p><a name="Footnote_53_53" id="Footnote_53_53"></a><a href="#FNanchor_53_53"><span class="label">[53]</span></a> +<i>Origin of Species</i>, 282-5.</p></div> + +<div class="footnote"><p><a name="Footnote_54_54" id="Footnote_54_54"></a><a href="#FNanchor_54_54"><span class="label">[54]</span></a> +<i>Elements of Geology</i>, p. 587.</p></div> + +<div class="footnote"><p><a name="Footnote_55_55" id="Footnote_55_55"></a><a href="#FNanchor_55_55"><span class="label">[55]</span></a> +<i>Origin of Species</i>, p. 289.</p></div> + +<div class="footnote"><p><a name="Footnote_56_56" id="Footnote_56_56"></a><a href="#FNanchor_56_56"><span class="label">[56]</span></a> +<i>Ibid.</i></p></div> + +<div class="footnote"><p><a name="Footnote_57_57" id="Footnote_57_57"></a><a href="#FNanchor_57_57"><span class="label">[57]</span></a> +See <i>Lay Sermons</i>, Lecture on Geological Reform.</p></div> + +<div class="footnote"><p><a name="Footnote_58_58" id="Footnote_58_58"></a><a href="#FNanchor_58_58"><span class="label">[58]</span></a> +See especially the following Presidential addresses:—Geol. Assoc. +Nov. 1876; Section D. Brit. Assoc., 1886; Lin. Soc., 1890.</p></div> + +<div class="footnote"><p><a name="Footnote_59_59" id="Footnote_59_59"></a><a href="#FNanchor_59_59"><span class="label">[59]</span></a> +<i>Elements of Geology</i>, p. 280.</p></div> + +<div class="footnote"><p><a name="Footnote_60_60" id="Footnote_60_60"></a><a href="#FNanchor_60_60"><span class="label">[60]</span></a> +<i>Origin of Species</i>, p. 332.</p></div> + +<div class="footnote"><p><a name="Footnote_61_61" id="Footnote_61_61"></a><a href="#FNanchor_61_61"><span class="label">[61]</span></a> +<i>Origin of Species</i>, p. 332.</p></div> + +<div class="footnote"><p><a name="Footnote_62_62" id="Footnote_62_62"></a><a href="#FNanchor_62_62"><span class="label">[62]</span></a> +<i>Ibid</i>. pp. 333-4.</p></div> + +<div class="footnote"><p><a name="Footnote_63_63" id="Footnote_63_63"></a><a href="#FNanchor_63_63"><span class="label">[63]</span></a> +<i>Evolution and its Relation to Religious Thought</i>, p. +194.</p></div> + +<div class="footnote"><p><a name="Footnote_64_64" id="Footnote_64_64"></a><a href="#FNanchor_64_64"><span class="label">[64]</span></a> +<i>On Truth</i>, p. 493.</p></div> + +<div class="footnote"><p><a name="Footnote_65_65" id="Footnote_65_65"></a><a href="#FNanchor_65_65"><span class="label">[65]</span></a> +<i>Nature</i>, vol. xli, p. 344.</p></div> + +<div class="footnote"><p><a name="Footnote_66_66" id="Footnote_66_66"></a><a href="#FNanchor_66_66"><span class="label">[66]</span></a> +<i>Entomologist</i>, March, 1890.</p></div> + +<div class="footnote"><p><a name="Footnote_67_67" id="Footnote_67_67"></a><a href="#FNanchor_67_67"><span class="label">[67]</span></a> +<i>Nature</i>, vol. xli, p. 394.</p></div> + +<div class="footnote"><p><a name="Footnote_68_68" id="Footnote_68_68"></a><a href="#FNanchor_68_68"><span class="label">[68]</span></a> +<i>Ibid.</i> vol. xli, pp. 559-560.</p></div></div> + +<div class='bbox'> +<h3>Transcriber’s Notes and Errata</h3> + +<p>The following words were found in both hyphenated and unhyphenated forms +in the text.</p> + + +<div class='center'> +<table border="1" cellpadding="4" cellspacing="0" summary=""> +<tr class='b'><td class='c' colspan='2'>Hyphenated</td><td class='c' colspan='2'>Unhyphenated</td></tr> +<tr class='b'><td class='c'>Word</td><td class='c'>Instances</td><td class='c'>Word</td><td class='c'>Instances</td></tr> +<tr><td class='l'>deer-hound</td><td class='r'>2</td><td class='l'>deerhound</td><td class='r'>1</td></tr> +<tr><td class='l'>fresh-water</td><td class='r'>13</td><td class='l'>freshwater</td><td class='r'>1</td></tr> +<tr><td class='l'>inter-relations</td><td class='r'>1</td><td class='l'>interrelations</td><td class='r'>1</td></tr> +<tr><td class='l'>re-action</td><td class='r'>1</td><td class='l'>reaction</td><td class='r'>1</td></tr> +<tr><td class='l'>sea-weed</td><td class='r'>7</td><td class='l'>seaweed</td><td class='r'>1</td></tr> +<tr><td class='l'>super-natural</td><td class='r'>2</td><td class='l'>supernatural</td><td class='r'>24</td></tr> +<tr><td class='l'>wood-cut</td><td class='r'>3</td><td class='l'>woodcut</td><td class='r'>3</td></tr> +<tr><td class='l'>wood-cuts</td><td class='r'>4</td><td class='l'>woodcuts</td><td class='r'>1</td></tr> +</table></div> + +<p>There were 9 instances of ‘larvae’ and 3 instances of ‘larvæ’.</p> + +<p>The following typographical errors were corrected:</p> + +<div class='center'> +<table border="1" cellpadding="4" cellspacing="0" summary=""> +<tr class='b'><td class='l'>Page</td><td class='l'>Error</td><td class='l'>Correction</td></tr> +<tr><td class='l'>44</td><td class='l'>arboresent</td><td class='l'>arborescent</td></tr> +<tr><td class='l'>75</td><td class='l'>the</td><td class='l'>The</td></tr> +<tr><td class='l'>337</td><td class='l'>dicussion</td><td class='l'>discussion</td></tr> +</table></div> + +<p>In the index, the page entry for “Lyell, Sir Charles … on +geological record” was changed from ‘420’ to +‘422’.</p> + +<p>Also, the page entry for “Natural selection … defintion +of” was changed from ‘275-376’ to +‘275-276’.</p> + +<p>Some illustrations were repositioned.</p> +</div> + + + + + + + + +<pre> + + + + + +End of the Project Gutenberg EBook of Darwin, and After Darwin (Vol. 1 and +3, of 3), by George John Romanes + +*** END OF THIS PROJECT GUTENBERG EBOOK DARWIN *** + +***** This file should be named 24800-h.htm or 24800-h.zip ***** +This and all associated files of various formats will be found in: + http://www.gutenberg.org/2/4/8/0/24800/ + +Produced by Marilynda Fraser-Cunliffe, LN Yaddanapudi and +the Online Distributed Proofreading Team at +http://www.pgdp.net + + +Updated editions will replace the previous one--the old editions +will be renamed. + +Creating the works from public domain print editions means that no +one owns a United States copyright in these works, so the Foundation +(and you!) can copy and distribute it in the United States without +permission and without paying copyright royalties. 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