Wallace, Darwin, and the Theory of Natural Selection

Wallace, Darwin, and the Theory of

Natural Selection

A Study in the Development of Ideas and Attitudes

BARBARA G. BEDDALL 2502 Bronson Road, Fairfield, Connecticut

INTRODUCTION

On 1 July 1908 the L innean Society of London c o m m e m o r a t e d the reading before the Society fifty years earlier of the Darwin- Wallace joint papers , "On the Tendency of Species to fo rm Varieties; and on the Perpetuat ion of Varieties and Species by Na tu ra l Means of Selection." 1 On the first occasion only some thir ty Fellows and guests had been present at a quiet, unhera lded meet ing; the authors themselves were absent. Now there was a large and dist inguished gather ing celebrat ing the historic event. Two of the original cast were present , the nat- u_ralist Alfred Russel Wallace ( 1 8 2 3 - 1 9 1 3 ) and the botanis t Sir Joseph Dal ton Hooker (1817 -1911) . The other two, the biologist Charles Robert Darwin ( 1 8 0 9 - 1 8 8 2 ) and the geologist Sir Charles Lyell ( 1 7 9 7 - 1 8 7 5 ) had been dead for m a n y years.

Hooker, now a venerable nonagenar ian , spoke of his "half- century-old real or fancied memor ies" of that June in 1858 when his old fr iend Darwin received Wallace 's paper on na tu ra l selection. He based his account on Sir Francis Darwin 's Li fe and Let ters o f Charles Darwin , r emark ing with some uneas iness that, beyond the letters f rom Darwin to h imsel f and to Lyell, no other documen ta ry evidence existed of the events of those turbulent weeks before the reading of the papers. Despite a search, the letters to Darwin f rom Hooker and Lyell could not be found, "and, mos t surpris ing of all, Mr. Wal lace 's letter and its enclosure have disappeared." 2

1. IAnnean Society of London, The Darwin-Wallace Celebration held on 1st July, 1908, by the Linnean Society of London (London: The Society, 1908).

2. Francis Darwin, ed., The Life and Letters of Charles Darwin, in- cludin9 an Autobiographical Chapter (London: Murray, 1887; reprinted

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Hooker was troubled by the meagerness of the evidence, but Marchant, Wallace's first biographer, was unconcerned, and most people have concurred in his opinion that the eight extant letters received f rom Darwin while Wallace was in the Malay Archipelago "explain themselves and reveal the inner story of the independent discovery of the theory of Natural Selection." z But do they?

A second question pertains to the relationship between Wal- lace and Darwin. Their recollections are often taken to be accurate reflections of their earlier thoughts and actions, but details may have been altered and the emphasis changed. Both men did come to play the roles assigned to them by history, but the making of the myth, in which they both participated, has obscured some of the facts.

This study will emphasize Wallace and the influences on him. It will at tempt to disentangle the various lines of evidence, to trace Wallace's progress toward the discovery of the theory of natural selection, to throw some light on what happened both before and after June 1858, and to suggest some alterna- tives to commonly accepted theories about these events. It is based on a re-evaluation and reinterpretation of contemporary sources, both published and unpublished. Because the aim has been to concentrate on pr imary source material, exhaustive reference to every author who has written since on these subjects has not been attempted.

In particular, three of Wallace's published papers are considered in detail: "On the Law which has regulated the Intro- duction of New Species," "Note on the Theory of Permanent and Geographical Varieties," and "On the Tendency of Varieties to depart indefinitely f rom the Original Type."

A number of previously published letters bear on questions raised here. For ease of reference, information about these letters has been arranged in tabular form in the Appendix and the letters numbered consecutively. They will be referred to by numbers in brackets in the text, with fur ther discussion when required in appropriate footnotes.

Use has also been made of a manuscr ip t notebook kept by Wallace during his travels in the Malay Archipelago, and I am grateful to Mr. Thomas O'Grady and the Council of the Linnean Society of London for permission to quote f rom it.

in 2 vols., New York, Basic Books, 1959); Linnean Society of London, The Darwin-Wallace Celebration, p. 16. See also notes 115 and 139.

3. James Marchant, Alfred Russel Wallace: Letters and Reminiscences (New York: Harper, 1916), p. 105.

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W a l l a c e , D a r w i n a n d the T h e o r y of N a t u r a l Se lec t ion

I. A N I N Q U I R I N G M I N D

He who in place of reasoning, employs authority, assumes that those to whom he addresses himself are incapable of forming a judgment of their own. If they submit to this insult, may it not be presumed they acknowledge the justice of it?

Wallace, "Notebook, 1855-1859," from Jeremy Bent_ham's Book of FaUacies4

N o t u n t i l 1841, w h e n h e was e i g h t e e n y e a r s of age, d id A l f r e d Russe l W a l l a c e , t he e m i n e n t n a t u r a l i s t , b e g i n h i s so l i t a ry s t u d y of the n a t u r a l w o r l d a r o u n d h im . As a f r e q u e n t l y u n e m p l o y e d a n d a l w a y s i m p e c u n i o u s su rveyor , he t u r n e d to the s t u d y of p l a n t s to fill h i s l e i su re t i m e :

But w h a t o c c u p i e d m e chief ly a n d b e c a m e m o r e a n d m o r e the so lace a n d de l i gh t of m y lone ly r a m b l e s a m o n g the m o o r s a n d m o u n t a i n s , w a s m y f i rs t i n t r o d u c t i o n to the var i - e ty, the b e a u t y , a n d the m y s t e r y of n a t u r e as m a n i f e s t e d in t he v e g e t a b l e k ingdom.5

W a l l a c e ' s e a r l y y e a r s a n d e d u c a t i o n were qu i te u n d i s t i n - gu i shed . T h e e i g h t h ch i ld of a n i n c r e a s i n g l y i m p o v e r i s h e d f a m i l y , he was b o r n on 8 January 1823 in t he r e m o t e v i l l age of Usk in M o n m o u t h s h i r e , Wa le s . W h e n he was five, the f a m i l y m o v e d to H e r t f o r d , n e a r L o n d o n , a n d i t was a t the H e r t f o r d G r a m m a r Schoo l t h a t he r e c e i v e d h i s "very o r d i n a r y e d u c a t i o n . " Th i s e n d e d w h e n h e was a l m o s t f o u r t e e n , a n d a f t e r t h a t he was m o r e or less on h is own. Desp i t e h is c o m m o n p l a c e u p b r i n g - ing, howeve r , he h a d r e c e i v e d a p r i ce l e s s gi f t f r o m h i s f a t h e r : a love of books a n d r e a d i n g - - a key to the wor ld fo r a n y o n e who w a n t s to use it.

A f t e r a f e w m o n t h s s p e n t w i t h h is b r o t h e r J o h n in L o n d o n i n t he s p r i n g of 1837, W a l l a c e j o i n e d h is o ldes t b r o t h e r , Wil - l i a m , to l e a r n l a n d su rvey ing . But these we re l e a n y e a r s fo r W i l l i a m , j u s t be fo re the r u s h of ac t iv i ty b r o u g h t on by the c o n s t r u c t i o n of r a i l r o a d s , a n d he o f t en h a d di f f icul ty in f i nd ing e n o u g h work fo r h i m s e l f a n d h i s y o u n g e r b ro the r . D u r i n g one lu l l in 1839, W a l l a c e s p e n t some m o n t h s l e a r n i n g the wa tch - m a k i n g t r ade . F o r t u n a t e l y , b u s i n e s s c h a n g e s b r o u g h t th i s to a n e n d be fo re W a l l a c e w a s f o r m a l l y a p p r e n t i c e d , a n d he re- t u r n e d to s u r v e y i n g w i th W i l l i a m .

4. AJ_fred Russel Wallace, "Notebook, 1855--1859," MS, Linnean So- ciety of London, p. 102. This and other quotations are reproduced with permission from the Wallace and other manuscript material in the Library of the Linnean Society of London.

5. Alfred Russel Wallace, My Life: A Record of Events and Opinions (New York: Dodd, Mead, 1905), I, 191.

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Two years later, in 1841, Wallace purchased his first book on na tu ra l history, a shilling pamph le t on botany published by the improbably n a m e d Society for Promot ing the Diffusion of Useful Knowledge. This quickly became his constant com- pan ion on his rambles through the countryside. Eager now to learn more, he was a t t racted by an adver t i sement for a textbook by one of England 's experts, The Elements of Botany by John Lindley. But this expensive purchase , which arrived in July 1842, was a d i sappoin tment because it described all the orders of p lants without indicat ing the British species. Not deterred, Wallace, with the aid of Loudon's Encyclopaedia of Plants, set about annota t ing his copy of Lindley, thereby giving himself the rud iments of a botanical education, s

How Wallace first happened upon Darwin 's Voyage of the Beagle is not known, but he t ipped quotations f rom the first edition into his copy of Lindley. He was apparent ly struck by Darwin 's c o m m e n t tha t to receive the fullest en joyment f rom the pass ing scene, "a traveller should be a botanist , for in all views plants fo rm the chief embel l ishment ." 7 Still another purchase made about the same t ime was Swainson's Treatise on the Geography and Classification of Animals, a remarkab le assembly indeed for someone with no biological background whatever.S

WiUiam's business did not improve, and Wallace again left to look for something else. Early in 1844 he settled upon teaching at the Collegiate School in Leicester. Teaching proved enjoyable enough, but this period is impor tan t for other reasons. The town of Leicester boasted a good library, and Wallace was soon spending his free t ime there reading, among other things, Humbold t ' s Personal Narrative of Travels and Malthus ' Essay on the Principle of Population)

6. J o h n Lindley, T h e E l e m e n t s o f Bo tany , S t ruc tural , Physiological , Sy s t ema t i ca l , and Medical : Be ing a Four th Ed i t ion of T h e Out l ine of the Firs t Pr inc ip les o f B o t a n y (London : Taylor and Walton, 1841 ) ; Wal lace 's copy is in the l ibrary of the L i n n e a n Society of London. J o h n Claudius Loudon, Encyc lopaed i a of Plants (London: L ongma n , 1829-1840).

7. Char les D a r w i n , Journa l and R e m a r k s , 1832 -1836 (vol. 3 of Robert Fitzroy's Narra t i ve of the S u r v e y i n g Voyages of His Majes ty ' s Sh ips "Ad- v en tu re ' and "Beagle," . . . London: H. Colburn, 1839), p. 604. Because the titles of the first and second edit ions differ, the shor t title used here for both will be Voyage of the Beagle. See also notes 39-41, 43 and Appendix, 69.

8. Wi l l iam Swainson , Treat i se on the Geography and Class i f icat ion o f A n i m a l s (London: L ongm an , 1835). Wallace 's copy is in the Library of the L i n n e a n Society of London.

9. Alexander von H um bo l d t and Aim6 Bonpland, Personal Narra t i ve o f Trave l s to the Equ inoc t ia l Reg ions of the N e w Con t inen t , tr. H. M. Wil- l i ams (London: L o n g m a n , 1814-1829) ; w h i c h edit ion Wallace read is

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Most i m p o r t a n t of all was the f r i e n d s h i p of a no t he r y o u n g a m a t e u r n a t u r a l i s t , the en tomolog i s t H e n r y W a l t e r Bates ( 1 8 2 5 - 1892) . Bates, two years y o u n g e r t h a n Wal l ace , h a d s imi la r ly f in i shed his f o r m a l e d u c a t i o n at the u s u a l ear ly age. But, t h o u g h app ren t i c ed to a hos iery m a n u f a c t u r e r i n Leicester , he h a d enro l led in the local Mechan ic s ' In s t i t u t e , one of m a n y such schools set up for the f u r t h e r e d u c a t i o n of w o r k i n g m e n . Here he m a d e the a c q u a i n t a n c e of several n a t u r a l i s t s a n d soon p l u n g e d e n t h u s i a s t i c a l l y in to the s tudy of en tomology. By the t ime he m e t Wa l l ace he had a l ready pub l i shed his first shor t paper . Io

For the first t ime Wa l l ace h a d someone wi th w h o m to ta lk over his discoveries. And Bates soon i n t r o d u c e d h i m to the whol ly n e w wor ld of insects . Wa l l ace was overjoyed. I n shor t order he equ ipped h i m s e l f wi th col lect ing bot t les a nd p in s a n d a copy of S tephens ' M a n u a l of Br i t i sh Coleoptera a nd em- ba rked on this f a s c i n a t i n g n e w study. 11

This happy per iod ended ab rup t ly w h e n Wal l ace ' s b ro the r W i l l i a m died u n e x p e c t e d l y i n F e b r u a r y 1846.12 Wa l l a c e lef t the school to help settle h is affairs, a n d i n J a n u a r y 1847 his second bro ther , John , j o ined h i m i n ca r ry ing on the bus iness . Wa l l ace was i rked by the detai ls of m a n a g e m e n t , fo rmer ly h a n d l e d by his oldest brother . And he was also cu t off f r o m his n e w - f o u n d f r i ends in Leicester , as he wrote p la in t ive ly to Bates [1]. 13

The t ime was no t was ted , however . Wal l ace ' s le t ters to Bates show the e x t r a o r d i n a r y progress i n h is r e a d i n g [1--4114: Dar-

not known. Thomas Robert Malthus, An Essay on the Principle of Popula- tion, 6th ed. (London: Murray, 1826). Which edition Wallace read is not known; in 1908 he took his references from the 6th. According to De Beer ("Darwin's Notebook," part 4, 3On), this is the edition that Darwin read.

10. Henry Walter Bates, "Note on Coleopterous Insects Frequenting Damp Places," Zoologist, 1 (1843), 114--115.

11. James Francis Stephens, Manual of British Coleoptera (London: Longman, 1839).

12. There is some question about the date of William's death. In Wal- lace's autobiography (My Life, I, 239), he gives it as 1846, but he also said he spent only one year at Leicester and correspondingly longer at Neath. In an attempt to straighten out this discrepancy, Poulton persuaded Wallace's son that William must have died in 1845; see E. B. P[oulton], "ALfred Russel Wallace, 1823--1913,'" Proc. Roy. Soc. London, [95B] (1923-1924), viiin.

13. Numbers in brackets refer to the letters Listed in the Appendix. Where additional discussion of the letters is required, it will be found in appropriate footnotes.

14. The dates of letters nos. 2 and 3 are uncertain, and there is no com- pelling evidence to settle the matter. McKinney, following Clodd, has placed them in 1845; see H. Lewis McKinney, "A_l_C-red Russel Wallace and the Discovery of Natural Selection," ]. Hist. Med. Allied Sci., 21 (1966),

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win aga in , LyeU's Principles o f Geology, Vest iges o f the Na tura l His tory o f Creat ion by the t h e n a n o n y m o u s au thor , Rober t Chamber s , Lawrence ' s Lectures on Man, a nd Pr ichard ' s Physical His tory of Man. ~5 Wal lace was a l ready c a u g h t up in the more ph i losophica l ques t ions of evolu t ion , the or ig in a n d d i s t r ibu t ion of species, a n d the d i f ferences be tween species a nd variet ies. W h a t a g ian t step fo rward f r o m the sh i l l ing p a m p h l e t on b o t a n y p u r c h a s e d only six years before l But a n even greater step was i n store.

Late i n 1847, Wal l ace and Bates r ead a n " u n p r e t e n d i n g volume ," A Voyage up the R iver A m a z o n , by a y o u n g A m e r i c a n a m a t e u r na tu r a l i s t , W. H. Edwards . l° Al ready dissat isf ied wi th the i r prospects at home, they wonde red if they could m a k e such a trip. I nqu i r i e s at the Bri t ish M u s e u m about the feasi- bi l i ty of the scheme b r o u g h t a s su rances tha t there was a r eady m a r k e t for a n y t h i n g they m i g h t collect i n this l i t t l e -known region. At once they d e t e r m i n e d to go.

But even as ear ly as this the i r in te res t s were no t l imi ted to collect ing. Wal l ace was a l ready in te res ted no t on ly in the dif- f e rences be tween species a n d var ie t ies , bu t also in the or ig in of species, a n d ju s t before they left he wrote to Bates tha t he h a d b e g u n to "feel r a t h e r dissat isf ied wi th a m e r e local col- lec t ion; l i t t le is to be l e a r n t by it. I should l ike to take some one f a m i l y to s tudy thoroughly , p r inc ipa l ly wi th a view to the theory of the or ig in of species. By tha t m e a n s I a m s t rongly of op in ion tha t some defini te resu l t s m i g h t be ar r ived at [3, 4]." 17

The two y o u n g a d v e n t u r e r s lef t E n g l a n d i n Apri l 1848, ar- r iv ing a m o n t h la te r at Pa rk ( n o w Bel6m) at the m o u t h of the

337 n25. See also note 12. Marchant, in his Wallace, pp. 73-74, puts them in 1847, and that date is used here.

15. Charles Lyell, Principles of Geology: or the Modern Changes of the Earth and Its Inhabitants (London: Murray, 1830-1833, and later editions). Robert Chambers, Vestiges of the Natural History of Creation (New York: Wiley and Putnam, 1845, from the 1st English ed., 1844). Darwin, among others, held a low opinion of the Vestiges; see Appendix, 14 and note 99,. William Lawrence, Lectures on Physiology, Zoology, and the Natural History of Man (London: Printed for J. Callow, 1819, and later editions). James Cowles Prichard, Researches into the Physical History of Man (London: J. & A. Arch, 1813, and later editions). Darwin also read both Lawrence and Prichard (De Beer, "Darwin's Notebook," part 2, 107n).

16. William Hem'y Edwards, A Voyage up the River Amazon (London: Murray, 1847).

17. See AppendLx; also note 97. McKinney ("Wallace and Natural Selection," p. 337) has also pointed out Wallaee's early interest in species, but he overlooked the significance of the related question of species and varieties; see See. HI and note 105.

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Amazon River. ( I t is worth not ing that they had no benefactors , f inancial or otherwise, and tha t they had no resource but themselves. Darwin, on the other hand, was a t tached to an official government miss ion and was able to pay his personal expenses h imsel f . ) Wallace r ema ined for four years, returnJ_ng to Eng- land in October 1852; Bates stayed eleven, not arr iving home unti l July 1859. For the first few months , while they were becoming famfliax with their new surroundings, they traveled together. Then they par ted company, each t ravel ing independently f rom then on.

Wallace journeyed up the Amazon and the Rio Negro to the place where the lat ter joins the Orinoco, a spot already made f amous as the far thes t point reached by Humbold t com- ing f rom the other direction. Bates chose to go to the Upper Amazon, spending considerable t ime at E t a , 1400 miles up- s t r eam f rom the Atlantic Ocean. These intrepid explorers were entirely on their own in the s t range and of tent imes forbidding tropics of Brazil, contending with an uncer ta in supply of food, hazardous travel, indolent or dishonest nat ives (spoiled, they thought, by being half-civil ized), illness, and isolation. They f inanced their exploits by the sale in Europe of duplicate collections, ma in ly of insects.

Unfor tunate ly , the scientific and f inancial results of Wal- lace's travels were disappoint ingly meager , but with reason. Owing to a misunders tanding , the specimens collected dur ing the last two years were still wai t ing unshipped when he returned to Bar ra (now M a n a u s ) on his way back to England. And so, unhappi ly , these as well as the r ichest pa r t of his pr ivate collections were on board with h im when the ship caught fire at sea. Near ly everything was lost as passengers and crew rushed to leave the burn ing vessel, some 700 miles off Ber- muda . Wallace and his companions were forced to spend ten days in open boats before they were finally rescued by a pass ing freighter.

