Chapter 11

{121}On rarity and extinction seeOrigin, Ed. i. pp. 109, 319, vi. pp. 133, 461.{122}In theOrigin, Ed. i. p. 346, vi. p. 493, the author begins his discussion on geographical distribution by minimising the effect of physical conditions. He lays great stress on the effect ofbarriers, as in the present Essay.{123}Note in the original, “Would it be more striking if we took animals, take Rhinoceros, and study their habitats?”{124}Note by Mr A. R. Wallace. “The want of similarity referred to, is, between the mountains of Brazil and Guiana and those of the Andes. Also those of the Indian peninsula as compared with the Himalayas. In both cases there is continuous intervening land.“The islands referred to were, no doubt, the Galapagos for dissimilarity from S. America; our own Islands as compared with Europe, and perhaps Java, for similarity with continental Asia.”{125}The arguments against multiple centres of creation are given in theOrigin, Ed. i. p. 352, vi. p. 499.{126}In theOrigin, Ed. i. p. 366, vi. p. 516, the author does not give his views on the distribution of alpine plants as original but refers to Edward Forbes’ work (Geolog. Survey Memoirs, 1846). In his autobiography, Darwin refers to this. “I was forestalled” he says, “in only one important point, which my vanity has always made me regret.” (Life and Letters, i. p. 88.){127}«The following is written on the back of a page of theMS.» Discuss one or more centres of creation: allude strongly to facilities of dispersal and amount of geological change: allude to mountain-summits afterwards to be referred to. The distribution varies, as everyone knows, according to adaptation, explain going from N. to S. how we come to fresh groups of species in the same general region, but besides this we find difference, according to greatness of barriers, in greater proportion than can be well accounted for by adaptation. «On representive species seeOrigin, Ed. i. p. 349, vi. p. 496.» This very striking when we think of cattle of Pampas, plants «?» &c. &c. Then go into discussion; this holds with 3 or 4 main divisions as well as the endless minor ones in each of these 4 great ones: in these I chiefly refer to mammalia &c. &c. The similarity of type, but not in species, in same continent has been much less insisted on than the dissimilarity of different great regions generically: it is more striking.«I have here omitted an incomprehensible sentence.» Galapagos Islands, Tristan d’Acunha,volcanicislands covered with craters we know lately did not support any organisms. How unlike these islands in nature to neighbouring lands. These facts perhaps more striking than almost any others. [Geology apt to affect geography therefore we ought to expect to find the above.] Geological-geographical distribution. In looking to past times we find Australia equally distinct. S. America was distinct, though with more forms in common. N. America its nearest neighbour more in common,—in some respects more, in some less allied to Europe. Europe we find «?» equally European. For Europe is now part of Asia though not «illegible». Africa unknown,—examples, Elephant, Rhinoceros, Hippopotamus, Hyaena. As geology destroys geography we cannot be surprised in going far back we find Marsupials and Edentata in Europe: but geology destroys geography.{128}Rinconin Spanish means anookorcorner, it is here probably used to mean a small farm.{129}The following is written across the page: “No one would expect a set of similar varieties to be produced in the different countries, so species different.”{130}«The following passage seems to have been meant to follow here.» The parent of an organism, we may generally suppose to be in less favourable condition than the selected offspring and therefore generally in fewer numbers. (This is not borne out by horticulture, mere hypothesis; as an organism in favourable conditions might by selection be adapted to still more favourable conditions.)Barrier would further act in preventing species formed in one part migrating to another part.{131}«The following notes occur on the back of the page.» Number of species not related to capabilities of the country: furthermore not always those best adapted, perhaps explained by creationists by changes and progress. «Seep. 34, note 1.»Although creationists can, by help of geology, explain much, how can he explain the marked relation of past and present in same area, the varying relation in other cases, between past and present, the relation of different parts of same great area. If island, to adjoining continent, if quite different, on mountain summits,—the number of individuals not being related to capabilities, or how &c.—our theory, I believe, can throw much light and all facts accord.{132}SeeOrigin, Ed. i. p. 390, vi. p. 543.{133}On oscillation seeOrigin, Ed. i. p. 291, vi. p. 426.{134}«From the back ofMS.» Effect of climate on stationary island and on continent, but continent once island. Moreover repeated oscillations fresh diffusion when non-united, then isolation, when rising again immigration prevented, new habitats formed, new species, when united free immigration, hence uniform characters. Hence more forms «on?» the island. Mountain summits. Why not true species. First let us recall in Part I, conditions of variation: change of conditions during several generations, and if frequently altered so much better [perhaps excess of food]. Secondly, continued selection [while in wild state]. Thirdly, isolation in all or nearly all,—as well to recall advantages of.[In continent, if we look to terrestrial animal, long continued change might go on, which would only cause change in numerical number «? proportions»: if continued long enough might ultimately affect all, though to most continents «there is» chance of immigration. Some few of whole body of species must be long affected and entire selection working same way. But here isolation absent, without barrier, cut off such «illegible». We can see advantage of isolation. But let us take case of island thrown up by volcanic agency at some distances, here we should have occasional visitants, only in few numbers and exposed to new conditions and «illegible» more important,—a quite new grouping of organic beings, which would open out new sources of subsistence, or «would» control «?» old ones. The number would be few, can old have the very best opportunity. «The conquest of the indigenes by introduced organisms shows that the indigenes were not perfectly adapted, seeOrigin, Ed. i. p. 390.» Moreover as the island continued changing,—continued slow changes, river, marshes, lakes, mountains &c. &c., new races as successively formed and a fresh occasional visitant.If island formed continent, some species would emerge and immigrate. Everyone admits continents. We can see why Galapagos and C. Verde differ «seeOrigin, Ed. i. p. 398»], depressed and raised. We can see from this repeated action and the time required for a continent, why many more forms than in New Zealand «seeOrigin, Ed. i. p. 389 for a comparison between New Zealand and the Cape» no mammals or other classes «see however,Origin, Ed. i. p. 393 for the case of the frog». We can at once see how it comes when there has been an old channel of migration,—Cordilleras; we can see why Indian Asiatic Flora,—[why species] having a wide range gives better chance of some arriving at new points and being selected, and adapted to new ends. I need hardly remark no necessity for change.Finally, as continent (most extinction «?» during formation of continent) is formed after repeated elevation and depression, and interchange of species we might foretell much extinction, and that the survivor would belong to same type, as the extinct, in same manner as different part of same continent, which were once separated by space as they are by time «seeOrigin, Ed. i. pp. 339 and 349».As all mammals have descended from one stock, we ought to expect that every continent has been at some time connected, hence obliteration of present ranges. I do not mean that the fossil mammifers found in S. America are the lineal successors «ancestors» of the present forms of S. America: for it is highly improbable that more than one or two cases (who will say how many races after Plata bones) should be found. I believe this from numbers, who have lived,—mere «?» chance of fewness. Moreover in every case from very existence of genera and species only few at one time will leave progeny, under form of new species, to distant ages; and the more distant the ages the fewer the progenitors. An observation may be here appended, bad chance of preservation on rising island, the nurseries of new species, appeal to experience «seeOrigin, Ed. i. p. 292». This observation may be extended, that in all cases, subsiding land must be, in early stages, less favourable to formation of new species; but it will isolate them, and then if land recommences rising how favourable. As preoccupation is bar to diffusion to species, so would it be to a selected variety. But it would not be if that variety was better fitted to some not fully occupied station; so during elevation or the formation of new stations, is scene for new species. But during elevation not favourable to preservation of fossil (except in caverns «?»); when subsidence highly favourable in early stages to preservation of fossils; when subsidence, less sediment. So that our strata, as general rule will be the tomb of old species (not undergoing any change) when rising land the nursery. But if there be vestige will generally be preserved to future ages, the new ones will not be entombed till fresh subsidence supervenes. In this long gap we shall have no record: so that wonderful if we should get transitional forms. I do not mean every stage, for we cannot expect that, as before shown, until geologists will be prepared to say that although under unnaturally favourable condition we can trace in future ages short-horn and Herefordshire «seenote 2, p. 26».