As a consequence of this disaster, Wal lace had to piece together the few papers and two books tha t he wrote on his re tu rn home f r o m letters, a few rescued notes, and his memory . He pursued his growing interest in the geographical distribution of an imals in papers on the monkeys and the butterflies of the Amazon Valley, r emark ing on l imits and barr iers to their distr ibution and on the impor tance of labelling specimens with the exact locality where they were found for the proper s tudy of this distribution. 18 Besides a small book on the pa lm

18. Alfred Russel Wallace, "On the Monkeys of the Amazon," Proc. Zool. Soc. London, 20 (1852) , 107-110; and "'On the Habits of the Butter-

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t r ees of the r eg ion , he a lso wro te the s to ry of h i s exped i t i on , A N a r r a t i v e o f T r a v e l s o n t h e A m a z o n a n d R i o N e g r o . 19 I t is a ta le of h igh a d v e n t u r e but , due to t he loss of so m u c h of h is m a t e r i a l , i t is less r e w a r d i n g f r o m a sc ient i f ic p o i n t of view.

Ba tes w a s m o r e f o r t u n a t e , a l t h o u g h he took good ca r e to s end h is l a s t co l l ec t ions h o m e on th ree sh ips i n s t e a d of one. On h is r e t u r n , he wro te a ser ies of p a p e r s on the i n sec t f a u n a of the A m a z o n Val ley , in one of w h i c h h e d e v e l o p e d h is f a m o u s theory of p ro t ec t i ve co lo ra t ion , s t i l l k n o w n as B a t e s i a n m i m - icry. 20 His a c c o u n t of h is t r ave l s , T h e N a t u r a l i s t o n t h e R i v e r

A m a z o n s , was m o r e sc ient i f ic t h a n W a l l a c e ' s a n d also, to Wa l - l ace ' s c h a g r i n , m o r e success fu l . 21

But W a l l a c e ' s m i s f o r t u n e p r o v e d a b o o n a f t e r all. H e h a d c o u n t e d on the m o n e y f r o m the sa le of the los t co l l ec t ions ; w i t h o u t i t he f a c e d a r e t u r n to s u r v e y i n g or a n o t h e r t r ip . For - t u n a t e l y for b o t h h i m s e l f a n d the wor ld , he chose the l a t t e r course , d e c i d i n g a f t e r m u c h s tudy t h a t the M a l a y a n r eg ion of fe red the o p p o r t u n i t i e s he sought . A n d so, in the s p r i n g of 1854, he set off aga in , on the e igh t -yea r e x p l o r a t i o n of the t rop ics on the o t h e r s ide of the wor ld "wh ich c o n s t i t u t e d the c e n t r a l a n d c o n t r o l l i n g i n c i d e n t of m y l ife." 22

Once a g a i n W a l l a c e h a d to m a k e do w i th t h i n g s as he f o u n d them, a c c o m m o d a t e h i m s e l f to loca l c u s t o m s , l e a r n n a t i v e l an- guages , f ind food a n d she l t e r w h e r e h e could , a n d use a n y a v a i l a b l e m e a n s of t rave l . As a p r a c t i c a l m a t t e r , the w e a t h e r a n d t r a v e l i n g a r r a n g e m e n t s l a r g e l y d e t e r m i n e d his i t i ne ra ry . T h e f irst two y e a r s were s p e n t a r o u n d S i n g a p o r e a n d in Borneo , the n e x t five in the a r e a f r o m Celebes to n o r t h e r n N e w G u ine a , a n d the l a s t y e a r in T imor , J a v a , a n d S u m a t r a .

A p a r t i c u l a r a i m th is t ime w a s a m o r e t h o r o u g h inves t iga - t ion of the p r o b l e m s of the g e o g r a p h i c a l d i s t r i b u t i o n of a n i m a l s . W a l l a c e h a d c h o s e n well , fo r the l e s sons of a n i s l a n d wor ld

flies of the Amazon Valley," Trans. Entomol. Soc. London, N.S. 2 (1852- 1853), 253-264.

19. Alfred Russel Wallace, Palm Trees of the Amazon and Their Uses (London: J. Van Voorst, 1853); and A Narrative of Travels on the Ama- zon and Rio Negro (London: Reeve, 1853).

20. Henry Walter Bates, "Contributions to an Insect Fauna of the Ama- zon Valley. Lepidoptera: Heliconidae,'" Trans. Linn. Soc. London, 23 (1862), 495-515.

21. Henry Walter Bates, The Naturalis t on the River Amazons (London: Murray, 1863). Many of Bates" more interesting comments on the origin of species and on geographical distribution were omitted from the 2rid abridged edition (1864).

22. Wallace, My Life, I, 336.

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axe even more vivid t h a n those of a c o n t i n e n t a l reg ion; D a r w i n had also discovered this w h e n he vis i ted the i s l ands of the Galapagos Archipelago. A l though Wa l l ace was aga in support - i ng h i m s e l f by the sale of dup l ica te col lect ions, his theore t ica l in te res t s were neve r f a r f r o m his m i n d .

The mos t spec tacu la r resu l t of this second tr ip was the in- d e p e n d e n t d iscovery of the theory of n a t u r a l select ion, b u t this was on ly one of m a n y i m p o r t a n t c o n t r i b u t i o n s to evolu- t iona ry theory. A n d happ i ly enough , h is d i s c e r n i n g a n d well- wr i t t en a c c o u n t of the region , T h e Malay Archipelago, b e c a m e a wor thy r iva l to D a r w i n ' s Voyage of the Beagle a n d Bates 's Natura l i s t on the R iver Amazons.e3

The three pape r s of specia l i m p o r t a n c e i n t r ac ing the de- v e l o p m e n t of Wa l l aee ' s ideas on n a t u r a l se lec t ion wil l n o w be cons ide red i n detail .

II. T H E "LAW"

To discover how the extinct species have from time to time been replaced by new ones down to the very latest geological period, is the most difficult, and at the same time the most interesting problem in the natural history of the earth. The present inquiry, which seeks to elim- inate from known facts a law which has determined, to a certain degree, what species could and did appear at a given epoch, may, it is hoped, be considered as one step in the right dixection towards a complete solution of it.

Wallace, "'On the Law which has regulated the Introduction of New Species"

Wal l ace ' s "powerfu l essay," e4 "On the Law w h i c h has regula ted the I n t r o d u c t i o n of New Species," was pub l i shed in Sep- t e m b e r 1855, b u t to h is d i s a p p o i n t m e n t i t a t t rac ted li t t le no t ice at the t ime. ~5 He h a d w r i t t e n it i n the p reced ing F e b r u a r y d u r i n g a lu l l i n his col lec t ing act ivi t ies , i n d u c e d by the pub- l i ca t ion of E d w a r d Forbes ' s theory of polar i ty , a n d he h a d hoped at the leas t for some c o m m e n t f r o m this b r i l l i a n t y o u n g n a t u r a l i s t ; u n k n o w n to Wal lace , however , Forbes ha d died i n N o v e m b e r 1854. Except for Bates 's , o ther r e sponse was mi n i - real , i f no t d i spa rag ing .

Wal l ace p h r a s e d his '~law" as fol lows: Every species has c o m e into ex i s tence co inc iden t both in space and t ime w i t h a pre-exis t ing closely allied species. 26 Al though the evo lu t iona ry

23. Alfred Russel Wallace, The Malay Archipelago: The Land of the Orang-utan, and the Bird of Paradise: a Narrative of Travel, with Studies of Man and Nature (London: Macmillan, 1869).

24. Thomas Hera'y Huxley, in Darwin, Life and Letters, I, 539. 25. Alfred Russel Wallace, "On the Law which has regulated the Intro-

duction of New Species," Ann. Mag. Nat. His~. [2], 16 (1855), 184--196.

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implications of this s tatement are obvious, Wallace at this thne had no suggestions on a mechan i sm of change. His speculations were rooted in his long-standing interest in the geographical distribution of animals and colored by his extensive practical experience in the tropics of two hemispheres.

The study of zoogeography, in which Wallace was to become an acknowledged master, was still clouded in mystery when he first came upon it in Swainson's T r e a t i s e on the subject :

We may, indeed, build a theory upon every thing in nature: but the more we investigate, the stronger will be our con- viction in the following deduction : - - T h a t the pr imary causes which have led to different regions of the earth being peopled by different races of animals, and the laws by which their dispersion is regulated, must be for ever hid f rom h u m a n research [to which Wallace wrote "no" in the mar- gin]. This conclusion is strengthened by the inference which will be drawn f rom the facts we shall subsequently state; an inference so well expressed by a very intelligent writer, that we shall give it nearly in his own words. "It appears that various tribes of organised beings were originally placed by the Creator in certain regions, for which they are by their nature peculiarly adapted [Wallace's underlining]." 2r

In a modification of the quinarian system of William Sharpe Macleay, Swainson divided the earth into five regions accord- Lug to what he believed to be the five major races of mankind; animal groups were likewise divided into fives. The divisions were mathematical , the reasons not only unknown but un- knowable. But Wallace questioned Swainson f rom the first, noting that "there appears not to be the slightest reason for believing a priori that all groups of animals are divided into the same number of types of forms or divisions . . ." 28

Over the years Wallace's ideas matured. Traces of this can be found, as mentioned earlier, in his papers on the monkeys and the butterflies of the Amazon Valley, as well as in his N a r r a t i v e , but he was undoubtedly handicapped by the loss of so much of his South American material. The appearance of Forbes's paper spurred Wallace to collect and organize his thoughts, for Forbes's theory of polarity as an explanation of organic changes through geologic time seemed as untenable

26. Ibid. , p. 186. 27. Swainson, Treat i se on Geography , p. 9. 28. Note written by Wallace in his copy of Swainson's Treat i se on

Geography , p. 223.

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to h im as had Swainson's theories about the present geographical distribution of animals.

A pervading influence on Wallace, one might almost say the pervading influence, was Charles Lyell's Principles of Geology. Wallace had read Lyell at least as early as 1846, and he had accepted wholeheartedly Lyell's application of the principle of unfformitar ianism to geology, But Lyell's influence was more profound than that. From the notebook kept by Wallace dur- Lug his Malayan travels, it seems that he had with h im a copy of the four th edition of Lyell's Principles, and the discussion here refers to this edition. 29

In a large measure, Wallace's 'T~lotebook" is a long, private a rgument with /_,yell, refuting m a n y of the latter's biological theories. Wallace was thus stimulated to broaden and deepen his own thinking. It should be noted in Lyell's defense, however, that he was neither a trained biologist nor a collector of living specimens (a l though he had collected insects as a youth) , and that he lacked the extensive first-hand experience with living things acquired through long years by a Lamarck, a Darwin, or a Wallace.

It has often been said that Lyell nearly stumbled onto evolutionary theory himself; it has even been suggested that he was an evolutionist in secret, at least at first. 30 But in fact he was far removed at m a n y critical points. He had missed altogether the crux of Wallace's p a p e r - - t h e relationship of species in both time and s p a c e - - a n d so a theory of descent was not only not a logical deduction, it was irrelevant. On his own terms he believed he had applied the principle of unfformi- tarianism to organic changes by showing the gradual extinction and creation of species. But he believed firmly in the stability of species, scarcely a good foundat ion for a theory of evolution.

Lyell was well acquainted, however, with Lamarck 's theory of the t ransmutat ion of species, having first read it in 1827. Although he found the ideas provocative, he confessed in a letter to Mantell that "I read him rather as I hear an advocate on the wrong side, to know what can be made of the case in good hands." 31 He included a lengthy summary in his Prin-

29. Lyell, Principles of Geology, 4th ed. (London: Murray, 1835). See a l s o n o t e 4 .

30. Gertrude Himmelfarb, Darwin and the Darwinian Revolution (Garden City, N.Y.: Doubleday, 1959), pp. 180-189. An early proponent of this point of view was Thomas Huxley, in Darwin, Life and Letters, I, 543--548.

31. Katherine Lyell, ed., Life, Letters and ]ournals of Sir Charles Lyell (London: Murray, 1881), I, 168.

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ciples, at first in protest against the pretension that species were "capable . . . of being indefinitely modified in the course of a long series of generations," but later to give due honor to Lamarck and his evolutionary theories. 32 Wallace and Dar- win both read Lyell's summary. Parenthetically, it might be observed that Lyell accepted to a limited degree what later generations have considered the hal lmark of Lamarckism, the inheri tance of acquired characteristics. In Lyell's case, this meant the possible inheritance of acquired habits that remained within the strict limits of predetermined variation. To a limited extent, Darwin was also to accept this Larnarckian tenet, but Wallace never did.

Lyell's view of geographical distribution was more compre- hensive than Swainson's. He had broadened the scope to in- clude time and change, al though there was no hint of evolution :

I f the views which I have taken are just, there will be no difficulty in explaining why the habitations of so m a n y species are now restrained within exceedingly narrow limits. Every local revolution, such as those contemplated in the preceding chapter, tends to circumscribe the range of some species, while it enlarges that of others; and ff we ale led to infer that new species originate in one spot only, each must require time to diffuse itself over a wide area. It will follow, therefore, f rom the adoption of this hypothesis, that the recent origin of some species, and the high antiquity of others, are equally consistent with the general fact of their limited distribution, some being local, because they have not existed long enough to admit of their wide dissemination; others, because circumstances in the animate or inanimate world have occurred to restrict the range which they may once have obtained.

As considerable modification in the relative levels of land and sea have taken place in certain regions since the existing species were in being, we can feel no surprise that the zoologist and botanist have hitherto found it difficult to refer the geographical distribution of species to any clear and determinate principles, since they have usually speculated on the phenomena, upon the assumption that the physical geography of the globe had undergone no material altera- tion since the introduction of the species now living. So long as this assumption was made, the facts relating to the

32. Lyell , Principles of Geology, 1st ed. ( 1 8 3 2 ) , II ,1; lOth ed. ( 1 8 6 8 ) , II, 246.

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geography of plants and animals appeared capricious in the extreme, and by many the subject was pronounced to be so full of mystery and anomalies, that the establ ishment of a satisfactory theory was hopeless. 33

Although Lyell's biological theories were ambiguous and ill- defined, his contributions to geology had added new dimensions to the history of the earth. In part icular, he had expanded on the unfformi tar ian theory of the Scottish geologist James Hut- ton (1726-1797) that a t tempted "to explain the fo rmer changes of the earth's crust by reference exclusively to na tura l agents," a4 and he had vigorously attacked the commonly accepted belief that life on the ear th had begun in the year 4004 B.C., as calculated by James Usher, Archbishop of Armagh. Wallace's efforts ( and Darwin's too) would have been crippled without Lyell.

Evolution was advocated, however, in a book published not long afterward, Vestiges of the Natural History of Creation. But its anonymous author, Robert Chambers, "a private person with limited opportunities for study," was no biologist either, and his popular work was ridiculed by scientists. Nev- ertheless, Wallace thought his hypothesis ingenious, as he wrote to Bates [2, 3].

For somewhat different reasons, two editions of Darwin's Voyage of the Beagle should also be included here. Wallace read the first edition (1839) quite early, perhaps as early as 1842 according to his first letter to Bates and the note in his copy of Lindley; and f rom evidence in the '~Iotebook,'" he took a copy of the second (1845) along with him to the Malay Archipelago.

Wallace began his paper "On the Law" by noting the long- cont inued series of geologic changes. Then, applying the uni- formi tar ian principle to organic changes, he proposed "'a like gradation and natura l sequence f rom one geological epoch to another." 35 (Lyell had suggested the slow and gradual extinction and creat ion of species, but with no hint of descent .) F rom a series of propositions relat ing to "organic geography and geology," Wallace then deduced his 'qaw," which supported a hypothesis that might explain the past and present distribution of life upon the ear th that had occurred to him, he said, about ten years earlier.

First of the four main questions i l luminated by Wallace's

33. Ibid., 4 th ed., III , 165-166 . 34. Ibid., 12th ed. ( 1 8 7 5 ) , I, 73. 35. Wa l l ace , "'On t he L aw, " p. 184.

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' l aw" was that perennial problem, "the system of natural af- fi_nities.'" z~ If the 'qaw" were true, species would be related to closely allied species which had preceded them. This relationship could rarely be expressed for long by a straight line. The divergence, uneven rates of change, and extinction of species, complicated by the f ragmentary fossil record, could be better represented by a branching "as intricate as the twigs of a gnarled oak." aT Artificial systems of classification based on circles or a fixed number of divisions were unnecessary con- trivances.

Lamarck would have agreed. He believed that species were defined by the gaps between them that were produced by extinction and by the f ragmentary record, and that two or more diverging species would merge going backward in time. To Lyell this would have been unthinkable, convinced as he was of the real and permanent existence of species in nature.

Next, in answer to the "singular phenomena" of the distribution of animals and plants in space, Wallace offered some original suggestions, as He clearly recognized the part played by geographical isolation in the origin of peculiar forms of life in long-isolated places, and also the divergence f rom a widespread "antitype" that results in two or more representa- tive forms in different regions of the world. He brought forward as examples the problems posed by the inhabitants of both ancient and recent island groups and mounta in ranges.

Of part icular interest are Wallace's suggestions regarding the Galapagos Islands, where the "phenomena . . . have not hitherto received any, even a conjectural explanation." a9 Dar- win had mentioned in the first edition of his Voyage of the Beagle the many peculiarities of distribution that he had found in this small archipelago, six hundred miles off the coast of Ecuador, concluding only: "But there is not space in this work, to enter on this curious subject." 40

Although Darwin expanded on this in the second edition of the Voyage, his remarks were again inconclusive :

The only light which I can throw on this remarkable difference in the inhabitants of the different islands, is, that very strong currents of the sea running in a westerly and W.N.W. direction must separate, as far as transportal by

36. Ibid., pp. 186--188. 37. "Divergence" was an i m p o r t a n t pa r t of Da r win ' s theory. See also

notes 105, 111-113, 125-127, and Wallace, "On the Law," p. 187. 38. Ibid., pp. 188-190. 39. Ibid., p. 188. 40. Darwin , Voyage of the Beagle (1839) , p. 475.

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sea is concerned, the southern islands f rom the nor thern ones; and between these nor thern islands a strong N.W. current was observed, which must effectually separate James and Albemarle Islands. As the archipelago is free to a most remarkable degree f rom gales of wind, neither the birds, insects, nor lighter seeds, would be blown f rom island to island. And lastly, the profound depth of the ocean between the islands, and their apparently recent ( in a geological sense) volcanic origin, render it highly unlikely that they were ever united; and this, probably, is a far more important consideration than any other, with respect to the geographical distribution of their inhabitants. Reviewing the facts here given, one is astonished at the amount of creative force, if such an expression may be used, displayed on these small, barren, and rocky islands; and still more so, at its diverse and yet analogous action on points so near each other. I have said that the Galapagos Archipelago might be called a satellite at tached to America, but it should rather be called a group of satellites, physically similar, organically distinct, yet intimately related to each other, and all related in a marked, though m u c h lesser degree, to the great American continent. 41

In other words, these islands were geologically recent, separated not only by deep ocean but also by currents, and without winds strong enough to have blown birds, insects, or seeds f rom one island to another. With no means of dispersal and a limited amount of time, it is no wonder that Darwin seemed puzzled. Actually, his private thinking was m u c h in advance of this public statement, for he had already worked out his theory of na tura l selection as an explanation of the "creative force" at work in the islands, but Wallace had no way of knowing this.42

Wallace suggested, on the contrary, that these were islands of high antiquity that h a d been peopled by the agency of wind and water, as other islands were peopled, and that, the pre- existing species having died out, only variously modified prototypes now remained. The islands of the Malay Archipelago differed in being separated by shallow seas (this was written shortly before Wallace discovered in the summer of 1856 the division between the Australian and Oriental Regions since

41. Charles Darwin, Journal of Researches into the Natural History and Geology of the Countries visited during the Voyage of H.M.S. Beagle round the World, 2rid ed. (London: Murray, 1fl45), p. 398.