{135}After “organs” is inserted, apparently as an afterthought:—“no, and instance metamorphosis, afterwards explicable.”{136}For analogical resemblances seeOrigin, Ed. i. p. 427, vi. p. 582.{137}“Practically when naturalists are at work, they do not trouble themselves about the physiological value of the characters.... If they find a character nearly uniform,... they use it as one of high value,”Origin, Ed. i. p. 417, vi. p. 573.{138}“We are cautioned ... not to class two varieties of the pine-apple together, merely because their fruit, though the most important part, happens to be nearly identical,”Origin, Ed. i. p. 423, vi. p. 579.{139}The whole of this passage is obscure, but the text is quite clear, except for one illegible word.{140}«The exact position of the following passage is uncertain:» “just as it is not likely every present breed of fancy birds and cattle will propagate, only some of the best.”{141}This suggests that the author was not far from the principle of divergence on which he afterwards laid so much stress. SeeOrigin, Ed. i. p. 111, vi. p. 134, alsoLife and Letters, i. p. 84.{142}That is to say the same conditions occurring in different parts of the globe.{143}The position of the following is uncertain, “greyhound and racehorse have an analogy to each other.” The same comparison occurs in theOrigin, Ed. i. p. 427, vi. p. 583.{144}Airis evidently intended; in theMS.wateris written twice.{145}Written between the lines occurs:—“extend to birds and other classes.”{146}Written between the lines occurs:—“many bones merely represented.”{147}In theOrigin, Ed. i. p. 434, vi. p. 595, the termmorphologyis taken as includingunity of type. The paddle of the porpoise and the wing of the bat are there used as instances of morphological resemblance.{148}The sentence is difficult to decipher.{149}In theOrigin, Ed. i. p. 436, vi. p. 598, the author speaks of the “general pattern” being obscured in the paddles of “extinct gigantic sea-lizards.”{150}SeeOrigin, Ed. i. p. 437, vi. p. 599.{151}The following passage seems to have been meant to precede the sentence beginning “These facts”:—“It is evident, that when in each individual species, organs are metamorph. a unity of type extends.”{152}This is, I believe, the first place in which the author uses the words “theory of descent.”{153}The sentence should probably run, “Let us take the case of the vertebrata: if we assume them to be descended from one parent, then by this theory they have been altered &c.”{154}That is “we should call it a morphological fact.”{155}In theOrigin, Ed. i. p. 438, vi. p. 602, the author, referring to the expressions used by naturalists in regard to morphology and metamorphosis, says “On my view these terms may be used literally.”{156}SeeOrigin, Ed. i. p. 439, vi. p. 605.{157}In theOrigin, Ed. i. p. 440, vi. p. 606, the author argues that the “loop-like course of the arteries” in the vertebrate embryo has no direct relation to the conditions of existence.{158}The following passages are written across the page:—“They pass through the same phases, but some, generally called the higher groups, are further metamorphosed.? Degradation and complication? no tendency to perfection.? Justly argued against Lamarck?”{159}An almost identical passage occurs in theOrigin, Ed. i. p. 440, vi. p. 606.{160}The following: “Deaths of brothers «when» old by same peculiar disease” which is written between the lines seems to have been a memorandum which is expanded a few lines lower. I believe the case of the brothers came from Dr R. W. Darwin.{161}See the discussion to this effect in theOrigin, Ed. i. pp. 443-4, vi. p. 610. The author there makes the distinction between a cause affecting the germ-cell and the reaction occurring at a late period of life.{162}Possibly the sentence was meant to end “is not visible till then.”{163}SeeOrigin, Ed. i. pp. 444-5, vi. p. 611. The query appended tomuch lessis justified, since measurement was necessary to prove that the greyhound and bulldog puppies had not nearly acquired “their full amount of proportional difference.”{164}«The following discussion, from the back of the page, is in large measure the same as the text.» I think light can be thrown on these facts. From the following peculiarities being hereditary, [we know that some change in the germinal vesicle is effected, which will only betray itself years after] diseases—man, goitre, gout, baldness, fatness, size, [longevity «illegible» time of reproduction, shape of horns, case of old brothers dying of same disease]. And we know that the germinal vesicle must have been affected, though no effect is apparent or can be apparent till years afterwards,—no more apparent than when these peculiarities appear by the exposure of the full-grown individual. «That is, “the young individual is as apparently free from the hereditary changes which will appear later, as the young is actually free from the changes produced by exposure to certain conditions in adult life.”» So that when we see a variety in cattle, even if the variety be due to act of reproduction, we cannot feel sure at what period this change became apparent. It may have been effected during early age of free life «or» fœtal existence, as monsters show. From arguments before used, and crossing, we may generally suspect in germ; but I repeat it does not follow, that the change should be apparent till life fully developed; any more than fatness depending on heredity should be apparent during early childhood, still less during fœtal existence. In case of horns of cattle, which when inherited must depend on germinal vesicle, obviously no effect till cattle full-grown. Practically it would appear that the [hereditary] peculiarities characterising our domestic races, therefore resulting from vesicle, do not appear with their full characters in very early states; thus though two breeds of cows have calves different, they are not so different,—grey-hound and bull-dog. And this is what is «to» be expected, for man is indifferent to characters of young animals and hence would select those full-grown animals which possessed the desirable characteristics. So that from mere chance we might expect that some of the characters would be such only as became fully apparent in mature life. Furthermore we may suspect it to be a law, that at whatever time a new character appears, whether from vesicle, or effects of external conditions, it would appear at corresponding time «seeOrigin, Ed. i. p. 444». Thus diseases appearing in old age produce children with do.,—early maturity,—longevity,—old men, brothers, of same disease—young children of do. I said men do not select for quality of young,—calf with big bullocks. Silk-worms, peculiarities which, appear in caterpillar state or cocoon state, are transmitted to corresponding states. The effect of this would be that if some peculiarity was born in a young animal, but never exercised, it might be inherited in young animal; but if exercised that part of structure would be increased and would be inherited in corresponding time of life after such training.I have said that man selects in full-life, so would it be in Nature. In struggle of existence, it matters nothing to a feline animal, whether kitten eminently feline, as long as it sucks. Therefore natural selection would act equally well on character which was fully «developed» only in full age. Selection could tend to alter no character in fœtus, (except relation to mother) it would alter less in young state (putting on one side larva condition) but alter every part in full-grown condition. Look to a fœtus and its parent, and again after ages fœtus and its «i. e. the above mentioned parents» descendant; the parent more variable «?» than fœtus, which explains all.