42. Charles Darwin and Alfred R. Wallace, Evolution by Natural Selec- tion (Cambridge: Cambridge University Press, 1958), pp. 116--121.

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BARBARA G. BEDDALL

immorta l ized as "Wallace's Line") , probably indicating an earlier land connect ion that could account for the basic similari- ties in their faunas . Is lands like Great Britain, which had only recent ly (geologically speaking) been separated f r o m continents, would have few groups peculiar to themselves.

Wallace 's reason for first writ ing Darwin in October 1856 can only be guessed at. Perhaps he hoped for some c o m m e n t f rom Darwin on these suggestions. (An entry in his "Notebook" shows that he was rereading Darwin about this t ime.) 43

Approaching the prob lem f rom another point of view, Wal- lace next considered the close geographical proximity of closely allied species in r ich g r o u p s - - s u c h as the hummingb i rds , toucans, pa lms , orchids, and various famil ies of bu t t e r f l i e s - - and asked, "why are these things so?" These facts of distribution would not only be explained by his "law," they would also be its necessary results. A corollary was tha t species have not ar isen more than once, in two widely separa ted places. LyeU, for all his shor tcomings in biology, had believed that species were "'created" in one place only, a l though he recognized that this might give a false impress ion of centers of creation.

Thirdly, Wallace inquired into the p h e n o m e n a of geological distribution, the distribution of species in t ime ra ther than in space. 44 Again he pointed out that proximity and gradual change were the rule and tha t species had been "created" only once. I t is in connect ion with geological distribution, however, that reasons for Lyell's fai lure to hit upon evolut ionary theory become clear. Especially impor tan t is the ma t t e r of the ex- t inction of species. On this point Lyell and Lamarck represented two opposite points of view. Lyell, convinced of the immutabi l i ty of species, could hardly believe otherwise than in their absolute extinction. He was applying the un i fo rmi ta r ian principle in extending backward in t ime the acknowledged present-day and probable fu ture extinct ion of species:

Although we have as yet considered one class only of the causes ( the organic) by which species m a y become extermin- ated, yet it cannot but appear evident that the cont inued action of these alone, throughout myr iads of fu ture ages, m u s t work an entire change in the state of the organic creation, not mere ly on the cont inents and islands, where the power of m a n is chiefly exerted, but in the great oceans, where his controul [sic] is a lmost unknown. The mind is pre-

43. Wal l ace , "Notebook," p. 60. See also no t e s 81, 103, a n d 162, a n d Append ix , 69.

44. Wal l ace , "On t he L aw, " pp. 190-195 .

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pa red by the contempla t ion of such fu ture revolutions to look for the signs of others, of an analogous nature , in the m o n u m e n t s of the past. Ins tead of being astonished at the proofs there mani fes ted of endless muta t ions in the an imate world, they will appear to one who has thought profoundly on the f luctuat ions now in progress, to afford evidence in favour of the un i formi ty of the system, unless, indeed, we are precluded f rom speaking of uni formi ty when we charac- terize a principle of endless variat ion. 45

Lamarck , on the other hand , considered tha t only a few large land an imals at mos t had become extinct, and then through the agency of man . Even earlier than Lyell he had disagreed with the catastrophis ts , who thought tha t life had been wiped out at var ious t imes by universa l cataclysms. Lyell suggested a course of gradual extinct ion and creat ion of species, But La- m a r c k had proposed tha t earlier species had gradual ly been changed into present-day species through t ransmuta t ion ; ex- t inction thus played no impor tan t par t in his system. LyeU, however, disagreed completely with Lam arck :

To pursue this t ra in of reasoning fa r the r is unnecessary; the geologist has only to reflect on wha t has been said of the habi ta t ions and stat ions of organic beings in general, and to consider them in relat ion to those effects which were contempla ted in the second book, as resul t ing f rom the ig- neous and aqueous causes now in action, and he will im- media te ly perceive that , amids t the vicissitudes of the ear th 's surface, species cannot be immorta l , bu t mus t perish, one af ter the other, like the individuals which compose them. There is no possibility of escaping f rom this conclusion, wi thout resort ing to some hypothesis as violent as tha t of Lamarck , who imagined, as we have before seen, tha t species are each of t hem endowed with indefinite powers of modify ing their organizat ion, in conformi ty to the endless changes of c i rcumstances to which they are exposed. 46

Wallace steered a middle course. He agreed with Lyell tha t some species migh t become extinct, but he also agreed with L a m a r c k tha t modified prototypes migh t remain . I f ext inct ion were the rule, as Lyell suggested, then some sort of "'creation" was necessary to fill the gaps. Here Lyell was quite vague, his hypothesis on the original introduct ion of species reading as follows :

Each species m a y have had its origin in a single pair , or

45. Lyell, Principles of Geology, 4th ed., III, 140--141. 46. Ibid., 155-156.

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individual, where an individual was sufficient, and species m a y have been created in succession at such t imes and in such places as to enable them to mult iply and endure for an appointed period, and occupy an appointed space on the globe.4~

This creat ion of new species was not readily seen because it was an inf requent occurrence taking place over a long period of t ime at different places on the earth.

Lamarck ' s solution to this problem was the mutabi l i ty of species, though the agency he sugges ted - - inhe r i t ance of characteristics acquired through the will of the a n i m a l - - w a s scoffed at even in his own day.

The author of the Vestiges had still another suggestion:

The idea, then, which I f o rm of the progress of organic life upon the g l o b e - - a n d the hypothesis is applicable to all s imilar theatres of vital be ing- - i s , that the simplest and most primitive type, under a law to which that of like-pro- duction is subordinate, gave birth to the type next above it, that this again produced the next higher, and so on to the very highest, the stages of advance being in all cases very s m a l l - - n a m e l y , f rom one species only to another; so that the phenomenon has always been of a s imple and modest character . 4s

Chambers ' proposal that one species gave bir th directly to the next higher brought derision upon his book. Nevertheless, he should at least be credited with having emphasized a na tu ra l method.

Wallace, however, had no theory on how extinct species were replaced. For the present , he confined himsel f to "what species could and did appear ." He saw that this was nei ther a r a n d o m process nor a s t ra ight line f rom simple to complex, as proposed by the proponents of the theory of progressive development. A theory of gradual change combined with his "law" would, he thought, account for the observed fac ts and even for apparen t retrogressions.

Lyell, as a geologist f ami la r with the anomalies of the fossil record, had protested against the theory of progressive development because he disputed the accuracy of the facts :

I t was before remarked , that the theory of progressive development arose f rom an a t tempt to ingraf t the doctrine of the t ransmuta t ionis ts upon one of the mos t popular gen- eralizations in geology. But m ode rn geological researches

47. Ibid., 99-100. 48. C h a m b e r s , Vestiges, 1st ed., p. 167.

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have almost destroyed every appearance of that gradation in the successive groups of animate beings, which was supposed to indicate the slow progress of the organic world from the more simple to the more compound structure. 49

Although evolutionary trends do exist (and Lyell later accepted a general progression), the record is far f rom straight- forward, and its message is not easily read. The theory of evolution, with natural selection as an agent of change, was to make this record more comprehensible, for it postulates suc- cessful adaptation rather than any necessary progression. Wal- lace was still several years away f rom this answer, but he was heading in the right direction.

It was Forbes's metaphysical theory of polarity that had prompted Wallace to set out his own though t s ) ° According to this theory, the distribution of organized beings (genera rather than species) in time manifested a quality known as polarity: an "arrangement in opposite directions with a development of intensity towards the extremes of each." 51 This relation was used to explain the larger number of generic forms in the earlier epochs of the Palaeozoic (Silurian and Devonian) and in the later epochs of the Neozoic (Cretaceous, Tertiary, and present) , compared with the smaller number to be found in the intervening time.

Wallace objected to this theory on several grounds. First of all, it invoked an obscure and hypothetical cause when "the facts may be readily accounted for on the principles already laid down." 52 In typical Lyellian style, he stressed the vast amounts of time involved and the action of natural causes, and he went on to propose that the rates of creation and extinction of species were related to unequal rates of geologic change, more species being created during quiet periods of the earth's long history and more becoming extinct during violent periods.

Even more to the point was Wallace's criticism that Forbes's theory presupposed the completeness of the geological record ( an error also made by those who believed in the theory of progressive development) . Wallace knew it was f ragmentary, but he would have been surprised to know that a century later fossils are still thought to represent less than one per cent of the species that have existed, Polarity's foundat ion was fragile at best.

49. Lyell, Principles of Geology, 4th ed., III, 14-15. 50. Edward Forbes, "'On the Manifestation of Polarity in the Distribu-

tion of Organized Beings in Time," Proc. Roy. Inst. London, 57 (October 1854), 332--337.

51. Ibid., p. 336. 52. Wallace, "On the Law," p. 192.

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BARBARA G. BEDDALL

The four th and final point of Wallace 's paper was another subject tha t had a t t rac ted m u c h attention, the puzzling problem of rud imen ta ry organs. 5~ LyeU later admit ted tha t he had missed their significance and as a result had omit ted f rom his s u m m a r y mos t of the examples given by Lamarck . 54 This is not surprising, however, because L a m a r c k had interpreted them in his own light, believing them to be the result of "the permanent disuse of an organ, arising from a change of habits, [which caused] a gradual shrinkage and ultimately the disap- pearance and even extinction of that organ." 55 Wallace, like Chambers , thought that rud imen ta ry organs showed relation- ships, but he mis interpre ted them, confusing vestigial with nascen t organs. He did, however, ask the r ight question. "If each species has been created independently, and without any necessary relat ions with pre-existing species, wha t do these rudiments , these apparen t imperfect ions m e a n ? " 56

Wallace ended his paper grandly: "Granted the law, and m a n y of the most impor tan t facts in Na tu re could not have been otherwise, bu t are a lmost as necessary deductions f rom it, as are the elliptic orbits of the planets f rom the law of gravitat ion." 57 The response to it was hardly encouraging. More than two years passed before Bates 's letter congratulat- ing h im arrived f rom the Upper Amazon [9]. In the mean t ime , his agent, Samuel Stevens, wrote tha t he had heard several natural is ts object to Wallace 's "theorizing" when what was needed was more facts•

III . THE "NOTE"

• . . w h y should a special act of c rea t ion be required to call into existence an o rgan i sm differing only in degree f r o m ano the r wh ich has been produced by exis t ing laws?

Wallace, "'Note on the Theory of P e r m a n e n t and Geographical Varieties'"

In the fal l of 1857 Wallace sent off to the Zoologist a short paper entit led "Note on the Theory of P e r m a n e n t and Geo- graphical Varieties." 58 The first pa r t of his hypothesis had been his ' Jaw" on "what species could and did appear"; the

53. Ibid., pp. 195--196. 54. Lyell, Principles of Geology, 12th ed., II , 274. 55. J e a n Baptis te Lamarck , Zoological Philosophy: an Exposition zvith

Regard to the Natural History of Animals, tr. H u g h Elliot (London: Mac- mi l lan , 1914; repr inted, N e w York: Hafne r , 1963), p. 115.

56. Wallace, "On the Law," p. 195. 57. Ibid., p. 196. 58. Alfred Russel Wallace, "Note on the Theory of P e r m a n e n t and

Geographica l Varieties," Zoologist, 16 (1858) , 5887-5888.

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second par t concerned the distinction be tween species and varieties, as he wrote to Bates in J a n u a r y 1858 [19]. This question still exists, of course, but the premises are entirely different. Then, species were real and pe rmanen t , "created" with cer tain relatively fixed characteris t ics; varieties, on the other hand , were produced by ordinary generat ion within strict l imits of variation. Varieties were, ff anything, an inconvenience, inter- fer ing with the rigid definition of species. Today, however, species have only a relative pe rmanence ; they also come about by ordinary generat ion, and variet ies m a y ( though they do not a lways) lead to the fo rmat ion of new species.

The definition of species has been a t roublesome problem at least as f a r back as the t ime of A.ristofle. Not unti l the t ime of the English natura l is t John Ray ( 1 6 2 8 - 1 7 0 5 ) was the t e rm limited to wha t we would today recognize as a breeding unit , including sex, color, and age variants . Ray concluded his discussion of species in his Historia Plantarum with the observa- tion that "an imals that differ in species preserve their distinct species pe rmanen t ly ; one species never springs f rom the seed of another nor vice versa." 59

Linnaeus ( 1 7 0 7 - 1 7 7 8 ) was strongly influenced by Ray. A clear notion of wha t was m e a n t by the t e rm "species" was a necessi ty in organizing his Sys tema Naturae, first publ ished in 1735. At first he was convinced of the p e r m a n e n c e of species, but later, af ter he had become acquainted with the vast number of new fo rms brought home by explorers f r o m all over the world, he was not so sure. Nevertheless , the belief in the creat ion of p e r m a n e n t species, which was closely inter twined with religion, was the general ly accepted opinion in the first ha l f of the n ine teenth century, in spite of the heresies of people like L a m a r c k and Chambers .

This vexed question had a t t racted Wallace 's at tent ion f rom the start, as can be seen f r o m his ear ly letters to Bates [3, 4]. I t became a ma t t e r of daily concern when, as a collector, he was confronted over and over again with the task of properly identifying specimens. U n h a m p e r e d by any strong re- ligious convictions, he worried at the prob lem as a dog does a bone, as his "Notebook" attests.

Lamarck had clearly grasped the gradual na tu re of change in species through time. Chambers , a l though he believed in evolution, thought tha t it proceeded by sudden leaps f rom species to species r a the r than by the accumula t ion of small changes. But Lyell stood on the opposite side of the fence,

59. Barbara G. Beddall, "Historical Notes o n A v i a n Class i f icat ion," Syst. Zooi., 6 (1957), 134.

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firmly convinced of the fixity of species and of strict limita- tions to variation. Had either Chambers or LyeU been trained biologists or collectors, exposed day after day to the endless variations to be found in nature, they might have modified their views. But Wallace benefited f rom LyeU's position, using it as a springboard f rom which to develop his own. Although Lyell was unaware that he had been cast in the role of principal antagonist, his importance to Wallace cannot be over- rated.

It seems likely that the '~Notebook" was intended as the basis of a projected book about which Wallace wrote to Darwin in the fall of 1857 and to Bates early in 1858 [16, 19]. 60 The entries of part icular interest here were written between June 1855 (not long after the formulat ion of his ' ]aw") and No- vember 1857, assuming that the dates scattered here and there are accurate indicators of the time and order of writing. They cover an assortment of topics relating to evolut ion--proofs of design, the theory of progressive development, t ransmutat ion and special creation, the "balance of species," geological changes and gaps in the fossil record, and the doctrine of the morphology of p l a n t s - - a n d reflect Wallace's progress toward a solution of the problem of the origin of species. ~1

The argument f rom design was teleological, presuming that a contrivance existed in accordance with a preconceived plan. Adaptation between structure and function was recognized, but it was thought that a structure was provided simply because a funct ion required it. Wallace wondered, however, how an animal could have necessities before it came into existence? And how could it "continue to exist unless its structure enabled it to obtain food?" B2 He thought that the arguments brought forward as proofs of design were absurd; not only were they insulting to the intelligence of a Supreme Being, but they also placed narrow limits on His power.

Wallace returned several times to the inconsistencies in the geological record that made the theory of progressive development so troublesome for Lyell, observing that "the supposed contradictions all arise f rom considering it necessary that the highest forms of one group should appear before the lowest of the next succeeding, not considering that each group goes on

60. The information that Wallace's plan for his book is on the r e v e r s e side of the fragment about the jaguars was sent me by Sydney Smith.

61. Wallace, "Notebook," pp. 12--100. See also McKinney ("Wallace and Natural Selection," pp. 342-347), who has also pointed out that this is essentially a long argument with LyeU.

62. Wallace, '¢Notebook," p. 12.

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progressing after other groups have branched f rom it." 0a LyeU's static concept of species did not allow for such a dynamic interpretation.

Were there connecting forms between groups, and if so, what were they? Wallace remarked that "as long as these most important characters [of groups] remain undiminished, no al- terations of external form or habits can be held to shew any signs of a transition." 64 He questioned the popularly held transitional status of the seal, asking, "is not the Cetaceous group rather a modification of mammal i a to an aquatic life than a link connect ing them with fishes?" 05 He noted that neither the bat nor the hummingbi rd is a transit ion form either, because each contains the characteristic features of m a m m a l or bird in a high state of development. The bat 's wing is even less like a bird's wing than are less modified forelimbs of other mammals . Therefore, the highest forms of one group cannot be a transition to the lowest forms of the next.

Lyell had also had trouble in accounting for the appearance of new species. Having decided in favor of the stability of species, he was obliged to settle for their extinction and "creation," admitt ing to only a limited amount of variation. Wallace was groping his way forward, however, questioning LyeU's assumptions :

Lyell says that varieties of some species may differ more than other species do f rom each other without shaking our confidence in the reality of species- -But why should we have that confidence? Is it not a nice prepossession or preju- dice like that in favour of the stability of the earth which he has so ably argued against? In fact, what positive evidence have we that species only vary within certain limits? . . . we have no proof how the varieties of dogs were produced. All varieties we know of are produced at birth, the offspring differing f rom the parent. This offspring propagates its kind. Who can declare that it shall not produce a variety, which process continued at intervals will account for all the facts? 6"

Not only did Lye]] believe that the amount of variation was strictly limited, but he also believed that it occurred over a brief period, after which no more changes took place no matter how long a time passed. Wallace thought that "'Mr. LyeU must be very perplexed to know this." 6v Lyell did concede

63. Ibid., p. 38. 64. Ibid., pp. 77-78; comma added. 65, Ibid., p. 76. 66. Ibid., pp. 39 -40 ; commas added. See also LyeH, Principles of

Geology, 4th ed., II, 435. 67. Wallace, "'Notebook," p. 42.

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BARBARA G. BEDDALL

that time might bring about any metamorphosis, but only ff such change were evidenced by a wholly new sense or organ. Wallace objected that "this would be taking a leap with a vengeance . We should have to get out of one class or order into another passing through m a n y thousand spec ies - - I f this is supposed to prove a change f rom one species to another, it can never be proved." 0s

Lyell opposed Lamarck 's theory of the t ransmutat ion of species, but Wallace defended his doctrine of their indefinite modifiability:

Many of Lamarck 's views arc quite untenable & it is easy to controvert them, but not so the simple question of a species being produced in time f rom a closely allied distinct species, which, however, may of course continue to exist as long or longer than the offshoot. Changes which we bring about artificially in short periods may have a tendency to revert to the parent stock, though this in animals is not proved. This is considered a grand test of a variety. But when the change has been produced by Nature during a long series of generations, as gradual as the changes of Geology, it by no means follows that they may not be permanent , & thus true species produced. ~°

Wallace was to re turn later to the "grand test of a variety"; it became the opening gun in the essay he sent to Darwin f rom Ternate in February 1858.

Lyell had made m a n y contributions to the tangle of questions surrounding the geographical distribution of animals. But a central p rob lem- - the "creation" of species- -muddled his conclusions, and Wallace put his finger on one of the inconsistencies to which this led :

LyeU occupies much space in shewing how the species which are c o m m o n to different & distant countries, might have been carried f rom one to the other by a variety of acci- dents. But this has never been felt to be a difficulty. The mat ter of wonder has always been that in distant countries of similar climate so m a n y should be di f ferent . This he gets over by special creation of the species each in one spot as they are wanted.

This is no doubt a very easy way of getting over it, but just as philosophical as to say that fossils of existing species

68. Ibid., p. 43; comma added. See also Lyell, Principles of Geology, 4th ed., II, 414.

69. Wallace, "'Notebook," pp. 44--45; some commas added. See also note 100.

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are remains of real animals while those which are not like any species now existing are special creations & not fossil animals at allJ °

Wallace turned again to the peculiarities of distribution to be found on the Galapagos Islands. Lyell's theories were inadequate to the task of explaining them, as Wallace pointed out :

In a small group of islands not very distant f rom the main land, like the Galapagos, we find animals & plants different f rom those of any other country but resembling those of the nearest land. I f they are special creations, why should they resemble those of the nearest land? Does not that fact point to an origin f rom that land? Again in these islands we find species peculiar to each island, & not one of them containing all the species found in the others as would be the case had one been peopled with new creations & the others left to become peopled by winds, currents, etc., f rom it. Here we must suppose special creations in each island of peculiar species though the islands are all exactly similar in structure, soil, & climate, & some of them within sight of each other, a work of supererogation one would suppose, as they must inevitably in time become peopled from each other, & contrary to what takes place e lsewhere- - l re land is peopled f rom England. It may be said it is a mystery which we cannot explain, but do we not thus make unnecessary systems and difficulties by supposing special creations contrary to the present course of nature? For we must conclude the course of nature in peopling islands in the ocean to be uni- form & that all islands distant f rom others should now be stocked with animals & plants equally peculiar.

But we know this not to be the case. Volcanic islands recently produced & coral islands far in the ocean contain stragglers f rom the nearest land & no other, nothing peculiarl Now we can hardly suppose that islands would be left for ages to become stocked in this manner , & then the new & peculiar creations be introduced just when they were not wanted. According to Mr. Lyelrs own arguments, they would hardly be able to hold their own against the previous occu- piers of the soil & there would have to be a special extermina- tion of them to make room for the new & peculiar species. We must therefore suppose that such islands as St. Helena & the Galapagos were stocked with their peculiar species

70. Ibid., pp. 45-46; see also Lye]l, Principles of Geology, 4th ed., III, 2 2 - 9 7 .

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immediately on their being raised f rom the ocean, & they would then have a chance of keeping out the new comers which might be blown accidentally on their shores. This supposition will certainly explain the present condition of those islands but it has the disadvantage of being contrary to the present order of nature, for none of the islands which we have any reason to believe have been formed, since a very late geological era, are inhabited by peculiar species. They generally have not one species peculiar to themselves. On the other hand, islands which are thus peculiarly inhabited, appear to be of a considerable antiquity [in a mar- ginal note, Wallace wrote, "this mus t be proved"]. A long succession of generations appears therefore to have been requisite, to produce those peculiar productions found nowhere else but allied to those of the nearest land. The change like every other change in nature was no doubt gradual, & the supposition that other species were successively produced closely allied to those previously existing, & that while this was going on, the original or some of the first formed species died out, exactly accords with the facts as we find them & the process of peopling new islands at the present day. 71

How far removed Lyell was f rom evolutionary theory is brought out at still another point. He thought that when cli- matic changes did cause the extinction of local species, any new inhabitants would be "perfectly dissimilar ha their forms, habits, and organization." 72 In other words, there would be a wholesale extinction followed by a renewal with altogether different species. Wallace pointed out, however, that the new species would more likely be modified forms of those previously existing: "'It would be an extraordinary thing if while the modifications of the surface took [place] by natural causes now in operation, & the extinction of species was the natural result of the same causes, yet the reproduction & introduction of new species required special acts of creation, or some process which does not present itself in the ordinary course of nature." (At this point in his '2qotebook," Wallace inserted the following note: "'Introduce this and disprove all Lyell's arguments first at the commencement of my last chapter"- -evidence that this was part of his proposed book.)Va

When environmental conditions changed, Lyell thought that new species already adapted to these new conditions would come in f rom surrounding areas before those in residence had

71. Wallace, "'Notebook," pp. 46-49; some commas added. 72. Lye]l, Principles of Geology, 4th ed., III, 154. 73. Wallace, "'Notebook," pp. 50-51.

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t ime to change ( i f they changed at all) . Wallace agreed tha t "if the change took place rapidly, the exact results LyeU pre- dicts migh t follow"; "but," he went on, "how the same results could follow f rom an excessively gradual change it is impossible to unders tand." 74 Lyell and L a m a r c k both agreed that plenty of t ime was available, bu t they differed on wha t would happen. Lyell believed that species could change, but only to a l imited degree and within the space of a few generations, Not only were they incapable of greater change, but such change would also be precluded by the immigra t ion of other species already adapted to the new environment . Lamarck , on the other hand , believed tha t species did change gradually through t ime in response to changes in the envi ronment , but his agency, the wills of the an imals themselves, was suspect. Whatever their deficiencies (pe rhaps because of t h e m ) , the beliefs of both these m e n were valuable s teppingstones along Wallace 's w a y .

In the mean t ime , Wallace was reading as widely as circumstances permi t ted: Edward Blyth on the classification of varieties, Richard Owen on the varieties of color in man , and Leopold yon Bueh on the flora of the Canary Islands. 75 Be- cause yon Buch's percept ive com m en t s also influenced Darwin, Wallace 's notes on them are worth quoting:

On continents the individuals of one kind of p lan t disperse themselves very far , and by the difference of stations of nour i shmen t & of soft produce varieties, which at such a distance not being crossed by other varieties & thus brought back to the pr imit ive type, become at length p e r m a n e n t & distinct species. Then if by chance in other directions they mee t with another variety equally changed in its march , the two have become very distinct species 8: are no longer sus- ceptible of in te rmixture . . . He then shews tha t p lants on the exposed peak of Teneriffe where they can mee t & cross do not fo rm varieties or species, while others such as Pyre- thrum & Cineraria l iving in sheltered valleys & low grounds often have closely allied species confined to one valley or one island. TM

74. Ibid., p. 52; c o m m a s added. See also Lyell , Principles of Geology, 4 t h ed., III , 161-163 .

75. C h r i s t i a n Leopold y o n B uch , Physicalische Beschreibung der Can- arisehen Inseln (Ber l in : K. Akade i n i e de r W i s s e n s c h a f t e n , 1825) . A n ear l ie r r epor t on t he f lora of t he C a n a r y I s l a n d s a p p e a r e d i n t he Abhand- lungen of t he Society ( 1 8 1 6 - 1 8 1 7 ) , 337 -384 .

76. Wa l l ace , " 'Notebook," p. 90.

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BARBARA G. BEDDALL

Darwin in his own "Notebook" says, 'Won Buch distinctly states that permanent varieties become species, pp. 147 -150 , - -no t being crossed with others." 77

Still another clue attracted Wallace's attention, and he asked:

What is the import of the doctrine of Morphology of plants? • . . For if s tamens & petals & carpels have been in every case independently created as such, it is absurd to say they are modifications or developments of any thing else, & the absurdity is still greater if that of which they are said to be the development came into existence after them. In that case all the beautiful facts of morphology are a delusion & a snare, as much so as fossils would be were they really not the remains of living things but chance imitations of them.

The natural inference of an unprejudiced person however would be that both are true records of the progress of the organic world. Nature seems to tell us that as organs are occasionally changed & modified now, in individual plants, we may learn how the actual changes have taken place in the species of plants. A key is offered us to a mystery we could otherwise never have laid open, why should we refuse to use it? 7s

Sometime during these musings, Wallace wrote out his "Note on the Theory of Permanent and Geographical Varieties," following up an interest expressed m a n y years earlier to Bates, and it was published early in 1858. Although he had been pondering a wide range of subjects, he limited the "Note" to showing the logical inconsistency in the suggestion that geographical varieties had permanent characters. I f varieties differ f rom species only in the minuteness of the permanent characters, then the difference between them is merely a quan- titative one and the dividing line becomes very difficult to dis- tinguish. The only qualitative difference that Wallace could discover was that of the permanence vs. the impermanence of variations. But this was no better, for some groups so formed were called special creations and others not. "Strange that such widely different origins produce such identical results." 79 If varieties are known to be produced by ordinary generation, why should species only slightly different be produced by special creation? "If there is no other character [than one of mere

77. Gavin de Beer, ed., " 'Darwin 's Notebooks on T r a n s m u t a t i o n of Species," Bull. Brit. Mus . (Nat . Hist . ) , Hist . Ser., 2 (1960) , 61. Da r win ' s first notebook w a s wr i t t en be tween Ju ly 1837 and Feb rua ry 1838.

78. Wallace, "'Notebook," pp. 97-100. 79. Wallace, "Note," p. 5888. See also Appendix , 3.

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W a l l a c e , D a r w i n a n d the T h e o r y of N a t u r a l Se l ec t ion

deg ree o f d i f fe rence] , t h a t f a c t is one of the s t r o n g e s t a rgu- m e n t s a g a i n s t t he i n d e p e n d e n t c r e a t i o n of species ." so

IV. A N D D A R W I N [Darwin] is now preparing for publication his great work on species and varieties, for which he has been collecting information twenty years. He may save me the trouble of writing the second part of my hypothesis by proving that there is no difference in nature between the origin of species and varieties, or he may give me trouble by arriving at another conclusion, but at all events his facts will be given me to work upon. Wallace to Bates, 4 January 1858, from Marcbant, Alfred Russel Wallace

E i g h t e e n h u n d r e d a n d f i f ty -e igh t was t he y e a r in w h i c h the c a r e e r s of W a l l a c e a n d D a r w i n col l ided. W a l l a c e h a d i n i t i a t e d the c o r r e s p o n d e n c e b e t w e e n t h e m w h e n he wro te h is f irst l e t t e r to D a r w i n in Oc tobe r 1856 [8]. sl W h y he wro te is n o t k n o w n ; p e r h a p s , as s u g g e s t e d ea r l i e r , h e h o p e d t h a t D a r w i n w o u l d be i n t e r e s t e d in h is s p e c u l a t i o n s on the G a l a p a g o s I s l a n d s puzzle. A l t h o u g h i t w a s W a l l a c e ' s t h i r d l e t t e r to D a r w i n t h a t b r o u g h t w i t h i t W a l l a c e ' s d i s cove ry of the t h e o r y of n a t u r a l se lec t ion , even th is f i rs t one m a y h a v e been d i squ ie t ing .

T h e c o n t e n t s of W a l l a c e ' s l e t t e r c a n be p a r t i a l l y s u r m i s e d f r o m D a r w i n ' s a n s w e r w r i t t e n in t he f o l l o w i n g M a y [11]: t he p a p e r in the A n n a l s , d o m e s t i c v e r s u s wi ld va r i e t i e s ( a c r u c i a l p o i n t in t he d e v e l o p m e n t of W a l l a c e ' s i d e a s ) , h y b r i d s te r i l i ty , a n d the ef fec ts of c l i m a t i c c h a n g e s . As to the p a p e r , D a r w i n a g r e e d "to the t r u t h of a l m o s t eve ry word" ; he h a d , in f ac t , a l r e a d y p o n d e r e d the s a m e p r o b l e m s - - c l a s s i f i c a t i o n , e x t i n c t i o n a n d c r ea t i on , a n d r u d i m e n t a r y o r g a n s - - a s c a n be seen f r o m h i s own n o t e b o o k s a n d h is e s s a y w r i t t e n in 1844. T h e r e w a s less a g r e e m e n t , h o w e v e r , on o t h e r sub jec t s .

D a r w i n l a t e r wro te Lye l l ( J u n e 1858) t h a t he a n d W a l l a c e d i f fe red on ly in " tha t I w a s l ed to m y v iews f r o m w h a t a r t i f i c ia l s e l ec t ion h a s done fo r d o m e s t i c a n i m a l s [23]." But t hey b o t h m a d e use of d o m e s t i c a t e d a n i m a l s , a l t h o u g h fo r d i f f e r en t e n d s : D a r w i n to show t h a t v a r i a t i o n e x i s t e d a n d cou ld be c h a n n e l e d , W a l l a c e to show t h a t t he u s u a l de f in i t i on of spec ies , b a s e d on d o m e s t i c a t e d a n i m a l s , d id n o t a p p l y to wi ld a n i m a l s . ( A n d , as W a l l a c e wro te m a n y y e a r s l a t e r , "it h a s a l w a y s b e e n c o n s i d e r e d a w e a k n e s s in D a r w i n ' s w o r k t h a t he b a s e d h is theory , pr i - m a r i l y , on the e v i d e n c e of v a r i a t i o n in d o m e s t i c a t e d a n i m a l s a n d c u l t i v a t e d p l a n t s . " ) s2 N o r d id W a l l a c e a n d D a r w i n eve r a g r e e on the t h o r n y p r o b l e m of the s te r i l i ty of hyb r id s .

80. Ibid., p. 5888. 81. See also Appendix, 69 and notes 43 and 162. 82. Alfred Russel Wallace, Darwinism: A n Exposi t ion of the Theory of

Natural Selection, w i th Some of its Applications (New York: Humboldt, 1889) , p. iv.

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BARBARA G. BEDDALL

There was also a difference of opinion on the effect of cli- matic changes. In 1844 Darwin had written "that probably such changes of external conditions would, f rom acting on the re- productive system, cause the organization of the beings most affected to become, as under domestication, plastic." 83 He had since modified his opinion, and in his letter to Wallace he agreed "on the little effect of 'clLmatal condit ions. '" Wallace, however, never did ascribe any such direct influence to cli- matic conditions in causing variations. Nevertheless, in spite of some disagreement it was plain, as Darwin said, that they had thought m u c h alike.

Darwin received Wallace's first letter late in April 1857. He was then deep in writing "his great work on species and varieties," having started almost exactly a year earlier at the urging of Lye]l, with the strong support of the botanist Joseph Hooker [5, 6, 7]. As Lyell wrote not long afterward, "Part of the MS. of [Darwin's] projected work was read to Dr. Hooker as early as 1844, and some of the principal results were com- municated to me on several occasions. Dr. Hooker and I had repeatedly urged him to publish without delay." 84 It has recently been proposed, on evidence from Lyell's own notebooks, that it was actually Wallace's paper in the A n n a l s that prompted Lyell to reconsider the subject of species and to prod Darwin to publication, and, fur thermore, that it was at this time that Darwin explained his theory to Lyell. s5

Darwin had protested to LyeU that he did not like writing for priority, but had admitted that he "certainly should be vexed if any one were to publish my doctrines before me [5]"; and to his cousin, W. D. Fox, he had confided that he wished he "could set less value on the bauble fame [10]." By 31 March 1857, he had completed six chapters of the projected work, and he may already have been at work on the seventh, com- paring species and varieties, when Wallace's first letter arrived in late April. The coincidence in their thinking may have put Darwin on his guard.

On 1 May 1857, Darwin wrote in answer (to some speculations by Wal lace?) that he had been working for nearly twenty years "on the question how and in what way do species and varieties differ f rom each other [11]." He did not elaborate on

83. Darwin and Wallace, Evolution, p. 119. Text differs slightly from that in joint papers; see note 133.

84. Charles Lyell, The Geological Evidences of the Ant iqui ty of Man, ~ i t h Remarks on Theories of the Origin of Species by Variation, 1st ed. (London: Murray, 1863), p. 408.

85. McKirmey, "Wallace and Natural Selection," pp. 350-352.

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the point, however, and left it up in the air. (Wallace had not yet written his own "Note" on the subject, and it is not known whether Darwin ever did see it.)

As was his usual practice, Darwin included several requests for information, asking Wallace, among other things, to let h im know "'ff you should, after receiving this, stumble on any curious domestic breed" of poultry. 86 Perhaps as a result of this inquiry, Wallace made a few notes in his "Notebook," some time before November 1857, on unusual breeds of ducks, s7 This small point will be brought up later in connection with Wallace's recollections of the beginning of his correspondence with Darwin.

More important at the momen t is Darwin's protestation that "it is really impossible to explain my views (in the compass of a letter) on the causes and means of variation in a state of nature." Was it really impossible?

Two years earlier, in April 1855, Darwin had begun to cor- respond with the noted American botanist Asa Gray, whom he had once met briefly at Kew, and Gray had been providing him with m a n y valuable comments. On 20 July 1857, not long after receiving his first letter f rom Wallace, Darwin wrote again to Gray, saying, "I should like to tell you (and I do not think I have) how I view my work," condensing into a few sentences the gist of his theory [12]. s8 A recent biographer of Gray suggests that Darwin felt it necessary to let Gray in on his secret to ensure the cont inuance of this useful correspondence, but this does not seem to be a necessary assumption. 89 More to the point, Darwin seemed rather to fear that Gray would despise h im and his crotchets, and in his next letter (5 September) he confessed that he had been afraid that Gray might think him "worth no more notice or assistance" because of his unorthodox views [14]. 9o (Even in March 1860 Darwin considered Gray to be a convert to his views only "to some extent.") 91

Along with this answer to Gray, Darwin sent a copy of an outline that he had made of his theory, "as you seem interested in the subject." This is the famous extract published the fol-

86. The paragraph containing this request is published in Marchant, Wallace, p. 108, but it was omitted ~rom the Darwin, Life and Letters, I, 454. See also note 87.

87. Wallace, "Notebook," p. 91. See also Appendix, 69, Marchant, Wallace, p. 86, and notes 161 and 162.

88. See also note 146. 89. A. Hunter Dupree, Asa Gray, 1810-1888 (Cambridge: Harvard Uni-

versity Press, 1959) , p. 244. 90. See also note 146. 91. Darwin, Life and Letters, II, 87.

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lowing year as part of the Darwin-Wal lace joint papers. Dar- win concluded his letter to Gray with the following request:

You will, perhaps, think it paltry in me, when I ask you not to ment ion my doctrine; the reason is, if any one, like the author of the "Vestiges,' were to hear of them, he might easily work them in, and then I should have to quote f rom a work perhaps despised by naturalists, and this would greatly injure any chance of my views being received by those alone whose opinion I value [14]. 92

Why did Darwin send this s tatement to Gray, who responded that it was "grievously hypothetical [15, 17]" rather than to Wallace (who was hardly at this time, however, a scientific peer) , who would have understood? Was it Wallace rather than Chambers of whom he was afraid? If so, a recent outline of his views, including the important addition on divergence, mailed to the eminent American botanist might protect his ideas. And, whatever his intentions may have been, this indeed was the result.

Finally, on 29 November 1857, Darwin thanked Gray for his help, remarking that "every criticism from a good m a n is of value to me [17]." s3 But the subject was apparently not pursued, and this concludes the series of letters about Darwin's theory.

Wallace was "much gratified" by Darwin's first letter, as he wrote Bates in January 1858, but he was no wiser than before ( h a d he asked?) about Darwin's opinion for or against a "difference in nature between the origin of species and varieties [19]." 04 In any case, he had pretty well made up his own mind already.

Judging f rom Darwin's next answer, Wallace in reply once again brought up his paper in the A n n a l s and also remarked on various problems relating to the geographical distribution of animals. But the only piece on Wallace's side of this early correspondence that is still in existence is a snippet f rom his answering letter of 27 September 1857, on the breeding habits of jaguars and his plan for his book [16]. 95 For the rest, Wal-

92. This paragraph has been variously interpreted; see Dupree, Asa Gray, p. 246 and Himmelfarb, Darwin, p. 207.

93. See note 146. 94. See also Loren Eiseley, Darwin's Century and the Men W h o Discov-

ered I t (Garden City, N.Y.: Doubleday, 1958), p. 291, where he considers this correspondence "stimulating'" to Wallace. But Wallace received no such positive statement as was given to Gray.

95. Unpublished fragment in the Cambridge University Library; s e e also note 60. For Darwin's methods of filing his letters, see notes 140--143.

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lace's opinions mus t be conjectured f rom Darwin's letters to him.