{165}Some of these examples occur inOrigin, Ed. i. pp. 450-51, vi. pp. 619-20.{166}The two following sentences are written, one down the margin, the other across the page. “Abortive organs eminently useful in classification. Embryonic state of organs. Rudiments of organs.”{167}I imagine the meaning to be that abortive organs are specific characters in contrast to monstrosities.{168}Minute hanging horns are mentioned in theOrigin, Ed. i. p. 454, vi. p. 625, as occurring in hornless breeds of cattle.{169}Linum flavumis dimorphic: thyme gynodiæcious. It is not clear what point is referred to underGeranium pyrenaicum.{170}The author’s work on duck’s wings &c. is inVar. under Dom., Ed. 2, i. p. 299.{171}The wordsvis medicatrixare inserted after “useless,” apparently as a memorandum.{172}In the male florets of certain Compositæ the style functions merely as a piston for forcing out the pollen.{173}«On the back of the page is the following.» If abortive organs are a trace preserved by hereditary tendency, of organ in ancestor of use, we can at once see why important in natural classification, also why more plain in young animal because, as in last section, the selection has altered the old animal most. I repeat, these wondrous facts, of parts created for no use in past and present time, all can by my theory receive simple explanation; or they receive none and we must be content with some such empty metaphor, as that of De Candolle, who compares creation to a well covered table, and says abortive organs may be compared to the dishes (some should be empty) placed symmetrically!{174}The author doubtless meant that the complex relationships between organisms can be roughly represented by a net in which the knots stand for species.{175}Between the lines occurs:—“one «?» form be lost.”{176}The original sentence is here broken up by the insertion of:—“out of the dust of Java, Sumatra, these «?» allied to past and present age and «illegible», with the stamp of inutility in some of their organs and conversion in others.”{177}Between the lines occur the words:—“Species vary according to same general laws as varieties; they cross according to same laws.”{178}“A cross with a bull-dog has affected for many generations the courage and obstinacy of greyhounds,”Origin, Ed. i. p. 214, vi. p. 327.{179}The simile of the savage and the ship occurs in theOrigin, Ed. i. p. 485, vi. p. 665.{180}In theOrigin, Ed. i. p. 486, vi. p. 665, the author speaks of the “summing up of many contrivances”: I have therefore introduced the above words which make the passage clearer. In theOriginthe comparison is with “a great mechanical invention,”—not with a work of art.{181}See a similar passage in theOrigin, Ed. i. p. 487, vi. p. 667.{182}See theOrigin, Ed. i. p. 488, vi. p. 668.{183}The following discussion, together with some memoranda are on the last page of theMS.“The supposed creative spirit does not create either number or kind which «are» from analogy adapted to site (viz. New Zealand): it does not keep them all permanently adapted to any country,—it works on spots or areas of creation,—it is not persistent for great periods,—it creates forms of same groups in same regions, with no physical similarity,—it creates, on islands or mountain summits, species allied to the neighbouring ones, and not allied to alpine nature as shown in other mountain summits—even different on different island of similarly constituted archipelago, not created on two points: never mammifers created on small isolated island; nor number of organisms adapted to locality: its power seems influenced or related to the range of other species wholly distinct of the same genus,—it does not equally effect, in amount of difference, all the groups of the same class.”{184}This passage is the ancestor of the concluding words in the first edition of theOrigin of Specieswhich have remained substantially unchanged throughout subsequent editions, “There is grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one; and that whilst this planet has gone cycling on according to the fixed law of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being, evolved.” In the 2nd edition “by the Creator” is introduced after “originally breathed.”{185}Compare theOrigin, Ed. i. p. 481, vi. p. 659, “The difficulty is the same as that felt by so many geologists, when Lyell first insisted that long lines of inland cliffs had been formed, and great valleys excavated, by the slow action of the coast-waves.”{186}The cumulative effect of domestication is insisted on in theOrigin, seee.g. Origin, Ed. i. p. 7, vi. p. 8.{187}This type of variation passes into what he describes as the direct effect of conditions. Since they are due to causes acting during the adult life of the organism they might be called individual variations, but he uses this term for congenital variations,e.g.the differences discoverable in plants raised from seeds of the same pod (Origin, Ed. i. p. 45, vi. p. 53).{188}«It is not clear where the following note is meant to come»: Case of Orchis,—most remarkable as not long cultivated by seminal propagation. Case of varieties which soon acquire, likeÆgilopsand Carrot (and Maize)a certain general characterand then go on varying.{189}Here, as in theMS.of 1842, the author is inclined to minimise the variation occurring in nature.{190}This is more strongly stated than in theOrigin, Ed. i. p. 30.{191}SeeOrigin, Ed. i. p. 13.{192}Origin, Ed. i. p. 86, vi. p. 105.{193}It is interesting to find that though the author, like his contemporaries, believed in the inheritance of acquired characters, he excluded the case of mutilation.{194}This corresponds toOrigin, Ed. i. p. 10, vi. p. 9.{195}Origin, Ed. i. p. 8, vi. p. 10.{196}ForplasticityseeOrigin, Ed. i. pp. 12, 132.{197}Var. under Dom., Ed. ii.I.p. 393.{198}Selection is here used in the sense of isolation, rather than as implying the summation of small differences. Professor Henslow in hisHeredity of Acquired Characters in Plants, 1908, p. 2, quotes from Darwin’sVar. under Dom., Ed. i.II.p. 271, a passage in which the author, speaking of the direct action of conditions, says:—“A new sub-variety would thus be produced without the aid of selection.” Darwin certainly did not mean to imply that such varieties are freed from the action of natural selection, but merely that a new form may appear withoutsummationof new characters. Professor Henslow is apparently unaware that the above passage is omitted in the second edition ofVar. under Dom.,II.p. 260.{199}See the Essay of 1842, p.3.{200}SeeOrigin, Ed. i. p. 33, vi. p. 38. The evidence is given in the present Essay rather more fully than in theOrigin.{201}Journal of Researches, Ed. 1860, p. 214. “Doggies catch otters, old women no.”{202}The effects of crossing is much more strongly stated here than in theOrigin. See Ed. i. p. 20, vi. p. 23, where indeed the opposite point of view is given. His change of opinion may be due to his work on pigeons. The whole of the discussion on crossing corresponds to Chapter VIII of theOrigin, Ed. i. rather than to anything in the earlier part of the book.{203}The parallelism between the effects of a cross and the effects of conditions is given from a different point of view in theOrigin, Ed. i. p. 266, vi. p. 391. See the experimental evidence for this important principle in the author’s work onCross and Self-Fertilisation. Professor Bateson has suggested that the experiments should be repeated with gametically pure plants.{204}The so-called Knight-Darwin Law is often misunderstood. See Goebel inDarwin and Modern Science, 1909, p. 419; also F. Darwin,Nature, Oct. 27, 1898.{205}Pallas’ theory is discussed in theOrigin, Ed. i. pp. 253, 254, vi. p. 374.{206}See Darwin’s paper on the fertility of hybrids from the common and Chinese goose inNature, Jan. 1, 1880.{207}Origin, Ed. i. p. 19, vi. p. 22.{208}Var. under Dom., Ed. ii. vol.II.p. 211.{209}This discussion corresponds to theOrigin, Ed. i. pp. 11 and 143, vi. pp. 13 and 177.{210}SeeOrigin, Ed. i. p. 7, vi. p. 7.{211}«Note in the original.» “Isidore G. St Hilaire insists that breeding in captivity essential element. Schleiden on alkalies. «SeeVar. under Dom., Ed. ii. vol.II.p. 244, note 10.» What is it in domestication which causes variation?”{212}«Note in the original.» “It appears that slight changes of condition «are» good for health; that more change affects the generative system, so that variation results in the offspring; that still more change checks or destroys fertility not of the offspring.” Compare theOrigin, Ed. i. p. 9, vi. p. 11. What the meaning of “not of the offspring” may be is not clear.

{121}On rarity and extinction seeOrigin, Ed. i. pp. 109, 319, vi. pp. 133, 461.

{122}In theOrigin, Ed. i. p. 346, vi. p. 493, the author begins his discussion on geographical distribution by minimising the effect of physical conditions. He lays great stress on the effect ofbarriers, as in the present Essay.

{123}Note in the original, “Would it be more striking if we took animals, take Rhinoceros, and study their habitats?”

{124}Note by Mr A. R. Wallace. “The want of similarity referred to, is, between the mountains of Brazil and Guiana and those of the Andes. Also those of the Indian peninsula as compared with the Himalayas. In both cases there is continuous intervening land.“The islands referred to were, no doubt, the Galapagos for dissimilarity from S. America; our own Islands as compared with Europe, and perhaps Java, for similarity with continental Asia.”

“The islands referred to were, no doubt, the Galapagos for dissimilarity from S. America; our own Islands as compared with Europe, and perhaps Java, for similarity with continental Asia.”

{125}The arguments against multiple centres of creation are given in theOrigin, Ed. i. p. 352, vi. p. 499.

{126}In theOrigin, Ed. i. p. 366, vi. p. 516, the author does not give his views on the distribution of alpine plants as original but refers to Edward Forbes’ work (Geolog. Survey Memoirs, 1846). In his autobiography, Darwin refers to this. “I was forestalled” he says, “in only one important point, which my vanity has always made me regret.” (Life and Letters, i. p. 88.)

{127}«The following is written on the back of a page of theMS.» Discuss one or more centres of creation: allude strongly to facilities of dispersal and amount of geological change: allude to mountain-summits afterwards to be referred to. The distribution varies, as everyone knows, according to adaptation, explain going from N. to S. how we come to fresh groups of species in the same general region, but besides this we find difference, according to greatness of barriers, in greater proportion than can be well accounted for by adaptation. «On representive species seeOrigin, Ed. i. p. 349, vi. p. 496.» This very striking when we think of cattle of Pampas, plants «?» &c. &c. Then go into discussion; this holds with 3 or 4 main divisions as well as the endless minor ones in each of these 4 great ones: in these I chiefly refer to mammalia &c. &c. The similarity of type, but not in species, in same continent has been much less insisted on than the dissimilarity of different great regions generically: it is more striking.«I have here omitted an incomprehensible sentence.» Galapagos Islands, Tristan d’Acunha,volcanicislands covered with craters we know lately did not support any organisms. How unlike these islands in nature to neighbouring lands. These facts perhaps more striking than almost any others. [Geology apt to affect geography therefore we ought to expect to find the above.] Geological-geographical distribution. In looking to past times we find Australia equally distinct. S. America was distinct, though with more forms in common. N. America its nearest neighbour more in common,—in some respects more, in some less allied to Europe. Europe we find «?» equally European. For Europe is now part of Asia though not «illegible». Africa unknown,—examples, Elephant, Rhinoceros, Hippopotamus, Hyaena. As geology destroys geography we cannot be surprised in going far back we find Marsupials and Edentata in Europe: but geology destroys geography.