By the time Darwin answered Wallace's second letter on 22 December 1857, he had almost completed the ninth chapter of his book, this one on hybridism. He assured Wallace that his paper had not gone unnoticed, Lyell and the zoologist Edward Blyth having called it to his attention; none of these men, however, had bothered to write Wallace about it. Darwin passed up his second and last opportunity to tell Wallace about his theory, saying that "though agreeing with you on your conclusions in that paper, I believe I go m u c h fur ther than you; but it is too long a subject to enter on my speculative no- tions [18]." It seems unlikely, however, that Wallace received this letter before the end of February 1858; most of the letters exchanged by the two men took anywhere f rom three to six months to reach the recipient.

In the meant ime, in his January 1858 letter to Bates, Wal- lace told him of his plans, explaining that "I have prepared the plan and written portions of an extensive work embracing the subject in all its bearings and endeavouring to prove what in the paper [in the Annals] I only indicated." He did not seem to be overly concerned about Darwin's conclusion on the origin of species and varieties, remarking that Darwin might save him the trouble of proving that there was no difference in their nature [19].

It has been claimed that Wallace, "duly warned off" by Dax- win's first letter, had nevertheless continued his own work on the subjectP s It should be objected that this was no private preserve of Darwin's. Many people were interested in it, as Wallace was well aware. Darwin gave Wallace no hint of a solution to the problem; why should he not continue with what had been a consuming interest for m a n y years? Bates himself, on his re turn home, wrote that one of their purposes in going to the Amazon was to "gather facts, as Mr. Wallace expressed it in one of his letters, ' towards solving the problem of the origin of species,' a subject on which we had conversed and corres- ponded m u c h together." 97 If Darwin had been working on the problem for twenty years, Wallace had been working on it for at least ten, the major difference being that Darwin had long had a theory against which he was collecting facts, while Wallace was still actively searching for one.

By now Wallace had concluded that there was no qualitative difference in the origin of species and varieties and that they

96. Himmelfarb, Darwin, p. 236. 97. Bates, The Naturalist, p. iii; see also Appendix, 4 and note 17.

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BARBARA G. BEDDALL

were formed gradually by natural means with a relationship in space and time to what had gone before, but without any necessary progression. Only a short distance separated him from his goal, a "theory of the origin of species.'"

In February 1858 the final step was taken. Wallace was ly- Lug ill in Ternate in the Moluccas, mulling as usual over his problem, when at last he found the key to the puzzle: the theory of natural selection. Much has been made of the fact that the solution came to him while he was ill but, as he im- patiently remarked later, he "had no idea whatever of "dying,' - - a s it was not a serious illness [69]." Indeed, he had frequently been sick, and sometimes m u c h sicker. Although not iden- tified at the time, the disease has recently been referred to as malaria.g8

As soon as he was able, Wallace wrote out his theory with the t ire, "'On the Tendency of Varieties to depart indefinitely f rom the Original Type," and sent it to Darwin with the request that it be shown to Lyell. 9° Had he chosen any other recipient, the results would have been different. Gray's response to Darwin's disclosure, for instance, had been that it was too hypothetical. But to Darwin, Wallace's paper was little short of a calamity.

In his "Note," Wallace had been struggling with the supposed distinction between permanent varieties and permanent ly invariable species. Now he saw his way clear. Belief in variation within strict limits and reversion to the original type was based on domestic animals and then applied to wild animals. But domestic animals are artificial and unable to maintain themselves without the help of man. If allowed to go wild, they must either re turn to something similar to the original type or become extinct. Wild animals, on the other hand, must be adapted to their environment. Their every faculty is con- stanfly exercised in keeping themselves alive. Any improve- ment in organization is quickly taken advantage of, and a new variety is thus superior to its predecessor. Being superior, it "cou ld n o t return to the original form; for that form is an inferior one, and could never compete with it for existence." 10o Quite the opposite is true of domestic animals, which are inferior f rom the point of view of mainta ining themselves in the wild. Thus varieties in nature tend to depart indefinitely f rom the original type.

98. Julian s. Huxley and H. B. D. Kettlewell, Charles Darwin and his World (New York: Viking Press, 1965), p. 74.

99. Alfred Russel Wallace, "'On the Tendency of Varieties to depart indefinitely from the Original Type," ]. Linn. Soc. London (Zooi.), 3 (1858), 53-62. 100. Ibid., p. 58. See also note 69.

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Several years earlier, Wallace had found puzzling Lyell's belief that the balance of species was preserved by plants, insects, mammals , and birds adapted to the purpose, and he had worried over this problem in his '¢Notebook":

This phrase is utterly without meaning. Some species are very rare, others very a b u n d a n t - - w h e r e is the balance? Some species exclude all others in particular t r ac t s - -where is the b a l a n c e - - W h e n the locust devastates vast regions, & causes the death of animals & man, what is the meaning of saying the balance is preserved . . . To h u m a n apprehension there is no balance but a struggle in which one often exterminates the o t h e r - - W h e n animals and plants become extinct, where is the balance. If any state can be imagined proving a want of balance, then a balance may perhaps be admitted, but what state is that? T M

Now, two years later, Wallace had his answer. In the "struggle for existence," m a n y individuals must perish annually. Even the least fecund species would soon overrun the earth ff its numbers went unchecked. Those that survive are the ones best adapted to obtain food and to withstand their enemies and the seasonal changes in the weather; those that die are the young, the old, and the sick. In applying this not only to individuals but also to species, Wallace thought he had an answer to why some species are rare while others are abundant. Besides this, the animal population of a country cannot increase materially ff conditions remain the same. In the "struggle for existence," therefore, those individuals and species best adapted to main ta in themselves survive.

It is well known that Wallace and Darwin both read Mal- thus' famous Essay on the Principle of Population, Wallace before leaving Leicester and Darwin after re turning home f rom his voyage. Malthus' interest was in the moral perfectibility of man, and it was in this light that he discussed the checks to population g rowth- - famine , war, disease, and v i ce - -bu t Wal- lace and Darwin were both impressed by the implications of these checks. The exact extent of Malthus' influence is hard to determine and has been the subject of m u c h debate, but at the least his forceful presentation was widely known.

Darwin read Malthus in the fall of 1838. There is no indication of this in the first edition of his Voyage of the Beagle,

101. Wal lace , "'Notebook," pp. 4 9 - 5 0 ; s o m e c o m m a s added. See also Lyell, Principles of Geology, 4th ed., III, 98-120, where he discusses the "checks and counter-checks which nature h a s appo inted to preserve the balance of p o w e r a m o n g s t species."

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however, because though published in 1839, the writing of it was largely finished by June 1837. Darwin had tried vainly to account for the extinction of so many peculiar South American forms that were related to the present inhabitants, concluding only that :

On such grounds it does not seem a necessary conclusion, that the extinction of species, more than their creation, should exclusively depend on the nature (altered by physical changes) of their country. All that at present can be said with certainty, is that, as with the individual, so with the species, the hour of life has run its course, and is spent. 1°2

This comment was greatly enlarged for the second edition (1845) and clearly shows the influence of Malthus:

Nevertheless, if we consider the subject under another point of view, it will appear less perplexing. We do not steadily bear in mind, how profoundly ignorant we are of the conditions of existence of every animal; nor do we always remember, that some check is constantly preventing the too rapid increase of every organized being left in a state of nature. The supply of food, on an average, remains constant; yet the tendency in every animal to increase by propagation is geometrical; and its surprising effects have nowhere been more astonishingly shown, than in the case of the European animals r u n wild during the last few c e n -

turies in America. Every animal in a state of nature regu- laxly breeds; yet in a species long established, any g r e a t in -

crease in numbers is obviously impossible, and must be checked by some means. We are, nevertheless, seldom able with certainty to tell in any given species, at what period of life, or at what period of the year, or whether only at long intervals, the check falls; or, again, what is the precise nature of the check. Hence probably it is, that we feel so little surprise at one, of two species closely allied in habits, being rare and the other abundant in the same district; or, again, that one should be abundant in one district, and another, filling the same place in the economy of nature, should be abundant in a neighboring district, differing very little in its conditions. If asked how this is, one immediately replies that it is determined by some slight difference in climate, food, or the number of enemies: yet how rarely, ff ever, we can point out the precise cause and manner of action of the checkl We are, therefore, driven to the conclusion, that

102, Darwin, Voyage of the Beagle (1839), p. '212.

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causes general ly quite inapprec iab le by us, de te rmine whe ther a given species shal l be abundan t or scan ty in numbers .

• . . If then, as appears probable , species first become ra re and then ext inc t ff the too rap id increase of every species, even the most favoured, is s teadi ly checked, as we mus t admit , though how and when it is ha rd to s a y - - a n d if we see, wi thout the smal les t surprise, though unable to ass ign the precise reason, one species abundan t and another closely-all ied species ra re in the same d i s t r i c t - - w h y should we feel such great a s ton i shment at the ra r i ty being car r ied a step fu r the r to ext inc t ion? 10a

Wal lace did not have Mal thus with him, but he did have Darwin. F a m i l i a r as he a l ready was wi th the Mal thus ian arguments , he mus t have not iced their inc lus ion in the second edi t ion of the Voyage of the Beagle. These pr inciples , first suggested in 1798, were now used by Wal lace and Darwin in a r a the r different context. But Mal thus alone was not enough. As Wal lace jus t ly pointed out m a n y years later , "along with Mal- thus I had read, and been even more deeply impressed by, Sir Char les Lyell 's immor t a l 'Pr inciples of Geology."" 104

The final and mos t impor t an t poin t of Wal lace ' s paper was the appl ica t ion of the concepts he had developed to varieties. Even sl ight var ia t ions would have an effect, e i ther favorable or unfavorable , and under changed phys ica l condit ions a better- adapted var ie ty migh t survive its pa ren t species. (This would be t rue only of wild variet ies , however, for domest ica ted an ima l s turned wild are ra re ly able to m a i n t a i n themselves . ) This process repea ted would lead to "progression and continued divergence." lo~ At las t Wal lace h a d a m e c h a n i s m tha t ex- p la ined the knot ty p rob lem of progress ion which had so baffled Lyell, and the equal ly puzzl ing problem of divergence; and it could also replace Lamarck ' s general ly discredi ted theory tha t progress ive changes were due to the wills of the an imals themselves.

Darwin ' s approach was a l i t t le different. Because Wal lace is the focus of this study, discussion here will be l imi ted to the two selections f rom Darwin ' s wri t ings tha t became pa r t of the jo int papers in 1858. The first was an ex t rac t f rom an essay wr i t ten in 1844. loo Darwin had r e m a r k e d there tha t De Can-

103. Ibid., (1845), pp. 174--176. 104. Linnean Society of London, The Darwin-Wallace Celebration, p.

118. 105. Wallace, "'On the Tendency," p. 59. See also note 37. 106. Charles Darwin, "Extract from an Unpublished Work on Species,

by C. Darwin, Esq., consisting of a portion of a Chapter entitled 'On the

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dolle's war of nature "is the doctrine of Malthus applied in most cases with ten-fold force" lo7 (he was to use a similar phrase in the Origin of Species). 1°8 Under these circumstances, slight variations caused directly by changed physical conditions might lead to small improvements in organisms through the natural selection of those better adapted. 1°9 Darwin also discussed sexual selection, another subject on which he and Wallace were to differ strongly.

The brief abstract enclosed with the letter to Asa Gray sum- marized Darwin's conclusions at this later date, 1857.11° Se- lection of variations by man is recognized in the propagation of domestic animals. Physical conditions are known to have changed over a great length of time. These changed conditions have caused variations to occur in organisms in a state of nature, al though Darwin was no longer certain that this was the sole cause. Finally, there is a natural power comparable to that of man which selects those that survive in the struggle for life, a power which Darwin called Natural Selection. In a country undergoing changes, these slight variations would be selected and gradually accumulated, leading to new varieties adapted to the new conditions.

At the end, Darwin added a paragraph on his principle of divergence. The solution to this p rob lem--" tha t the varying offspring of each species will try (only few will succeed) to seize on as m a n y and as diverse places in the economy of nature as possible" 111--had not occured to him until 1852,112 "long after I had come to Down,"l la and so was not yet a part of a formal statement of his views. He knew from Wal- lace's paper in the Annals that he too was trying to solve this problem.

Varia t ion of Organic Beings in a state of Na tu re ; on the Na tu r a l Means of Selection; on the Compar i son of Domest ic Races and t rue Species,"" ]. Linn. Soc. London (Zool.), 3 (1858) , 46-50. See also note 133.

107. Ibid., p. 47. 108. Charles Darwin , On the Origin of Species by Means of Natural

Selection, or the Preservation of favoured Races in the Struggle for Life (London: Murray , 1859; facsLrn.ile repr int , Cambridge; Harva rd Univers i ty Press, 1964), p. 63.

109. See also note 83. 110. Charles Darwin , " 'Abstract of a Letter f r o m C. Darwin , Esq., to

Prof. Asa Gray, Boston, U.S., dated Down, September 5th, 1857," ]. Linn. Soc. London (Zool.), 3 (1858) , 50-53. See also Appendix, 14 and notes 88-93.

111. Ibid., pp. 52-53. See also note 37. 112. Gavin de Beer, Charles Darwin: a Scientific Biography (Garden

City, N.Y.: Doubleday, 1964), p. 140. 113. Darwin , Life and Letters, I, 69.

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V. THE JOINT PAPERS

I was not aware before that your father had been so distressed--or rather disturbed--by my sending him my essay from Ternate . . .

Wallace to Francis Darwin, 20 November 1887

"June 14th Pigeons ( i n t e r rup t ed ) . " 114 Such is the cryptic note in Darwin ' s "Journal" ind ica t ing the receipt of Wal lace ' s paper on 18 June 1858. By this t ime he h a d comple ted twelve chapte rs of his book and was at work on the th i r teenth , and his distress at being thus fores ta l led can easi ly be imagined. The story of this most d r ama t i c m o m e n t has of ten been re- counted. But, as Hooker observed, the detai ls s tem ent i re ly f rom some of the let ters wri t ten at the t ime by Darwin to h imsel f and to Lyell [22-26, 29, 33-36]; 115 all other docu- m e n t a r y evidence, the let ters f rom Wal lace , Lyell, and Hooker to Darwin, as well as the manusc r i p t of Wal lace ' s paper , has d isappeared. The facts , consequent ly , are difficult to de termine , and the c i rcumstances have been var iously interpreted.

There is no way of ascer ta in ing exact ly why Wal lace sent his paper to Darwin; cer ta in ly he could not have an t ic ipa ted the result . Wi th no h in t f rom Darwin, he could not have real ized tha t he had s tumbled onto the very founda t ion of Dar- win 's work. Darwin, on the other hand , mus t have had a fa i r not ion of Wal lace ' s progress f rom his publ ished papers and pe rhaps a warn ing of this d isas ter f rom his let ters , the first of which m a y indeed have prec ip i ta ted the sketch sent to Gray.

Wal lace , who had been away f rom England for m a n y years , was a se l f -educated collector f rom outside the regula r establish- ment , and he had few persona l contacts among the scientific elite. By d in t of his own efforts, he had finally es tabl ished a cor respondence with one of its members , Charles Darwin. He m a y well have hoped for some useful cr i t ic ism f rom Darwin and Lyell, bu t he could ha rd ly have expected to be ca tapu l t ed into the f ront ranks himself .

F rom the evidence, i t appears tha t Wal lace sent his paper to Darwin with the request tha t it be fo rwarded to Lyell, "should he th ink it sufficiently novel and in teres t ing [20]." 116 Darwin ' s own le t ter to Lyell, wr i t ten on 18 June, said only

114. Gavin de Beer, ed., "'Darwin's Journal," Bull. Brit. Mus. (Nat , Hist.), Hist. Ser., 2 (1959), 14. 14 June 1858 was the clay on which Darwin began this chapter.

115. See also note 139. 116. Charles Lyell and Joseph D. Hooker, "'[Letter communicating the

Darwin-Wallace Papers to the Linnean Society]," 30 June 1858, ]. L inn . Soe. London (Zool.), 3 (1858), 46.

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BARBARA G. BEDDALL

that Wallace "has to-day sent me the enclosed, and asked me to forward it to you [22]." Lyell's recollection later was that the paper had been brought to him by Hooker, who then suggested some sort of joint publ icat ion. 117 In Leonard Huxley's biography of Hooker, it is stated that Darwin "had first confided Wallace's unexpected letter" to him, and that he first suggested joint publication and also the getting of LyeU's opinion. 11s Whatever the precise details, Hooker, Darwin's "most intimate friend," accepted the larger share of the re- sponsibility for what happened.

Darwin was now in a di lemma: "He does not say he wishes me to publish, but I shall, of course, at once write and offer to send to any journal. So all my originality, whatever it may amount to, will be smashed [22]." In fact, Darwin did start a letter to Wallace giving up his claims to priority, but he never finished it, for his old friends, Hooker and LyeU, suggested a compromise. If Darwin's and Wallace's roles had been reversed, as they could have been, Wallace would have had no one to help him resolve the difficulty.

A week later, on 25 and 26 June, Darwin wrote Lyell again. A joint publication of some kind had apparently been proposed, al though "Wallace says nothing about publication" and Dar- win was properly hesitant about the proprieties involved. He mentioned a copy of his sketch sent to Gray "about a year ago . . . (owing to correspondence on several points) ." ("Cor- respondence" here probably means actual letter-writing rather than agreement, because the sketch "gives most imperfectly only the means of change," a subject new to Gray.) He also enclosed the letter f rom Wallace and requested that Lyell get Hooker's opinion; some of the confusion on this point has already been mentioned [20, 9.3, 24, 26]. By 29 June, Darwin had their answers. He was to send to Hooker both Wallace's paper (which Lyell must already have returned to h im) and his own sketch sent to Asa Gray. Although they had apparently not asked for it, he also sent along his much more extensive sketch (230 pages) , written in 1844, to show by notes in Hook- er's handwri t ing that he had read it [25, 26]. 119

Darwin's letters were usually written with an intensity of feeling lacking in the more pedestrian efforts of m a n y of his contemporaries. At this momen t they were shrill with anxiety and doubt. But Darwin was being sorely tried. He was troubled not only by Wallace's communicat ion but also by severe illness

117. Lyell , Principles of Geology, 10th ed., II, 278. 118. L e o n a r d Hux l ey , ed., Life and Letters of Sir ]oseph Dalton Hooker

( L o n d o n : M u r r a y , 1918) , II, 465. 119. See also no te 84.

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in his family. An infant son died of scarlet fever on 28 June, and a daughter was seriously ill with diphtheria.

Finding a suitable fo rum for the papers at this time of year would ordinarily have presented still another problem. But both Hooker and Lyell, as Fellows of the Linnean Society of London, knew that one had unexpectedly become available. Robert Brown, a leading botanist, former President and then Council Member of the Society, had died on 10 June. Out of respect, the last meet ing of the old session, held on 17 June, was ad- journed before the reading of the scheduled papers. But Brown had to be replaced on the Council within three months, and, as the new session would not start until November, it was decided to hold the extra meeting on 1 July. Without consulting anyone else, Hooker and Lyell t ransmitted their selections to the Secretary of the Society on 30 June, to be read by him the next day. As Hooker said in 1908 :

It cannot fail to be noticed that all these inter-communica- tions between Mr. Darwin, Sir Charles Lyell, and myself were conducted by correspondence, no two of us having met in the interval between June the 18th and July the 1st, when I met Lyell at the evening meeting of the Linnean Society; and no fourth individual had any cognisance of our proceedings. 120

This was not an occasion of "mutual nobility," 121 nor was it "a m o n u m e n t to the natural generosity of both the great biologists," 122 as is so often claimed. It was clearly not mutual because Wallace's paper was read without his knowledge or consent, and he knew nothing about it until October. Nor does it seem to have been particularly noble. However just Darwin's claims to priority, he was a gainer, not a loser, f rom the decision. Wallace had no opportunity to be either noble or generous.