«I have here omitted an incomprehensible sentence.» Galapagos Islands, Tristan d’Acunha,volcanicislands covered with craters we know lately did not support any organisms. How unlike these islands in nature to neighbouring lands. These facts perhaps more striking than almost any others. [Geology apt to affect geography therefore we ought to expect to find the above.] Geological-geographical distribution. In looking to past times we find Australia equally distinct. S. America was distinct, though with more forms in common. N. America its nearest neighbour more in common,—in some respects more, in some less allied to Europe. Europe we find «?» equally European. For Europe is now part of Asia though not «illegible». Africa unknown,—examples, Elephant, Rhinoceros, Hippopotamus, Hyaena. As geology destroys geography we cannot be surprised in going far back we find Marsupials and Edentata in Europe: but geology destroys geography.

{128}Rinconin Spanish means anookorcorner, it is here probably used to mean a small farm.

{129}The following is written across the page: “No one would expect a set of similar varieties to be produced in the different countries, so species different.”

{130}«The following passage seems to have been meant to follow here.» The parent of an organism, we may generally suppose to be in less favourable condition than the selected offspring and therefore generally in fewer numbers. (This is not borne out by horticulture, mere hypothesis; as an organism in favourable conditions might by selection be adapted to still more favourable conditions.)Barrier would further act in preventing species formed in one part migrating to another part.

Barrier would further act in preventing species formed in one part migrating to another part.

{131}«The following notes occur on the back of the page.» Number of species not related to capabilities of the country: furthermore not always those best adapted, perhaps explained by creationists by changes and progress. «Seep. 34, note 1.»Although creationists can, by help of geology, explain much, how can he explain the marked relation of past and present in same area, the varying relation in other cases, between past and present, the relation of different parts of same great area. If island, to adjoining continent, if quite different, on mountain summits,—the number of individuals not being related to capabilities, or how &c.—our theory, I believe, can throw much light and all facts accord.

Although creationists can, by help of geology, explain much, how can he explain the marked relation of past and present in same area, the varying relation in other cases, between past and present, the relation of different parts of same great area. If island, to adjoining continent, if quite different, on mountain summits,—the number of individuals not being related to capabilities, or how &c.—our theory, I believe, can throw much light and all facts accord.

{132}SeeOrigin, Ed. i. p. 390, vi. p. 543.

{133}On oscillation seeOrigin, Ed. i. p. 291, vi. p. 426.

{134}«From the back ofMS.» Effect of climate on stationary island and on continent, but continent once island. Moreover repeated oscillations fresh diffusion when non-united, then isolation, when rising again immigration prevented, new habitats formed, new species, when united free immigration, hence uniform characters. Hence more forms «on?» the island. Mountain summits. Why not true species. First let us recall in Part I, conditions of variation: change of conditions during several generations, and if frequently altered so much better [perhaps excess of food]. Secondly, continued selection [while in wild state]. Thirdly, isolation in all or nearly all,—as well to recall advantages of.[In continent, if we look to terrestrial animal, long continued change might go on, which would only cause change in numerical number «? proportions»: if continued long enough might ultimately affect all, though to most continents «there is» chance of immigration. Some few of whole body of species must be long affected and entire selection working same way. But here isolation absent, without barrier, cut off such «illegible». We can see advantage of isolation. But let us take case of island thrown up by volcanic agency at some distances, here we should have occasional visitants, only in few numbers and exposed to new conditions and «illegible» more important,—a quite new grouping of organic beings, which would open out new sources of subsistence, or «would» control «?» old ones. The number would be few, can old have the very best opportunity. «The conquest of the indigenes by introduced organisms shows that the indigenes were not perfectly adapted, seeOrigin, Ed. i. p. 390.» Moreover as the island continued changing,—continued slow changes, river, marshes, lakes, mountains &c. &c., new races as successively formed and a fresh occasional visitant.If island formed continent, some species would emerge and immigrate. Everyone admits continents. We can see why Galapagos and C. Verde differ «seeOrigin, Ed. i. p. 398»], depressed and raised. We can see from this repeated action and the time required for a continent, why many more forms than in New Zealand «seeOrigin, Ed. i. p. 389 for a comparison between New Zealand and the Cape» no mammals or other classes «see however,Origin, Ed. i. p. 393 for the case of the frog». We can at once see how it comes when there has been an old channel of migration,—Cordilleras; we can see why Indian Asiatic Flora,—[why species] having a wide range gives better chance of some arriving at new points and being selected, and adapted to new ends. I need hardly remark no necessity for change.Finally, as continent (most extinction «?» during formation of continent) is formed after repeated elevation and depression, and interchange of species we might foretell much extinction, and that the survivor would belong to same type, as the extinct, in same manner as different part of same continent, which were once separated by space as they are by time «seeOrigin, Ed. i. pp. 339 and 349».As all mammals have descended from one stock, we ought to expect that every continent has been at some time connected, hence obliteration of present ranges. I do not mean that the fossil mammifers found in S. America are the lineal successors «ancestors» of the present forms of S. America: for it is highly improbable that more than one or two cases (who will say how many races after Plata bones) should be found. I believe this from numbers, who have lived,—mere «?» chance of fewness. Moreover in every case from very existence of genera and species only few at one time will leave progeny, under form of new species, to distant ages; and the more distant the ages the fewer the progenitors. An observation may be here appended, bad chance of preservation on rising island, the nurseries of new species, appeal to experience «seeOrigin, Ed. i. p. 292». This observation may be extended, that in all cases, subsiding land must be, in early stages, less favourable to formation of new species; but it will isolate them, and then if land recommences rising how favourable. As preoccupation is bar to diffusion to species, so would it be to a selected variety. But it would not be if that variety was better fitted to some not fully occupied station; so during elevation or the formation of new stations, is scene for new species. But during elevation not favourable to preservation of fossil (except in caverns «?»); when subsidence highly favourable in early stages to preservation of fossils; when subsidence, less sediment. So that our strata, as general rule will be the tomb of old species (not undergoing any change) when rising land the nursery. But if there be vestige will generally be preserved to future ages, the new ones will not be entombed till fresh subsidence supervenes. In this long gap we shall have no record: so that wonderful if we should get transitional forms. I do not mean every stage, for we cannot expect that, as before shown, until geologists will be prepared to say that although under unnaturally favourable condition we can trace in future ages short-horn and Herefordshire «seenote 2, p. 26».