Wallace, "a gent leman attached to the study of Natural His- tory," was not unknown to the Linnean Society. The first two volumes of the Society's 1ournal (Zoology) were largely taken up with descriptions (writ ten by others) of the collections of insects he had sent home f rom Singapore, Malacca, and Sara- wak. The Society was later to publish some of Wallace's most

120. L i n n e a n Society of L o n d o n , The Darwin-Wallace Celebration, p. 15. T h i s a n d t he i n f o r m a t i o n of B r o w n (Ibid., pp. 1 4 - 1 5 ) were omi t t ed i n M a r c h a n t , Wallace, p. 98.

121. Eiseley, Darwin's Century, p. 292. 122. J u l i a n S. H u x l e y , " 'Alfred R u s s e l W a l l ace , " Dictionary of National

Biography, S u p p l e m e n t 1912-1921 ( L o n d o n : Oxfo rd U n i v e r s i t y Press, 1927) , p. 547.

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i m p o r t a n t papers , bu t he h imse l f was no t elected a Fel low u n t i l 1872. Darwin , on the other h a n d , was a l ready a Fellow, and h a d ju s t b e e n elected to the Counc i l i n May.

Some th i r ty people, pe rhaps more , out of a m e m b e r s h i p of over four h u n d r e d were p r e sen t at the mee t ing . Al though some of the i r n a m e s m i g h t no t be recognized today, fu l ly ha l f of those l is ted in the Minu te s as a t t e n d i n g have ra ted not ices i n Br i t a in ' s r e n o w n e d Dictionary of National Biography, cer- t a in ly a d i s t i ngu i shed audience . 123

The e v e n i n g was a fu l l one. The bus ine s s of the m e e t i n g was t r ansac ted , and t h e n c a m e the r e a d i n g of the papers , the jo in t papers by D a r w i n a n d Wa l l ace fol lowed by five of the six previous ly scheduled for 17 June . The jo in t papers were i n t roduced by a le t ter f rom Lyell a n d Hooker exp la inJng w h a t they had done and why. The first se lect ion was f r o m Darwin ' s essay of 1844, a n "Extrac t f r o m a n u n p u b l i s h e d Work on Species, by C. Da rwin , Esq., cons i s t ing of a por t ion of a Chap- ter en t i t l ed 'On the Var i a t ion of Organ ic Beings in a s tate of N a t u r e ; on the N a t u r a l Means of Select ion; on the C o m p a r i s o n of Domest ic Races a n d t rue Spec ies . ' " D a r w i n appended to the pub l i shed vers ion a no te t ha t "this MS. work was neve r i n t e n d e d for pub l i ca t ion , and therefore was no t wr i t t en wi th care." This was no has t i ly w r i t t e n s u m m a r y , however , for D a r w i n had h a d it copied a n d b o u n d , a nd he h a d also left i n s t r u c t i o n s to his wife for its p u b l i c a t i o n i n the even t of his p r e m a t u r e death. 124

Secondly c a m e the "Abst rac t of a Let ter f r o m C. D a r w i n ,

123. Listed in the minutes of the meeting, in the Society's Darwin- Wallace Celebration, pp. 81-86. The following can be found in the D.N.B.: Baird, William (1803-1872), zoologist; Ball, John (1818-1889), botanist; Baly, William (1814-1861), physician (visitor); Bell, Thomas (1792- 1880), dental surgeon and zoologist (President); Bennett, John Joseph (1801-1876), botanist (Sole Secretary), not listed as present, although he presumably read the papers; Bentham, George (1800--1884), botanist; Burchell, William John (1782?-1863), explorer and naturalist; Busk, George (1807-1886), physician and scientist (Under- (Zoological) Secre- tary), not listed as present, although Hooker later recalled that he was; Carpenter, William Benjamin (1813-1885), naturalist and physician; Currey, Frederick (1819-1881), mycologist; Fitton, William (1780-1861), physician and geologist; Henfrey, Arthur (1819--1859), botanist; Hooker, Joseph Dalton (1817-1911), botanist; Lyell, Charles (1797-1875), geologist; Salter, John William (1820-1869), geologist; Seeman, Berthold Carl (1825--1871), botanist and traveler; Ward, Nathaniel Bagshaw (1791-1868), botanist and physician. Others may have been present as the list ends with "etc., etc."

124. The date is incorrectly given as 1842 in the published papers. Darwin further confused the issue by using the date 1839 in a letter to Wallace; see Appendix, 43. For details of Darwin's plans, see Himmelfarb, Darwin, pp. 190-191.

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Esq., to Prof. Asa Gray, Boston, U.S., dated Down, September 5th, 1857," the outline of his theory of na tu ra l selection tha t Dar- win had sent to Gray. Francis Darwin was later of the opinion that the reason for the inclusion of this note was the discussion of the "principle of divergence," an impor tan t par t of Darwin 's theory not included in the 1844 essay. 125 Hooker was aware tha t Darwin gave divergence equal p rominence with na tu ra l selection as "the keystone of m y book [21]," a l though he apparently did not unders tand the connect ion between them. In his own essay, "On the Flora of Australia," publ ished at a lmost the same t ime as Darwin 's Origin of Species, Hooker wrote tha t "the tendency of varieties, both in na ture and under culti- vation, when fu r the r varying, is ra ther to depar t more and more widely f rom the original type than to rever t to it." z26 Darwin objected tha t this was "without selection doubtful." z27

Third and last was Wallace 's paper , "On the Tendency of Varieties to depar t indefinitely f r o m the Original Type." But no note was added to indicate tha t Wallace had not wri t ten for publicat ion either.

I t was Hooker 's recollection twenty-eight years la ter (whe the r accurate ly or no; certainly public react ion to the publicat ion of the papers was a lmost n i l ) tha t the interest was intense, although there was no discussion. Thom as Bell, the President, though a personal f r iend of Darwin's , "was hostile to the end of his life." Nei ther of the Secretaries, George Busk and John J. Bennett , said anything, nor did the botanis t George Bent- h a m [67]. ( T h o m a s Huxley, la ter to be '~Darwin's bulldog," was not present , not being elected a Fellow unti l December 1858.)

Ben tham m a y have been silent, but his feelings were those of "severe pa in and disappointment ." His was the only one of the six previously scheduled papers tha t was not read. Many years la ter he recalled the events in a letter to Francis Darwin:

On the day tha t his [C. Darwin's] celebrated paper was read at the L innean Society, July 1st, 1858, a long paper of mine had been set down for reading, in which, in comment ing on the British Flora, I had collected a n u m b e r of observa- tions and fac ts i l lustrat ing wha t I then believed to be a fixity in species, however difficult it migh t be to assign their limits, and showing a tendency of abnormal fo rms produced by cult ivation or otherwise, to wi thdraw within those orig-

125. D a r w i n a n d W a l l ace , Evolution, p. 34. See also no te 37. 126. Quo ted i n F r a n c i s D a r w i n , ed., More Letters of Charles Darwin: a

Record of his W o r k in a Series of Hitherto Unpubl ished Letters ( N e w York: App le ton , 1903) , I, 134.

127. Ibid.

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BAIIBABA G. BEDDALL

inal limits when left to themselves. Most fortunately my paper had to give way to Mr. Darwin's and when once that was read, I felt bound to defer mine for reconsideration; I began to entertain doubts on the subject, and on the appearance of the 'Origin of Species,' I was forced, however re- luctantly, to give up my long-cherished convictions, the results of much labour and study, and I cancelled all that part of my paper which urged original fixity, and published only portions of the remainder in another form, chiefly in the 'Natural History Review.' I have since acknowledged on various occasions my full adoption of Mr. Darwin's views, and chiefly in my Presidential Address of 1863 [to the Lin- nean Society], and in my thirteenth and last address, issued in the form of a report to the British Association at its meeting at Belfast in 1874 [66].

Bentham had not given in easily; even in his Presidential Address in 1862 he was still struggling against the new doctrine :

I do not refer to those speculations on the origin of species, which have excited so m u c h controversy; for the discussion of that question, when considered only with reference to the comparative plausibility of opposite hypothesis, is beyond the province of our Society. Attempts to bring it forward at our meetings were very judiciously checked by my predecessor [Bell] in this Chair, and I certainly should be sorry to see our time taken up by theoretical arguments not accompanied by the disclosure of new facts or observa- tions.12s

Bell was a dental surgeon and zoologist, but "as a naturalist he was more at home in his study than in the field, and he made few original contributions of special value to zoology. As a writer, his chief merit is that of agreeable compilation." 120 In his own Presidential Address in 1859, he dismissed the joint papers altogether:

The year which has passed . . . has not been unproductive in contributions of interest and value, in those sciences to which we are professedly more particularly addicted, as well as in every other walk of scientific research. It has not, indeed, been marked by any of those striking discoveries which at once revolutionize, so to speak, the depar tment of

128. Proc. Linn. Soc. London (1 November 1860-19 June 1862), p. lxxxi.

129. G. Y. Bettany, "Thomas Bell," D.N.B., II, 175.

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science on which they bear; it is only at remote intervals that we can reasonably expect any sudden and brilliant in- novation which shall produce a marked and permanent impress on the character of any branch of knowledge, or confer a lasting and important service on mankind. A Bacon or a Newton, an Oersted or a Wheatstone, a Davy or a Daguerre, is an occasional phenomenon, whose existence and career seem to be especially appointed by Providence, for the purpose of effecting some great important change in the condition or pursuits of man, 130

Darwin's immediate reaction to WaUace's paper was, hu- manly enough, great distress, followed by relief at the solution worked out by Hooker and Lyell. "But," he wrote to Hooker, "in truth it shames me that you should have lost time on a mere point of priority." ( I f it was not a point of priority, what, indeed, was the hu r ry? ) Although he was not yet clear on exactly which of his papers had been read at the Linnean Society meeting, he was glad that Hooker planned to write Wallace about the affair, "as it would quite exonerate me [29]."

Darwin was "more than satisfied" when he discovered what had been done: the strictly chronological ( and alphabetical) a r rangement of the papers meant that his preceded Wallace's (as does his name in the references to the published papers) , when "I had thought that your letter and mine to Asa Gray were to be only an appendix to WaUace's paper [33]." lal

On 13 July Hooker and Darwin both sent letters to Wallace explaining the turn of events [31, 32]. Unhappily, these letters are missing, al though Wallace carefully saved most of Dar- win's letters to h im (and it is f rom them that we know of Wallace's early letters to Darwin) . A few days later Darwin also thanked Lyell for his part, again expressing himself as "far more than satisfied [34]." He was pleased to have the public backing of men like Lyell and Hooker. It was only after some years of struggle, however, that LyeU became a "convert," while Hooker, al though convinced of the action of natural selection, nevertheless vacillated on its importance. Even Darwin hedged as time went on. Of the four men, Wallace was to be the most steadfast, mainta in ing to the end his belief in "the overwhelm-

130. J. L inn . Soc. L o n d o n (Zool.), 4 (1859) , vii i- ix. 131. The collective title for the joint pape r s and the accompany ing

letter is entered as fol lows: Charles D a r w i n and Alfred Russel Wallace, "On the Tendency of Species to f o r m Varieties; and on the Pe rpe tua t ion of Varieties and Species by N a t u r a l Means of Selection," J. L inn . Soc. L o n d o n (Zool.), 3 (1858) , 45-62.

305

BARBARA G. BEDDALL

Lug impor tance of Na tu ra l Selection over all other agencies in the product ion of new species." 132

By 20 July Darwin had received the proof sheets, and he re turned them to Hooker the next day, with "only a few corrections in the style [35]." In fact , judging f rom the original texts as published in Evolution by Natural Selection and the Life and Letters of Charles Darwin, several hundred changes were made in both the 1844 sketch and the letter to Asa Gray, not only in the punc tua t ion and wording but even in whole phrases. 1~3 To have been scrupulously fair, it would seem that no changes should have been made at all. Darwin complained to Hooker that he had not been wri t ing for publicat ion [36], and a note to this effect was inserted in the Journal, as already ment ioned.

Wallace did not have these opportunities. I t is not known who read the proof of his paper nor wha t became of his manuscript. In later reprintLugs he added phrases in footnotes that he would have inserted in the text, but he made no corrections then because of the historical impor tance of the document . 134 The joint papers were published on 20 August 1858 in No. 9 of the third volume of the LLunean Society's ]ournal (Zoology).

The sudden confronta t ion with Darwin threw Wallace into the limelight. But he had not s tumbled upon the theory of na tura l selection by accident; he was to be nei ther a hanger-on nor a blind follower of Darwin's , and he was to make m a n y valuable and original contr ibutions of his own to evolut ionary theory. His interests often paral leled those of Darwin, but his point of view frequent ly differed. Because of Darwin 's illness and isolation at Down, their long and frui t ful association is recorded in a correspondence tha t continued unti l Darwin 's death. 135

Wallace 's first in t imat ion of what had happened came when

132- Wallace, Darwinism, p. iv. 133. Compare the text of the jo in t papers w i th the 1844 excerpt in

D a r w i n and Wallace, Evolution, pp. 116-121, and the letter to Asa Gray in Darwin , Life and Letters, I, 479-482. See also notes 83, 138, 150, and 158.

134. Wallace added two footnotes in the first r ep r in t ing in h is Contri- butions to the Theory of Natural Selection (London: Macmil lan , 1870); he omit ted these and added a third in the second repr in t ing in Natural Selection and Tropical Nature (London: Macmil lan , 1891), p. 27n, wi th the c o m m e n t tha t "'it m u s t be r e m e m b e r e d tha t the wri ter had no opportu- n i ty of correct ing the proofs of this paper. '"

135. The complete ex tan t cor respondence is in Marchan t , Wallace, pp. 107-262. The texts differ s o m e w h a t f r o m those in Darwin , Life and Letters, besides the publ ica t ion of sect ions omit ted there. See also note 158.

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he received the letters f rom Hooker and Darwin in early Octo- ber 1858. At last he had aroused a t ten t ion--a t ten t ion that assured him, as he wrote his mother, "the acquaintance and assistance of these eminent men" on his re turn home. What- ever else Darwin's letter conveyed to Wallace, it was not the pain and anguish of those two final weeks in June. Wallace, unaware of the flurry he had aroused, told his mother that he was "highly gratified . . . I sent Mr. Darwin an essay on a subject on which he is now writing a great work. He showed it to Dr. Hooker and Sir C. LyeU, who thought so highly of it that they immediately read it before the Linnean Society [37]."

And to an old boyhood friend, George Silk, Wallace crowed: "if you have any acquaintance who is a fellow of the Linnean Society, borrow the Journal of Proceedings for August last, and in the last article you will find some of my latest lucubra- tions, and also some complimentary remarks thereon by Sir Charles Lyell and Dr. Hooker, which (as I know neither of them) I am a little proud of [42]." Almost thirty years passed before Wallace learned some of the details of Darwin's side of the story.

Darwin, who received Wallace's answering letter in January 1859, was "extremely m u c h pleased" with it; he had been "anxious to hear what your impression would be." He incorrectly referred to his own extracts as having been written in 1839, when in fact the first was written in 1844 and the second, the letter to Asa Gray, had been written in 1857, only a year and a half before [38, 43].

By November 1858 Wallace had received a copy of the ]ournal containing the papers and could read for himself Dar- win's "distinct and tangible idea." The tenor of Wallace's comments can be judged f rom Darwin's answer in the following April. Darwin agreed that Wallace was right in "that I came to the conclusion that selection was the principle of change from study of domesticated productions; and then, reading Malthus, I saw at once how to apply this principle. Geographi- cal distribution and geographical relations of extinct to recent inhabitants of South America first led me to the subject: especially the case of the Galapagos Islands [41, 45]." (But the letter in which this latter s tatement appeared was not published until 1903. )

Again Darwin expressed his admiration for the manner in which Wallace had taken the publication of the papers, Actu- ally, it is hard to imagine what else Wallace could have done. Whatever reservations he might have had (and there is no indication that he had any) , he was the forestaller, not the

307

BARBARA G. BEDDALL

forestalled, and he had at last achieved recognition f rom "two of the most eminent naturalists in England," a remarkable accomplishment for a self-educated collector.

Darwin had set to work again almost immediately after the publication of the papers, making an "Abstract of Species book," and by November 1859 he could write Wallace that his publisher was sending him a copy of the Origin of Species, adding that "I do not think your share in the theory will be overlooked by the real judges, as Hooker, Lye]l, Asa Gray, etc. [47]." 136 Darwin had hoped that Wallace would feel that his Linnean paper was "fairly noticed" in the short Introduct ion and added that he would "allude" to the paper in the Annals in the body of the work [45]-- this he did, but without giving either its title or the date (he always thought of this work as an abstract of the one he intended to write, and consequently he never gave proper references). 13~ But Darwin has never been accused of being overgenerous in his credits, particularly in the "Historical Sketch" later appended to the Origin of Species, and it may be that Wallace was too modest in his claims.

Wallace wrote to congratulate Darwin on his book in Febru- ary 1860 [49], and Darwin sent the letter on to Lyell, remarking on "how admirably free [he was] f rom envy or jealousy. He must be a good fellow [53]." This letter is among those missing, and so for Wallace's opinions it is necessary to turn to letters written to Bates in December 1860 and to his brother-in-law, Thomas Sims, in the following April.

Perhaps better than anyone else, Wallace could appreciate the extraordinary amount of work involved, and he wrote Bates that he was thankful it had not been left for him to do. "Mr. Darwin has created a new science and a new philosophy, and I believe that never has such a complete illustration of a new branch of h u m a n knowledge been due to the labours and researches of a single m a n [55]."

The letter to Sims is both a defense of Darwin and an explanation. Sims had apparently taken exception to Darwin's references to Wallace, and Wallace reproved him :

You quite misunders tand Mr. D.'s s tatement in the preface and his sentiments. I have, of course, been in correspondence with him since I first sent h im my little essay. His conduct has been most liberal and disinterested. I think anyone who reads the Linnean Society papers and his book will

136. Darwin had sent copies of the joint papers to Wallace the preceding year; see Appendix, 40.

137. Darwin, Origin of Species, 1st ed., pp. 1-2, 355.

3 0 8


see it. I do back him up in his whole round of conclusions and look upon him as the Newton of Natural History.

Sims objected not only to the contents of the Origin of Spe- cies but even to its title. After explaining that Darwin had originally given h im a different title (presumably in the letter of 13 July 1858), Wallace went on to give his own judgment of Darwin's accomplishment : "It is the vast chaos of facts, which are explicable and fall into beautiful order on the one theory, which are inexplicable and remain a chaos on the other, which I think must ult imately force Darwin's views on any and every reflecting mind." The letter is worth reading carefully because, in trying to convince Sims, Wallace showed himself a strong and articulate champion of these views [56].

It is a pity that Wallace was not at home to take an active part in the controversy over the Origin of Species. He would have enjoyed the dispute which Darwin found so distasteful. But by the time he returned to England in 1862 the first heat of the battle was past. Darwin, at any rate, was pleased with Wallace's enthusiasm, and he thanked him for his "too high approbation of my book . . . most persons would in your position have felt bitter envy and jealousy [54]." 13s

But the documentat ion of this famous episode leaves something to be desired. There are gaps in the record that not only are rarely noticed but also are being gradually obliterated in the frequent retelling of this episode. Most people agree with Marchant that Darwin's letters tell the whole story, but they form only part of the evidence. Missing are all the letters (except for one f r agment ) that Wallace sent to Darwin f rom the Malay Archipelago, Wallace's manuscript , the letters written to Darwin by Hooker and Lyell during those hectic weeks in June 1858, and the pert inent letters f rom Asa Gray. 1~9

The evidence on the disposition of the letters to Darwin is contradictory. Francis Darwin later recalled that his father "made a rule of keeping all letters that he received; this was a habit which he learnt f rom his father, and which he said had been of great use to him." 14o But, in describing his own work in compiling the Life and Letters of Charles Darwin, he gave a different account :

138. "Bi t ter e n v y " c h a n g e d to " s o m e e n v y " i n D a r w i n , Life and Letters; see also no t e 150.