[In continent, if we look to terrestrial animal, long continued change might go on, which would only cause change in numerical number «? proportions»: if continued long enough might ultimately affect all, though to most continents «there is» chance of immigration. Some few of whole body of species must be long affected and entire selection working same way. But here isolation absent, without barrier, cut off such «illegible». We can see advantage of isolation. But let us take case of island thrown up by volcanic agency at some distances, here we should have occasional visitants, only in few numbers and exposed to new conditions and «illegible» more important,—a quite new grouping of organic beings, which would open out new sources of subsistence, or «would» control «?» old ones. The number would be few, can old have the very best opportunity. «The conquest of the indigenes by introduced organisms shows that the indigenes were not perfectly adapted, seeOrigin, Ed. i. p. 390.» Moreover as the island continued changing,—continued slow changes, river, marshes, lakes, mountains &c. &c., new races as successively formed and a fresh occasional visitant.

If island formed continent, some species would emerge and immigrate. Everyone admits continents. We can see why Galapagos and C. Verde differ «seeOrigin, Ed. i. p. 398»], depressed and raised. We can see from this repeated action and the time required for a continent, why many more forms than in New Zealand «seeOrigin, Ed. i. p. 389 for a comparison between New Zealand and the Cape» no mammals or other classes «see however,Origin, Ed. i. p. 393 for the case of the frog». We can at once see how it comes when there has been an old channel of migration,—Cordilleras; we can see why Indian Asiatic Flora,—[why species] having a wide range gives better chance of some arriving at new points and being selected, and adapted to new ends. I need hardly remark no necessity for change.

Finally, as continent (most extinction «?» during formation of continent) is formed after repeated elevation and depression, and interchange of species we might foretell much extinction, and that the survivor would belong to same type, as the extinct, in same manner as different part of same continent, which were once separated by space as they are by time «seeOrigin, Ed. i. pp. 339 and 349».

As all mammals have descended from one stock, we ought to expect that every continent has been at some time connected, hence obliteration of present ranges. I do not mean that the fossil mammifers found in S. America are the lineal successors «ancestors» of the present forms of S. America: for it is highly improbable that more than one or two cases (who will say how many races after Plata bones) should be found. I believe this from numbers, who have lived,—mere «?» chance of fewness. Moreover in every case from very existence of genera and species only few at one time will leave progeny, under form of new species, to distant ages; and the more distant the ages the fewer the progenitors. An observation may be here appended, bad chance of preservation on rising island, the nurseries of new species, appeal to experience «seeOrigin, Ed. i. p. 292». This observation may be extended, that in all cases, subsiding land must be, in early stages, less favourable to formation of new species; but it will isolate them, and then if land recommences rising how favourable. As preoccupation is bar to diffusion to species, so would it be to a selected variety. But it would not be if that variety was better fitted to some not fully occupied station; so during elevation or the formation of new stations, is scene for new species. But during elevation not favourable to preservation of fossil (except in caverns «?»); when subsidence highly favourable in early stages to preservation of fossils; when subsidence, less sediment. So that our strata, as general rule will be the tomb of old species (not undergoing any change) when rising land the nursery. But if there be vestige will generally be preserved to future ages, the new ones will not be entombed till fresh subsidence supervenes. In this long gap we shall have no record: so that wonderful if we should get transitional forms. I do not mean every stage, for we cannot expect that, as before shown, until geologists will be prepared to say that although under unnaturally favourable condition we can trace in future ages short-horn and Herefordshire «seenote 2, p. 26».

{135}After “organs” is inserted, apparently as an afterthought:—“no, and instance metamorphosis, afterwards explicable.”

{136}For analogical resemblances seeOrigin, Ed. i. p. 427, vi. p. 582.

{137}“Practically when naturalists are at work, they do not trouble themselves about the physiological value of the characters.... If they find a character nearly uniform,... they use it as one of high value,”Origin, Ed. i. p. 417, vi. p. 573.

{138}“We are cautioned ... not to class two varieties of the pine-apple together, merely because their fruit, though the most important part, happens to be nearly identical,”Origin, Ed. i. p. 423, vi. p. 579.

{139}The whole of this passage is obscure, but the text is quite clear, except for one illegible word.

{140}«The exact position of the following passage is uncertain:» “just as it is not likely every present breed of fancy birds and cattle will propagate, only some of the best.”

{141}This suggests that the author was not far from the principle of divergence on which he afterwards laid so much stress. SeeOrigin, Ed. i. p. 111, vi. p. 134, alsoLife and Letters, i. p. 84.

{142}That is to say the same conditions occurring in different parts of the globe.

{143}The position of the following is uncertain, “greyhound and racehorse have an analogy to each other.” The same comparison occurs in theOrigin, Ed. i. p. 427, vi. p. 583.

{144}Airis evidently intended; in theMS.wateris written twice.

{145}Written between the lines occurs:—“extend to birds and other classes.”

{146}Written between the lines occurs:—“many bones merely represented.”

{147}In theOrigin, Ed. i. p. 434, vi. p. 595, the termmorphologyis taken as includingunity of type. The paddle of the porpoise and the wing of the bat are there used as instances of morphological resemblance.

{148}The sentence is difficult to decipher.

{149}In theOrigin, Ed. i. p. 436, vi. p. 598, the author speaks of the “general pattern” being obscured in the paddles of “extinct gigantic sea-lizards.”

{150}SeeOrigin, Ed. i. p. 437, vi. p. 599.

{151}The following passage seems to have been meant to precede the sentence beginning “These facts”:—“It is evident, that when in each individual species, organs are metamorph. a unity of type extends.”

{152}This is, I believe, the first place in which the author uses the words “theory of descent.”

{153}The sentence should probably run, “Let us take the case of the vertebrata: if we assume them to be descended from one parent, then by this theory they have been altered &c.”

{154}That is “we should call it a morphological fact.”

{155}In theOrigin, Ed. i. p. 438, vi. p. 602, the author, referring to the expressions used by naturalists in regard to morphology and metamorphosis, says “On my view these terms may be used literally.”

{156}SeeOrigin, Ed. i. p. 439, vi. p. 605.

{157}In theOrigin, Ed. i. p. 440, vi. p. 606, the author argues that the “loop-like course of the arteries” in the vertebrate embryo has no direct relation to the conditions of existence.

{158}The following passages are written across the page:—“They pass through the same phases, but some, generally called the higher groups, are further metamorphosed.? Degradation and complication? no tendency to perfection.? Justly argued against Lamarck?”

{158}The following passages are written across the page:—“They pass through the same phases, but some, generally called the higher groups, are further metamorphosed.

? Degradation and complication? no tendency to perfection.

? Justly argued against Lamarck?”

{159}An almost identical passage occurs in theOrigin, Ed. i. p. 440, vi. p. 606.

{160}The following: “Deaths of brothers «when» old by same peculiar disease” which is written between the lines seems to have been a memorandum which is expanded a few lines lower. I believe the case of the brothers came from Dr R. W. Darwin.

{161}See the discussion to this effect in theOrigin, Ed. i. pp. 443-4, vi. p. 610. The author there makes the distinction between a cause affecting the germ-cell and the reaction occurring at a late period of life.

{162}Possibly the sentence was meant to end “is not visible till then.”

{163}SeeOrigin, Ed. i. pp. 444-5, vi. p. 611. The query appended tomuch lessis justified, since measurement was necessary to prove that the greyhound and bulldog puppies had not nearly acquired “their full amount of proportional difference.”

{164}«The following discussion, from the back of the page, is in large measure the same as the text.» I think light can be thrown on these facts. From the following peculiarities being hereditary, [we know that some change in the germinal vesicle is effected, which will only betray itself years after] diseases—man, goitre, gout, baldness, fatness, size, [longevity «illegible» time of reproduction, shape of horns, case of old brothers dying of same disease]. And we know that the germinal vesicle must have been affected, though no effect is apparent or can be apparent till years afterwards,—no more apparent than when these peculiarities appear by the exposure of the full-grown individual. «That is, “the young individual is as apparently free from the hereditary changes which will appear later, as the young is actually free from the changes produced by exposure to certain conditions in adult life.”» So that when we see a variety in cattle, even if the variety be due to act of reproduction, we cannot feel sure at what period this change became apparent. It may have been effected during early age of free life «or» fœtal existence, as monsters show. From arguments before used, and crossing, we may generally suspect in germ; but I repeat it does not follow, that the change should be apparent till life fully developed; any more than fatness depending on heredity should be apparent during early childhood, still less during fœtal existence. In case of horns of cattle, which when inherited must depend on germinal vesicle, obviously no effect till cattle full-grown. Practically it would appear that the [hereditary] peculiarities characterising our domestic races, therefore resulting from vesicle, do not appear with their full characters in very early states; thus though two breeds of cows have calves different, they are not so different,—grey-hound and bull-dog. And this is what is «to» be expected, for man is indifferent to characters of young animals and hence would select those full-grown animals which possessed the desirable characteristics. So that from mere chance we might expect that some of the characters would be such only as became fully apparent in mature life. Furthermore we may suspect it to be a law, that at whatever time a new character appears, whether from vesicle, or effects of external conditions, it would appear at corresponding time «seeOrigin, Ed. i. p. 444». Thus diseases appearing in old age produce children with do.,—early maturity,—longevity,—old men, brothers, of same disease—young children of do. I said men do not select for quality of young,—calf with big bullocks. Silk-worms, peculiarities which, appear in caterpillar state or cocoon state, are transmitted to corresponding states. The effect of this would be that if some peculiarity was born in a young animal, but never exercised, it might be inherited in young animal; but if exercised that part of structure would be increased and would be inherited in corresponding time of life after such training.I have said that man selects in full-life, so would it be in Nature. In struggle of existence, it matters nothing to a feline animal, whether kitten eminently feline, as long as it sucks. Therefore natural selection would act equally well on character which was fully «developed» only in full age. Selection could tend to alter no character in fœtus, (except relation to mother) it would alter less in young state (putting on one side larva condition) but alter every part in full-grown condition. Look to a fœtus and its parent, and again after ages fœtus and its «i. e. the above mentioned parents» descendant; the parent more variable «?» than fœtus, which explains all.

I have said that man selects in full-life, so would it be in Nature. In struggle of existence, it matters nothing to a feline animal, whether kitten eminently feline, as long as it sucks. Therefore natural selection would act equally well on character which was fully «developed» only in full age. Selection could tend to alter no character in fœtus, (except relation to mother) it would alter less in young state (putting on one side larva condition) but alter every part in full-grown condition. Look to a fœtus and its parent, and again after ages fœtus and its «i. e. the above mentioned parents» descendant; the parent more variable «?» than fœtus, which explains all.

{165}Some of these examples occur inOrigin, Ed. i. pp. 450-51, vi. pp. 619-20.

{166}The two following sentences are written, one down the margin, the other across the page. “Abortive organs eminently useful in classification. Embryonic state of organs. Rudiments of organs.”

{167}I imagine the meaning to be that abortive organs are specific characters in contrast to monstrosities.

{168}Minute hanging horns are mentioned in theOrigin, Ed. i. p. 454, vi. p. 625, as occurring in hornless breeds of cattle.

{169}Linum flavumis dimorphic: thyme gynodiæcious. It is not clear what point is referred to underGeranium pyrenaicum.

{170}The author’s work on duck’s wings &c. is inVar. under Dom., Ed. 2, i. p. 299.

{171}The wordsvis medicatrixare inserted after “useless,” apparently as a memorandum.

{172}In the male florets of certain Compositæ the style functions merely as a piston for forcing out the pollen.

{173}«On the back of the page is the following.» If abortive organs are a trace preserved by hereditary tendency, of organ in ancestor of use, we can at once see why important in natural classification, also why more plain in young animal because, as in last section, the selection has altered the old animal most. I repeat, these wondrous facts, of parts created for no use in past and present time, all can by my theory receive simple explanation; or they receive none and we must be content with some such empty metaphor, as that of De Candolle, who compares creation to a well covered table, and says abortive organs may be compared to the dishes (some should be empty) placed symmetrically!

{174}The author doubtless meant that the complex relationships between organisms can be roughly represented by a net in which the knots stand for species.

{175}Between the lines occurs:—“one «?» form be lost.”

{176}The original sentence is here broken up by the insertion of:—“out of the dust of Java, Sumatra, these «?» allied to past and present age and «illegible», with the stamp of inutility in some of their organs and conversion in others.”