139. T h e m i s s i n g le t t e r s a re nos . 8, 13, 15, 20, 27, 28, 30, 38, 39, 41, 44, 48, 49, m a r k e d w i t h a n a s t e r i sk i n t he Append ix . Le t te r s nos . 31 a n d 32 f r o m D a r w i n a n d Hooker to W a l l a c e a re also m i s s i n g .

140. D a r w i n , Life and Letters, I, 97.

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BARBARA G. BEDDALL

Of letters addressed to my father I have not made much use. It was his custom to file all letters received, and when his slender stock of files ("spits" as he called them) was ex- hausted, he would burn the letters of several years, in order that he might make use of the liberated "spits." This process, carried on for years, destroyed nearly all letters received before 1862. After that date he was persuaded to keep the more interesting letters, and these are preserved in an ac- cessible form. 141

Many of the letters received before 1862 do exist, however, as can easily be checked in the published collections of Darwin's letters and in the list of unpublished material in the Cambridge University Library. 142

Another version of what happened to the letters is given in a biography of Hooker:

In one of his letters Darwin makes special ment ion of pre- serving his friend's [i.e. Hooker's] letters. The answers to scientific questions are detached and placed among the memoranda of that subject; the other parts are put away among his general correspondence, so that it would only be a matter of half an hour to rearrange them in case of need. In spite of his care, however, a large number of the earlier letters f rom Hooker have disappeared wholly or in part.14a

The one snippet f rom Wallace's letter of 27 September 1857 shows that Darwin must have followed this practice with his letters also, at least at first [16]. But Darwin was meticulous (indeed, it is f rom his own care in answering Wallace that the dates of Wallace's letters to h im can be so easily determined) . It seems surprising that all the material relating to the most dramatic (not to say t raumat ic) moment in his life should disappear.

The dating of Darwin's own letters presents still another problem. As his son observed:

He rarely dated his letters, so that but for the Diary [Jour- nal] it would have been all but impossible to unravel the history of his books. It has also enabled me to assign dates to m a n y letters which would otherwise have been shorn of half their value, 144 [and]

141. Ibid., xviii-xix. 142. Cambridge University Library, H a n d l i s t o f D a r w i n Papers at the

Univers i t y Library , Cambr idge (Cambridge: Cambridge University Press, 1960).

143. L. Huxley, Li f e and Let ters . . . Hooher, I, 436. 144. Darwin, Li f e a n d Let ters , I, xvi.i.i.

310

W a l l a c e , D a r w i n a n d t h e T h e o r y o f N a t u r a l S e l e c t i o n

Mr . D a r w i n , w h o w a s c a r e f u l i n o t h e r t h i n g s , g e n e r a l l y o m i t t e d t h e d a t e i n f a m i l i a r c o r r e s p o n d e n c e , a n d i t i s o f t e n o n l y b y t r e a t i n g a l e t t e r a s a d e t e c t i v e s t u d i e s a c r i m e t h a t w e c a n m a k e s u r e o f i t s d a t e . F o r t u n a t e l y , h o w e v e r , S i r J o s e p h H o o k e r a n d o t h e r s o f D a r w i n ' s c o r r e s p o n d e n t s w e r e a c c u s t o m e d to a d d t h e d a t e o n w h i c h t h e l e t t e r s w e r e r e - ceived.145

S o m e o f t h e c r u c i a l l e t t e r s o n t h e t h e o r y o f n a t u r a l s e l e c t i o n f r o m D a r w i n to A s a G r a y h a v e r e c e n t l y b e e n r e d a t e d b y D u - p r e e i n a d i f f e r e n t c o n n e c t i o n , a r e d a t i n g t h a t c a n b e s u p - p o r t e d b y a n e x a m i n a t i o n o f t h e t e x t s . T h e s e l e t t e r s h a v e a p p e a r e d i n d i f f e r e n t c o l l e c t i o n s o f D a r w i n ' s l e t t e r s a n d h a v e b e e n d a t e d 1 8 5 6 [12] , 1 8 5 7 [14] , a n d 1 8 5 9 [17] , r e s p e c t i v e l y , b u t t h e y s e e m a l l to h a v e b e e n w r i t t e n i n 1 8 5 7 , a f t e r D a r w i n r e c e i v e d W a l l a c e ' s f i r s t l e t t e r . P u t t o g e t h e r a n d r e a d i n se- q u e n c e , t h e y s e e m to f o r m a n a t u r a l u n i t , l e n d i n g s u p p o r t to t h e t h e o r y t h a t t h e r e v e l a t i o n to G r a y w a s i n d u c e d b y W a l l a c e . 145

VI. E P I L O G U E

I feel m u c h sat isfact ion in hav ing thus aided in br inging about the publ ica t ion of this celebrated book, and wi th the ample recognit ion by Darwin h imsel f of my independen t discovery of "na tu ra l selection."

Wallace, Natural Selection and Tropical Nature

T o a l a r g e e x t e n t t h e w o r l d h a s a c c e p t e d a t f a c e v a l u e b o t h W a l l a c e ' s a n d D a r w i n ' s r a t h e r p i o u s r e c o l l e c t i o n s o f t h e i r c l o s e

145. Darwin, More Letters, I, x. 146. There are two ways to date the letters to Gray. One is by the

sequence in the Darwin-Gray correspondence, and the other is f rom in te rna l evidence. Both indicate tha t no. 12 was wri t ten in 1857. In particular, Darwin refers to a chapter on the cont inuous dis t r ibut ion of species, which "Hooker kindly read . . . over." According to Darwin 's " 'Journal," he f inished this section on 13 October 1856, hav ing asked Hooker on 13 July 1856 if he would read i t for ]aim (Darwin, More Letters, I, 95). See also note 88.

Al though there is no quest ion about the year (1857) of no. 14, Darwin did not date the copy he kept, and he thought he had sent i t i n October (Darwin, Life and Letters, I, 477n) . The date is given as October in the letter of t r ansmi t t a l to the L i nnean Society but as 5 September 1857 on the letter itself. See also note 90.

Number 17 was probably pu t in 1859 because i t concerns na tu ra l selection. But Darwin 's book was publ ished on 24 November, and Gray could not have received a copy, read it, and given Darwin his opinion in 5 days. Again, the sequence of letters shows tha t i t belongs in 1857. I t is followed by two others tha t have also been dated 1859 bu t which should be dated 1858, one on 21 February (Darwin, Life and Letters, I, 463-464) and another on 4 April (ibid., 510). The la t ter refers to a jus t f inished chapter on ins t inc t that , according to Darwin 's "Journal ," was f inished on 9 March 1858. See also note 93.

3 1 1

B A R B A R A G . B E D D A L L

bu t n o t a lways unruf f led r e l a t i onsh ip . The v e t of V ic to r i an p r o p r i e t y t h r o u g h w h i c h they c a m e to v iew e a c h o t h e r ha s , howeve r , o b s c u r e d some of t he i r m o r e h u m a n r e a c t i o n s to w h a t m u s t a t t imes h a v e b e e n a t r y i n g e n t a n g l e m e n t .

Pub l i c r e c o g n i t i o n by D a r w i n of W a l l a c e ' s a c h i e v e m e n t c a n be d iv ided in to two p a r t s , be fo re a n d a f t e r p u b l i c a t i o n of the L i f e a n d L e t t e r s o f C h a r l e s D a r w i n i n 1887, the f irst p a r t c o n s i s t i n g m a i n l y of r e m a r k s in the "His to r i ca l Ske tch" ap- p e n d e d to the O r i g i n o f S p e c i e s . This ske t ch ( w h i c h h a s been ca l l ed " the m o s t u n r e l i a b l e a c c o u n t t h a t eve r wi l l be wr i t - t e n " ) 147 was a d d e d to t he t h i r d ed i t i on in 1861 a n d e x p a n d e d s o m e w h a t fo r the f o u r t h in 1866.148 W a l l a c e a n d the L i n n e a n p a p e r s were g iven one sen t ence , whi l e W a l l a c e ' s p a p e r in the A n n a l s was n o t m e n t i o n e d a t a l l a n d Lye l l was o m i t t e d alto- ge ther . A m o n g the n a m e s a d d e d in 1866 were those of P a t r i c k M a t t h e w a n d W i l l i a m Wel l s , bo th c o n t e n d e r s fo r t he t i t le of d i s cove re r of the t heo ry of n a t u r a l se lec t ion .

M a t t h e w h a d w r i t t e n on n a t u r a l se lec t ion in 1831, a n d he p r e s s e d h is c l a i m in t he G a r d n e r s " C h r o n i c l e i n 1860.149 Dar - w i n p u b l i c l y a c k n o w l e d g e d h is a n t i c i p a t i o n of the theory , w r i t i n g to W a l l a c e t h a t 'the g ives m o s t c l e a r l y b u t ve ry br ie f ly • . . our v i ew of N a t u r a l Se lec t ion [51, 52, 54]. ''15° W e l l s ' c l a i m was m a d e in 1865 b y a "Mr. Rowley , of the U n i t e d Sta tes ," fo r " a n a c c o u n t of a f e m a l e . . . p a r t of w h o s e sk in r e s e m b l e s t ha t of a negro , " a p a p e r f i rs t r e a d be fo re the Roya l Socie ty in 1813 a n d p u b l i s h e d p o s t h u m o u s l y in 1818.151 W a l l a c e was s u r p r i s e d " ' that i t shou ld h a v e s t r u c k no one t h a t [his sugges- t ion] w a s a g r e a t p r i n c i p l e of u n i v e r s a l a p p l i c a t i o n in Na- tu re [60] I"

Shor t ly be fo re th is , W a l l a c e h a d m o d e s t l y r e f e r r e d to "Mr. D a r w i n ' s c e l e b r a t e d t h e o r y of ' N a t u r a l S e l e c t i o n . ' " 152 D a r w i n h a d d e m u r r e d , b u t W a l l a c e h a d i n s i s t e d t h a t h e w o u l d " a lw a ys

147. C. D. Darlington, "'The Origin of Darwinism," Sci. Amer . , 200 (May 1959), 61.

148. Darwin, Origin of Species , 3rd ed., pp. v-xi; 4th ed., pp. xiii-xxi. 149. Patrick Matthew, N a v a l T i m b e r and Arbor icu l ture (Edinburgh:

Longman, 1831); "Nature's Law of Selection," Gard. Chron. Agr icul . Gaz. , 20 (7 April 1860), pp. 312-313.

150. Darwin, Li fe and Let ters , II, 95-96n. The remark on Patrick Matthew was omitted from Darwin, Li fe a n d Let ters; see also note 138.

151. William Wells, T w o Essays: one u p o n Single V i s ion w i t h two Eyes; the o ther on Dew. A let ter to . . . Lord K e n y o n and an A c c o u n t o f a Fema le . . . pa r t o f w h o s e S k i n r e semb l e s t ha t o f a Negro (London: A. Constable, 1818).

152. Alfred Russel Wallace, "'The Origin of Human Races and the Antiquity of Man deduced from Natural Selection," ]. An thropo l . Soc. London , 2 (1864), clix.

312


main ta in it to be actually yours, and yours only [57, 58]." And yet in 1869 Wallace's reply to a request f rom the anthropolo- gist A. B. Meyer for his recollections of his part in the theory was brusque. After stating that he "was led to it by Malthus' views on population applied to animals," his terse account ended with the remark that his paper "was printed without my knowledge, and of course without any correction of proofs. I should, of course, like this fact to be stated [63]."

Wallace objected to being classed with Matthew and Wells, "who made no fur ther use of that principle, and failed to see its wide and immensely important applications," and he published a collection of his papers in 1870 to make the extent of his own contribution clear. 15a But he did not then, nor did he ever, claim that he had worked out the theory in the detail that Darwin had. Darwin thanked Wallace in glowing terms for the kind words in his preface, having missed altogether the point about Matthew and Wells [65].

Lyell, in the meant ime, thought Darwin had given short shrift not only to Wallace and himself, but also to Lamarck. He was astonished to find no ment ion of Wallace's paper in the Annals in Darwin's "Historical Sketch" [61], and he discussed this "next important effort to determine the manner in which new species may have originated" in some detail in the new edition of his famous Principles of Geology, published in 1867-1868. T M He had already credited Wallace with thinking out, "independently for himself, one of the most novel and important of Mr. Darwin's theories," in discussing the joint papers in his Antiquity of Man. ~55

And in his Principles of Geology, he now reprinted in its entirety his original summary of Lamarck, not to protest against his theory of the t ransmutat ion of species as before, but to "show how nearly the opinions taught by him at the commencement of this century resembled those now in vogue." ~58

As for himself, Lyell wrote in a letter to Ernst Haeckel in November 1868 that he was obliged to h im "for pointing out [in his History of Creation] how clearly I advocated a law of continuity even in the organic world, so far as possible without adopting Lamarck 's theory of t ransmutat ion . . . I had certainly prepared the way in this country . . . for the reception of Darwin's gradual and insensible evolution of species [62]," and Huxley later agreed with him. T M In 1870, apparently in

153. Wallace, Contributions, p. iv. 154. Lyell, Principles of Geoloyy, 10th ed., II, 276---281. 155. Lyell, Geological Evidences, 1st ed., pp. 408---409. 156. LyeU, Principles of Geology, 10th ed., II, 246n. 157. Darwin, Life and Letters, I, 543-544.

313

BARBARA G. BEDDALL

r e f e r e n c e to the p r e f a c e to W a l l a c e ' s c o m p i l a t i o n of h i s p a p e r s , Lyel l a g a i n s u p p o r t e d W a l l a c e , w r i t i n g h i m t h a t "it is h i g h t ime this m o d e s t a s s e r t i o n of you r c l a i m s as a n i n d e p e n d e n t o r i g i n a t o r of N a t u r a l Se lec t ion shou ld be p u b l i s h e d [64]."

N o t u n t i l the p u b l i c a t i o n of the Li fe and Le t t ers o f Charles D a r w i n i n 1887, five y e a r s a f t e r D a r w i n ' s d e a t h , d id W a l l a c e r ece ive ful l p u b l i c r e c o g n i t i o n for h is p a r t in the t heo ry of n a t u r a l se lec t ion . T h e l e t t e r s f r o m D a r w i n to Hooker , Lyel l , a n d W a l l a c e h i m s e l f were t h e r e fo r a l l to r e a d ( a l t h o u g h nei- t he r the c o m p l e t e t ex t s n o r al l of the ea r ly l e t t e r s f r o m D a r w i n to W a l l a c e we re i n c l u d e d 15s); a n d as f u r t h e r p r o o f of Wa l - l ace ' s ro le in i n d u c i n g D a r w i n to p u b l i s h the Origin o f Species , t he re w a s also D a r w i n ' s " A u t o b i o g r a p h y . " F o r the f irst t ime , W a l l a c e l e a r n e d some of t he de t a i l s of those long-ago events . He was s u r p r i s e d to f ind t h a t D a r w i n ' ~ a d b e e n so d i s t r e s s e d - - or r a t h e r d i s t u r b e d " by h is e s say , a n d he wro te apo loge t i ca l l y to F r a n c i s D a r w i n t h a t he h a d a l w a y s fe l t t ha t he h a d r ece ived too m u c h c r ed i t "'for m y m e r e ske tch of a t heo ry [68]."

D a r w i n ' s a t t i t u d e h a d a lso so f tened . N e a r l y t w e n t y y e a r s a f t e r the even t , in a n a u t o b i o g r a p h i c a l ske t ch w r i t t e n m a i n l y for the eyes of h is ch i l d r en , he cou ld s ay t h a t he " ca r ed ve ry l i t t le w h e t h e r m e n a t t r i b u t e d m o s t o r i g i n a l i t y to m e or Wa l - lace ." 159 He h a d w o r r i e d w h e t h e r W a l l a c e w o u l d cons ide r the who le p r o c e e d i n g jus t i f i ab le ( a b o u t w h i c h b o t h he a n d H o o k e r s e e m e d to have some l i n g e r i n g d o u b t s ) , n o t t h e n knowing '~how g e n e r o u s a n d nob le was h is d i spos i t ion . " 100 He m e n t i o n e d once a g a i n t h a t h is o w n p a r t s of the j o i n t p a p e r s h a d no t b e e n i n t e n d e d for p u b l i c a t i o n , wh i l e W a l l a c e ' s was a m o d e l of c la r i ty . H e n e v e r r e c o g n i z e d in a n y w a y t ha t W a l - l ace ' s was h a s t i l y wr i t t en , t ha t i t h a d no t b e e n i n t e n d e d fo r p u b l i c a t i o n e i the r , or t h a t W a l l a c e h a d h a d no c h a n c e to proof- r e a d it, b u t h e d id c red i t W a l l a c e w i th g iv ing h i m the i m p e t u s t h a t p r o d u c e d the Origin o f Species .

N o w t h a t W a l l a c e ' s pos i t i on was secure , he b e g a n to em- b r o i d e r h is own reco l lec t ions . At the t ime he wro te h is essay , n e a r l y t h i r t y y e a r s be fo re , he h a d h a d no i d e a e i t he r of i t s i m p o r t a n c e or of i t s i m p a c t on D a r w i n , a n d the e a r l i e s t re-

158. o f the first eight extant letters from Darwin to Wallace, six are included in Darwin, Life and Letters (nos. 11, 18, 43, 46, 47, and 54), but only one (47) is complete. The seventh (45) is in Darwin, More Letters, and the eighth (50) is in Marchant, Wallace. Marehant includes the complete extant text of all of them, but his book was not published until 1916. The texts also differ slightly from those in Darwin, Life and Letters. See also notes 3, 87, 135, 138 -143 , 146, 150, and 171.

159. Darwin, Life and Letters, I, 71. 160. Ibid., 69.

314

Wal lace , D a r w i n and the T h e o r y of N a t u r a l Se lec t ion

ques t fo r h is m e m o r i e s h a d c o m e f r o m Meye r in 1869. But n o w the c i r c u m s t a n c e s of t h a t w e e k so long ago a s s u m e d a n e w in teres t . A l f r ed N e w t o n , o rn i tho log i s t and zoologist , wro t e i n q u i r i n g fo r de ta i l s to i n c o r p o r a t e in his r e v i e w of the Li f e and Le t t e r s o f Charles Darwin , and W a l l a c e obl ig ingly responded.

But the l e t t e r to N e w t o n [69], w r i t t e n in 1887, c o n t a i n s a n u m b e r of ques t i onab le s t a t emen t s . W a l l a c e was n o w uncer - t a in w h e t h e r he h a d e v e n r e a d D a r w i n ' s Voyage o f the Beagle at the t ime, w h e n he h a d in f a c t r e a d bo th ed i t ions and h a d the s econd one w i t h h i m in the Ma lay Arch ipe l ago ; s ca t t e r ed r e f e r e n c e s to D a r w i n in h is o w n ear ly pub l i shed works also show tha t he h a d r e a d h i m wi th s o m e care . He t h o u g h t he h a d s t a r t ed the c o r r e s p o n d e n c e over some p e c u l i a r va r i e t i e s of ducks . H o w e v e r , the le t te r i t se l f shows tha t he was con- su l t ing the Li fe and Le t t e r s o f Charles D a r w i n to r e f r e s h h is m e m o r y , and the p a r a g r a p h c o n t a i n i n g the r e q u e s t fo r "any cu r ious b reed" in D a r w i n ' s first l e t te r to h i m was om i t t ed there . E v i d e n c e f r o m W a l l a c e ' s "Notebook" also ind ica tes tha t D a r w i n b r o u g h t the sub jec t up first [11]. T M Not ices in the A t h e n a e u m t ha t D a r w i n was i n t e r e s t ed i n species and va r i e t i e s s e e m improbab le . 162 W a l l a c e aga in pa id h is r e spec t s to Mal- thus , f u r t h e r e n s h r i n i n g h i m in the a n n a l s of science. A n d f inal ly, he r e f e r r e d to a "hot fit" of i n t e r m i t t e n t f eve r , a l t h o u g h he l a t e r sa id the idea h a d c o m e to h i m d u r i n g a "cold fit."