{177}Between the lines occur the words:—“Species vary according to same general laws as varieties; they cross according to same laws.”

{178}“A cross with a bull-dog has affected for many generations the courage and obstinacy of greyhounds,”Origin, Ed. i. p. 214, vi. p. 327.

{179}The simile of the savage and the ship occurs in theOrigin, Ed. i. p. 485, vi. p. 665.

{180}In theOrigin, Ed. i. p. 486, vi. p. 665, the author speaks of the “summing up of many contrivances”: I have therefore introduced the above words which make the passage clearer. In theOriginthe comparison is with “a great mechanical invention,”—not with a work of art.

{181}See a similar passage in theOrigin, Ed. i. p. 487, vi. p. 667.

{182}See theOrigin, Ed. i. p. 488, vi. p. 668.

{183}The following discussion, together with some memoranda are on the last page of theMS.“The supposed creative spirit does not create either number or kind which «are» from analogy adapted to site (viz. New Zealand): it does not keep them all permanently adapted to any country,—it works on spots or areas of creation,—it is not persistent for great periods,—it creates forms of same groups in same regions, with no physical similarity,—it creates, on islands or mountain summits, species allied to the neighbouring ones, and not allied to alpine nature as shown in other mountain summits—even different on different island of similarly constituted archipelago, not created on two points: never mammifers created on small isolated island; nor number of organisms adapted to locality: its power seems influenced or related to the range of other species wholly distinct of the same genus,—it does not equally effect, in amount of difference, all the groups of the same class.”

{184}This passage is the ancestor of the concluding words in the first edition of theOrigin of Specieswhich have remained substantially unchanged throughout subsequent editions, “There is grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one; and that whilst this planet has gone cycling on according to the fixed law of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being, evolved.” In the 2nd edition “by the Creator” is introduced after “originally breathed.”

{185}Compare theOrigin, Ed. i. p. 481, vi. p. 659, “The difficulty is the same as that felt by so many geologists, when Lyell first insisted that long lines of inland cliffs had been formed, and great valleys excavated, by the slow action of the coast-waves.”

{186}The cumulative effect of domestication is insisted on in theOrigin, seee.g. Origin, Ed. i. p. 7, vi. p. 8.

{187}This type of variation passes into what he describes as the direct effect of conditions. Since they are due to causes acting during the adult life of the organism they might be called individual variations, but he uses this term for congenital variations,e.g.the differences discoverable in plants raised from seeds of the same pod (Origin, Ed. i. p. 45, vi. p. 53).

{188}«It is not clear where the following note is meant to come»: Case of Orchis,—most remarkable as not long cultivated by seminal propagation. Case of varieties which soon acquire, likeÆgilopsand Carrot (and Maize)a certain general characterand then go on varying.

{189}Here, as in theMS.of 1842, the author is inclined to minimise the variation occurring in nature.

{190}This is more strongly stated than in theOrigin, Ed. i. p. 30.

{191}SeeOrigin, Ed. i. p. 13.

{192}Origin, Ed. i. p. 86, vi. p. 105.

{193}It is interesting to find that though the author, like his contemporaries, believed in the inheritance of acquired characters, he excluded the case of mutilation.

{194}This corresponds toOrigin, Ed. i. p. 10, vi. p. 9.

{195}Origin, Ed. i. p. 8, vi. p. 10.

{196}ForplasticityseeOrigin, Ed. i. pp. 12, 132.

{197}Var. under Dom., Ed. ii.I.p. 393.

{198}Selection is here used in the sense of isolation, rather than as implying the summation of small differences. Professor Henslow in hisHeredity of Acquired Characters in Plants, 1908, p. 2, quotes from Darwin’sVar. under Dom., Ed. i.II.p. 271, a passage in which the author, speaking of the direct action of conditions, says:—“A new sub-variety would thus be produced without the aid of selection.” Darwin certainly did not mean to imply that such varieties are freed from the action of natural selection, but merely that a new form may appear withoutsummationof new characters. Professor Henslow is apparently unaware that the above passage is omitted in the second edition ofVar. under Dom.,II.p. 260.

{199}See the Essay of 1842, p.3.

{200}SeeOrigin, Ed. i. p. 33, vi. p. 38. The evidence is given in the present Essay rather more fully than in theOrigin.

{201}Journal of Researches, Ed. 1860, p. 214. “Doggies catch otters, old women no.”

{202}The effects of crossing is much more strongly stated here than in theOrigin. See Ed. i. p. 20, vi. p. 23, where indeed the opposite point of view is given. His change of opinion may be due to his work on pigeons. The whole of the discussion on crossing corresponds to Chapter VIII of theOrigin, Ed. i. rather than to anything in the earlier part of the book.

{203}The parallelism between the effects of a cross and the effects of conditions is given from a different point of view in theOrigin, Ed. i. p. 266, vi. p. 391. See the experimental evidence for this important principle in the author’s work onCross and Self-Fertilisation. Professor Bateson has suggested that the experiments should be repeated with gametically pure plants.

{204}The so-called Knight-Darwin Law is often misunderstood. See Goebel inDarwin and Modern Science, 1909, p. 419; also F. Darwin,Nature, Oct. 27, 1898.

{205}Pallas’ theory is discussed in theOrigin, Ed. i. pp. 253, 254, vi. p. 374.

{206}See Darwin’s paper on the fertility of hybrids from the common and Chinese goose inNature, Jan. 1, 1880.

{207}Origin, Ed. i. p. 19, vi. p. 22.

{208}Var. under Dom., Ed. ii. vol.II.p. 211.

{209}This discussion corresponds to theOrigin, Ed. i. pp. 11 and 143, vi. pp. 13 and 177.

{210}SeeOrigin, Ed. i. p. 7, vi. p. 7.

{211}«Note in the original.» “Isidore G. St Hilaire insists that breeding in captivity essential element. Schleiden on alkalies. «SeeVar. under Dom., Ed. ii. vol.II.p. 244, note 10.» What is it in domestication which causes variation?”

{212}«Note in the original.» “It appears that slight changes of condition «are» good for health; that more change affects the generative system, so that variation results in the offspring; that still more change checks or destroys fertility not of the offspring.” Compare theOrigin, Ed. i. p. 9, vi. p. 11. What the meaning of “not of the offspring” may be is not clear.

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