W a l l a c e h a d r e t u r n e d on ly a f e w m o n t h s be fo re f r o m a speak ing tour of the U n i t e d States. I n t e r e s t i n g l y enough , he h a d m e t Asa Gray d u r i n g a m o n t h - l o n g s tay in Boston, and Gray h a d inv i t ed h i m to a t t end a m e e t i n g of the C a m b r i d g e Scient i f ic Club. There , Gray showed his c o r r e s p o n d e n c e wi th D a r w i n be fo re the p u b l i c a t i o n of the Origin o f Species ,

161. See also notes 86 and 87. 162. No such notices located. Although it is possible that Wallace

heard of Darwin's interest through his agent, Stevens (who had been a member with Darwin since 1837 of the Entomological Society), and he could have used them as an opening for his first letter in 1856 (8), this still would not account for the range of subjects he apparently discussed. Darwin answered: "By your letter and even still more by your paper in the Annals . . . ,'" indicating that the letter itself must have contained some related remarks (11).

Sydney Smith has suggested (personal communication) that two letters from Darwin to W. B. Tegetmeier, dated 21 Nov. and 29 Nov. 1857, show that it was about the second date that Darwin received some poultry specimens collected by Wallace. Since Wallace received Darwin's letter (11) with this request in July (see 19), the shipment could have been in response to that request. This would leave unaffected the first letter Wallace wrote to Darwin in October 1856 (8), and the reason for it would then remain an open question.

315

BARBARA G. BEDDALL

and Wal lace "related what led h im to his theory of N a t u r a l Selection . . . The wri t ings of Spencer, Vestiges of Creation, Lamarck? , but pa r t i cu la r ly of Mal thus on popula t ion suggested his own view." 103 This also seems inaccura t e : Wal lace omitted Lyell and added Spencer, whose First Principles he read af ter his r e tu rn f rom the Malay Archipelago, in September 1862.184

At any rate , pe rhaps hea r tened by the publ ic recogni t ion he had now received, Wal lace expanded his series of lectures into a book, publ i shed in 1889, to which he gave the title, Darwinism: An Exposition of the Theory of Natural Selection. To h im the terms were synonymous. Fur the rmore , he was still convinced of the pr ime impor tance of na tu ra l selection, although Darwin had s taged a g radua l re t rea t f rom this position.

Next , in the in t roductory note to a chapte r in a new com- pi la t ion of his papers , Natural Selection and Tropical Nature, Wal lace added fu r the r detai ls of tha t now never-to-be-forgotten week, even to the t empera tu re outside. Again he consul ted the Life and Letters of Charles Darwin, and another probable error crept into the story. F rom the dates on which the first two Darwin let ters were wri t ten, Wal lace p re sumed tha t he had received them before sending his paper to Darwin, and he quoted f rom them both. I t is unlikely, however, tha t the second let ter could have reached h im so quickly. But he was now satisfied, as can be seen f rom his concluding remark , "with the ample recogni t ion by Darwin himself ." ~5

Wal lace gave an even more deta i led account in his own autobiography, wri t ten nea r ly fifty years after the event, adding, among other things, that he had asked Darwin to show his paper to Lyell "who had thought so h ighly of my fo rmer paper ." This aga in p resumes that Wal lace had a l ready received Darwin ' s second letter, which is doubtful . This frequent ly quoted-from account should be t rea ted with caution. 166

In thank ing Wal lace for a copy of his Life, Hooker, now eighty-eight years old, r e m a r k e d tha t "your ci ta t ion of my letters and their contents are like d reams to me; but to tell you the t ruth, I am gett ing dull of memory as well as of hear ing , and wha t is worse, in read ing : wha t goes in at one eye goes out at the o t h e r " - - p e r h a p s an honest eva lua t ion tha t could be appl ied to most octogenar ians , inc luding Wallace. 167

163. Cambridge Scientific Club, "'Record Book," 17 November 1886, MS, Harvard University Archives; quoted with the permission of Harvard University. See also Dupree, Asa Gray, pp. 380, 473 n52.

164. Marchant, WaUace, p. 122. 165. Wallace, Natural Selection, pp. 20-21. 166. Wallace, My Life, I, 357-363. 167. Marchant, Wallace, p. 332.

316


On 1 July 1908, the Linnean Society held a jubilee to com- memora te the reading of the joint papers fifty years before. It seems extraordinary that two of the protagonists, Wallace and Hooker, were still alive and able to participate. (LyeU had died in 1875 and Darwin in 1882.) Wallace, the first of the two to speak, repeated his story with some new elaborations, helping to perpetuate the (perhaps fictitious) report that his paper had come to Darwin ' l ike a thunderbol t f rom a cloud- less sky," but his account was modest and self-effacing. ~6s He was, af ter all, a modest man. He had refused some of the honors offered h im and had accepted others ( such as an honor- ary degree f rom Oxford and a Fellowship in the Royal Society) only after strong urging. He had long since received the credit he felt due him as an independent originator of the theory of na tura l selection.

In the published report of the proceedings, Wallace added some selections f rom the sixth edition of Malthus that he thought might have influenced him, remarking, however, that it was the over-all effect ra ther than part icular details that he remembered. He concluded with a well-deserved tribute to Lyell who had, as he said, impressed h im even more deeply than Malthus. 189

Then it was Hooker's turn. It is startling to discover that Hooker's first biographer, Leonard Huxley, refer red to Hooker as the "sole survivor of those immediately concerned," dismiss- ing Wallace altogether ( an oversight repeated by his latest biographer) , 179 and he ment ioned just as casually that "one or two of the letters that then passed were missing." 1~1 But Hooker, in accepting the invitat ion to speak, was ra ther anxious not only about the expediency and propriety of telling the public what he had done, but also about the accuracy of his recollections. He appealed to Sir Francis Darwin and to Sir Leonard Lyell, Sir Charles's nephew, for help in Finding additional documen ta ry evidence. But none could be found, and he was forced to rely entirely on the part ial story in the Life and Letters of Charles Darwin. He carefully noted this in his speech, apologiz- ing at the end for "the half-century-old real or fancied memories of a nonagenar ian ." 17z

But Wallace had saved the early letters he received f rom

168. L in n ean Society of London, T h e D a r w i n - W a l l a c e C e l e b r a t i o n , pp. 5-11.

16g. Ib id . , pp. 111-118. 170. Mea Allan, T h e H o o k e r s o f K e w , 1 7 8 5 - 1 9 1 1 (London: Michael

Joseph, 1967), p. 248. 171. L. Huxley, L i f e a n d L e t t e r s . . . H o o k e r , II, 465. 172. L in n ean Society of London, T h e D a r w i n - W a l l a c e Ce/ebr,,tion,

pp, 11-16.

317

BARBARA G. BEDDALL

Darwin (except for one) , al though the manuscr ip t of his famous paper was never found. On the outside of an envelope in which he kept the letters, Wallace wrote:

The first 8 letters I received f rom Darwin- -whi le in the Malay Archipelago. NB. The MSS. of my Paper sent to Darwin and printed in the Journal of the Linnean Society, was not returned to me, and seems to be lost. The proofs with the MSS. were perhaps sent to Sir Charles Lyell, or to the Secretary of the Linn. S o c . & may some day be found. It was written on thin foreign note paper. 173

However, "neither Wallace's part of this correspondence, nor the original MS. of his essay . . . has been discovered," as Marchant wrote after Wallace's death. 174

And so the story rests, with some questions still unanswered. Why did Wallace first write to Darwin? Why did Darwin send the outline of his theory of natural selection to Asa Gray? What became of the letters Darwin received f rom Wallace, Lyell, Hooker, and Gray? Where is Wallace's manuscr ipt? The answers are in the missing material, and what really happened must remain speculation. The fact that m u c h other material is also missing does not invalidate the point that evidence to support some commonly accepted explanations is inadequate or lacking and that other explanations are clearly in error.

ACKNOWLEDGMENTS

I would like to express my appreciation to Dr. Ernst Mayr for critically reading the manuscript , and to Dr. Everett Men- delsohn for editorial help in preparing it for publication. I would also like to thank Miss Sandra Raphael, Librarian of the Linnean Society of London, for her kind assistance. Fi- nally, I would like to acknowledge my husband's encourage- ment and financial support, without which this would not have been done.

173. Marchant, Wallace, p. 106. 174. Ibid., p. 105.

318

A P P E N D I X • L E T T E R S

No.l F r o m - to Date2 Sources3 C o m m e n t s

1. A . R . Wal l ace - 11 April 1846 W, 1 ,255-256 ; Omiss ion in M. H. W. Bates M, 21.

2. A. 1t. Wa l l ace - 9 Nov. [1847] W, 1 ,254 ; For dating, see H. W. Bates M, 73. note 14.

3. A. It. Wal l ace - 28 Dec. [1847] W, 1, 254-255; Omiss ion in M; for H. W. Bates M, 73-74. dat ing, see note 14.

4. A. 1t. Wa l l ace - [Eaxly 1848] W, 1 ,256-257 ; Omiss ion in Y. H. W. Bates M, 74-75.

5. C. D a r w i n - 3 May [1856] DLL, 1 ,426-427 ; C. Lye11 DLLE, 2, 67-68.

6. C. D a r w i n - 9 May [1856] DLL, 1,427--428; J. D. Hooker DLLE, 2, 66-69.

7. C. D a r w i n - 11 May [1856] DLL, 1,428--430; J. D. Hooker DLLE, 2, 69--71.

*8. A. 11. Wal l ace - [10 Oct. 1856] Received late April C. Daxwm 1857; see 11 and 69,

and notes 43, 81, and 162.

Received July 1857; see 19.

9. H . W . Ba te s - 19 Nov. 1856 A. 11. Wallace

10. C. D a r w i n - 22 Feb. 1857 W. D. Fox

11. C. D a r w i n - 1 May 1857 A. 11. Wallace

12. C. D a r w i n - 20 July [1857] A. Gray

"13. A. Gray- [Aug. 1857] C. Daxwin

M, 52--53.

DLL, 1 ,452; DLLE, 2, 94-95.

DLL, 1,452--454; DLLE, 2, 95-96; M, 107-109.

DLL, 1, 437--438; DLLE, 2, 78-80; Du, 244-245, 458 n22.

Answer to 8, received July 1857; see 19. Omiss ions in DLL; see notes 86, 87, and 162.

See also 14 and 17. For dating, see note 146.

See 14.

3 1 9

L e t t e r s ( c o n t i n u e d )

No.l F r o m - to Date2 Sources3 Comments

14. C. D a r w i n - 5 Sept. [1857] JLZ, 3, 50-53; See also 12 and 17. A. Gray DLL, I, 477-482; Many differences in

DLLE, 2, 122--125; texts between JLZ Du, 246, 458-459 and DLL. For dating, n23. see note 146;

see also note 92.

"15. A. Gray- [Au tumn 1857] C. Darwin

16. A .R . Wal l ace - [27 Sept. 1857] C. Darwin

CUL

17. C. D a r w i n - 29 Nov [1857] DML, 1 ,126-127; A. Gray Du, 247, 459 n24.

18. C. D a r w i n - 22 Dec. 1857 DLL, 1,465--467; A. R. Wallace DLLE, 2, 108-110;

M, 109-111.

19. A .R . Wal l ace - 4 & 25 Jan. 1858 W, 1 ,358-359; H. W. Bates M, 53-55.

*20. A .R . Wal l ace - [Feb. 1858] C. Darwin

21. C. Daxwin- 8 June [1858] DML, 1,109. J. D. Hooker

22. C. D a r w i n - 18 June [1858] DLL, 1,473; C. Lyell DLLE, 2, 116-117.

23. C. D a r w i n - [25 June 1858] DLL, 1 ,474-475; C. / ,yel l DLLE, 2, 117-118.

24. C. D a r w i n - 26 [June 1858] DLL, 1,475; C. Lyell DLLE, 2, 118-119.

25. C. D a r w i n - [29 June 1858] DLL, 1,476; J. D. Hooker DLLE, 2, 119.

26. C. D a r w i n - [29 June 1858] DLL, 1 ,476-477; J. D. Hooker DLLE, 2, 119-120.

*27. C. Lyel l - [June 1858] C. Darwin

*28. J .D. Hooker- [June 1858] C. Darwin

See 17.

Unpubl i shed frag- m e n t of answer to 11. Received Dec. 1857; see 18. See also notes 60, 95, 142, and 143.

Not publ ished unt21 1903. See also 12 and 14. For dating, see note 146.

Answer to 16. Omiss ion in DLL.

Answer to 9. Omissions in W, and texts differ slightly; here quoted f rom M. See also note 94,

Received 18 June 1858; see 22. See also note 116.

Not publ ished unt i l 1903.

See 25 and notes 3, 139-143, and 171.

See 25 and 26, and notes 3, 139-143, and 171.

3 2 0

Letters (continued)

NO. 1 F r o m -- t o D a t e 2 S o u r c e s 3 C o m m e n t s

29. C. Darwin - 5 July [1858] J. D. Hooker

*30. J .D. Hooker- [July 1858] C. Darwin

"31. J .D. Hooker- [13? July 1858] A. R. Wallace

*32. C. Darwin- [13 July 1858] A. R. Wallace

DLL, 1,482--484; DLLE, 2, 124-127.

33. C. Darwin- [13 July 1858] DLL, 1,484--485; J. D. Hooker DLLE, 2, 128-129.

34. C. Darwin - 18 July [1858] DLL, 1,485-486; C. Lyell DLLE, 2, 129-130.

35. C. Darwin- 21 July [1858] DLL, 1,486--487; J. D. Hooker DLLE, 2, 130-131.

36. C. Darwin- [5 Aug. 1858] DLL, 1,489--490; J. D. Hooker DLLE, 2, 133.

37. A .R . w a l l a c e - 6 Oct. 1858 W, 1,365; his mother M, 57-58.

*38. A.R. Wal lace- [Oct. 1858J C. Daxwin

*39. A.R. Wallace- [Oct. 1858] J. D. Hooker

40. C. Darwin- 12 Oct. 1858 DLL, 1,494; J. D. Hooker DLLE, 2, 138.

"41. A.R. Wallace- [30 Nov. 1858] C. Darwin

42. A .R. Wal lace- [Nov. 1858] W, 1,365-367. G. Silk

43. C. Darwin- 25 Jan. [1859] DLL, 1,501-502; A. R. Wallace DLLE, 2, 145-147;

M, 111-112. *44. A.R. Wallace- ?

C. Darwin

45. C. Darwin- 6 April 1859 DML, 1,118-120; A. R. Wallace M, 112--114.

46. C. Darwin- 9 Aug. 1859 DLL, 1, 516--517; A. R. Wallace DLLE, 2, 161-162;

M, 114-115.

47. C. Darwin- 13 Nov. 1859 DLL, 2, 16--17; A. R. Wallace DLLE, 2, 220-221;

M, 115-116.

See 33.

Received early October 1858; see 33, 37, and 42.

Answer to 20, received early October 1858; see 33, 37, and 42.

Omissions in W.

Received [22] January 1859; see 43.

Received [22] January 1859; see 43.

Received 6 April 1859; see 45.

Answer to 38; omissions in DLL.

Received 7 August 1859; see 46.

Answer to 41, but not published until 1903.

Answer to 44; omission in DLL.

321

Letters (continued)

NO.I F r o m - to Date2 Sources3 C o m m e n t s

*48. A.R. Wal lace- ? C. Darwin

*49. A . R . Wallace- [16Feb. 1860] C. Darwin

50. C. Darwin- 7 March 1860 M, 116. A. R. Wallace

51. C. Darwin- 10 April [1860] DLL, 2, 93-95; C. Lyell DLLE, 2, 300-301.

52. C. Darwin- [13 April 1860] DLL, 2, 95-96; J. D. Hooker DLLE, 2, 301-303.

53. C. Darwin- 18 May [1860] DLL, 2, 101-102; C. Lyell DLLE, 2, 308-309.

64. C. Darwin- 18 May 1860 DLL, 2, 102-103; A. R. Wallace DLLE, 2, 309-310;

M, 117-118.

55. A.R. Wal lace- 24 Dee. 1860 H. W. Bates

56. A .R. Wallace-- 15 March 1861 T. Sims

57. C. Darwin- 28 [May? 1864] A. R. Wallace

58. A.R. Wallace-- 29 May [1864] C. Darwin

59. C. Darwin- [Oct. 1865] J. D. Hooker

60. A.R. Wal lace- 19 Nov. 1866 C. Darwin

61. C. Lyell- 4 April 1867 A. R. Wallace

62. C. Lyell- 23 Nov. 1868 E. Haeckel

63. A.R. Wal lace- 22 Nov. 1869 A. B. Meyer

64. C. Lyell- 15 Feb. [1870?] A. R. Wallace

65. C. Darwin- 20 April [1870] A. R. Wallace

66. G. B en tham - 30 May 1882 F. Darwin

W, 1,373-375; M, 59.

M, 59-67.

Received March ? 1860; see 50.

Received 18 May 1860; see 54.

Answer to 48; not published unti l 1916.

Answer to 49. Omissions and changes in DLL; see notes 138, 150.

Omissions in M.

DLL, 2, 271-273; DLLE, 3, 89-91; DML, 2, 32-34; M, 127-128.

DML, 2, 34-37; M, 128-131.

DLL, 2, 225; DLLE, 3, 41.

M, 145-146.

M, 279-281.

LLL, 2, 435--437.

Nat . , 52 (1895), 415. Not published in its entirety unti l 1895.

M, 288-289 .

DLL, 2, 301-302; DLLE, 3, 121; M, 206-207.

DLL, 2, 87-88; DLLE, 2, 293-294.

3 2 2

Letters (continued)

No.1 F r o m - to Date2 Sources3 Comments

67. J .D. Hooker- 22 Oct. 1886 HLL, 2, 300-302. F. Darwin

68. A .R . Wa l l ace - 20 Nov. 1887 M, 295. F. Darwin

69. A . R . Wal l ace - 3 Dec. 1887 DAU, 200-201; A. Newton DAUE, 189-190.

See also notes 7, 39--41, 43, 81, 86, 87, 161, and 162.

1. Miss ing letters, known to have exSsted f rom other correspondence, are marked wi th an asterisk; see "Comments" .

2. Uncer t a in dates of special interest here are discussed in the notes; see " 'Comments".

3. Sources are a r ranged chronologically. Most of t hem are ful ly cited in the notes, the rest here. They are abbreviated as follows:

CUL DAU

DAUE DLL

DLLE DML

Du HLL ILZ LLL

M Nat.

W

Cambridge Univers i ty Library. Franc is Darwin, ed., Charles Darwin: His Life Told in an Auto- biographical Chapter and in a Selected Series of His Published Letters (New York: D. Appleton, 1892). Engl i sh edition of above (London: Murray, 1892). F. Darwin, Life and Letters of Charles Darwin. Engl ish 3-volume edition of above (London: Murray, 1888). F. Darwin, More Letters of Charles Darwin. Dupree, Asa Gray. L. Huxley, Life and Letters of Sir Joseph Dalton Hooker. Journal of the Linnean Society of London (Zoology). K. Lyell, Life, Letters and Journals of Sir Charles Lyell. Marchant , Alfred Russel Wallace. Nature. Wallace, My Life.

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Wallace, Darwin, and the Theory of Natural Selection