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Chapter 5

GRADUAL APPEARANCE AND DISAPPEARANCE OF SPECIES[326]

[326] This chapter corresponds to ch. X of Origin, Ed. i., vi. ch. XI, "On the geological succession of organic beings."


In the Tertiary system, in the last uplifted beds, we find all the species recent and living in the immediate vicinity; in rather older beds we find only recent species, but some not living in the immediate vicinity[327]; we then find beds with two or three or a few more extinct or very rare species; then considerably more extinct species, but with gaps in the regular increase; and finally we have beds with only two or three or not one living species. Most geologists believe that the gaps in the percentage, that is the sudden increments, in the number of the extinct species in the stages of the Tertiary system are due to the imperfection of the geological record. Hence we are led to believe that the species in the Tertiary system have been gradually introduced; and from analogy to carry on the same view to the Secondary formations. In these latter, however, entire groups of species generally come in abruptly; but this would naturally result, if, as argued in the foregoing chapter, these Secondary deposits are separated by wide epochs. Moreover it is important to observe that, with our increase of knowledge, the gaps between the older formations become fewer and smaller; geologists of a few years standing remember how beautifully has the Devonian system[328] come in between the Carboniferous and Silurian formations. I need hardly observe that the slow and gradual appearance of new forms follows from our theory, for to form a new species, an old one must not only be plastic in its organization, becoming so probably from changes in the conditions of its existence, but a place in the natural economy of the district must [be made,] come to exist, for the selection of some new modification of its structure, better fitted to the surrounding conditions than are the other individuals of the same or other species[329].

[327] Origin, Ed. i. p. 312, vi. p. 453.

[328] In the margin the author has written "Lonsdale." This refers to W. Lonsdale's paper "Notes on the age of the Limestone of South Devonshire," Geolog. Soc. Trans., Series 2, vol. V. 1840, p. 721. According to Mr H. B. Woodward (History of the Geological Society of London, 1907, p. 107) "Lonsdale's 'important and original suggestion of the existence of an intermediary type of PalŠozoic fossils, since called Devonian,' led to a change which was then 'the greatest ever made at one time in the classification of our English formations'." Mr Woodward's quotations are from Murchison and Buckland.

[329] [[Note in original.]] Better begin with this. If species really, after catastrophes, created in showers over world, my theory false. [[In the above passage the author is obviously close to his theory of divergence.]]

In the Tertiary system the same facts, which make us admit as probable that new species have slowly appeared, lead to the admission that old ones have slowly disappeared, not several together, but one after another; and by analogy one is induced to extend this belief to the Secondary and PalŠozoic epochs. In some cases, as the subsidence of a flat country, or the breaking or the joining of an isthmus, and the sudden inroad of many new and destructive species, extinction might be locally sudden. The view entertained by many geologists, that each fauna of each Secondary epoch has been suddenly destroyed over the whole world, so that no succession could be left for the production of new forms, is subversive of my theory, but I see no grounds whatever to admit such a view. On the contrary, the law, which has been made out, with reference to distinct epochs, by independent observers, namely, that the wider the geographical range of a species the longer is its duration in time, seems entirely opposed to any universal extermination[330]. The fact of species of mammiferous animals and fish being renewed at a quicker rate than mollusca, though both aquatic; and of these the terrestrial genera being renewed quicker than the marine; and the marine mollusca being again renewed quicker than the Infusorial animalcula, all seem to show that the extinction and renewal of species does not depend on general catastrophes, but on the particular relations of the several classes to the conditions to which they are exposed[331].

[330] Opposite to this passage the author has written "d'Archiac, Forbes, Lyell."

[331] This passage, for which the author gives as authorities the names of Lyell, Forbes and Ehrenberg, corresponds in part to the discussion beginning on p. 313 of Origin, Ed. i., vi. p. 454.

Some authors seem to consider the fact of a few species having survived[332] amidst a number of extinct forms (as is the case with a tortoise and a crocodile out of the vast number of extinct sub-Himalayan fossils) as strongly opposed to the view of species being mutable. No doubt this would be the case, if it were presupposed with Lamarck that there was some inherent tendency to change and development in all species, for which supposition I see no evidence. As we see some species at present adapted to a wide range of conditions, so we may suppose that such species would survive unchanged and unexterminated for a long time; time generally being from geological causes a correlative of changing conditions. How at present one species becomes adapted to a wide range, and another species to a restricted range of conditions, is of difficult explanation.

[332] The author gives Falconer as his authority: see Origin, Ed. i. p. 313, vi. p. 454.



Extinction of species.

The extinction of the larger quadrupeds, of which we imagine we better know the conditions of existence, has been thought little less wonderful than the appearance of new species; and has, I think, chiefly led to the belief of universal catastrophes. When considering the wonderful disappearance within a late period, whilst recent shells were living, of the numerous great and small mammifers of S. America, one is strongly induced to join with the catastrophists. I believe, however, that very erroneous views are held on this subject. As far as is historically known, the disappearance of species from any one country has been slow--the species becoming rarer and rarer, locally extinct, and finally lost[333]. It may be objected that this has been effected by man's direct agency, or by his indirect agency in altering the state of the country; in this latter case, however, it would be difficult to draw any just distinction between his agency and natural agencies. But we now know in the later Tertiary deposits, that shells become rarer and rarer in the successive beds, and finally disappear: it has happened, also, that shells common in a fossil state, and thought to have been extinct, have been found to be still living species, but very rare ones[334]. If the rule is that organisms become extinct by becoming rarer and rarer, we ought not to view their extinction, even in the case of the larger quadrupeds, as anything wonderful and out of the common course of events. For no naturalist thinks it wonderful that one species of a genus should be rare and another abundant, notwithstanding he be quite incapable of explaining the causes of the comparative rareness[335]. Why is one species of willow-wren or hawk or woodpecker common in England, and another extremely rare: why at the Cape of Good Hope is one species of rhinoceros or antelope far more abundant than other species? Why again is the same species much more abundant in one district of a country than in another district? No doubt there are in each case good causes: but they are unknown and unperceived by us. May we not then safely infer that as certain causes are acting unperceived around us, and are making one species to be common and another exceedingly rare, that they might equally well cause the final extinction of some species without being perceived by us? We should always bear in mind that there is a recurrent struggle for life in every organism, and that in every country a destroying agency is always counteracting the geometrical tendency to increase in every species; and yet without our being able to tell with certainty at what period of life, or at what period of the year, the destruction falls the heaviest. Ought we then to expect to trace the steps by which this destroying power, always at work and scarcely perceived by us, becomes increased, and yet if it continues to increase ever so slowly (without the fertility of the species in question be likewise increased) the average number of the individuals of that species must decrease, and become finally lost. I may give a single instance of a check causing local extermination which might long have escaped discovery[336]; the horse, though swarming in a wild state in La Plata, and likewise under apparently the most unfavourable conditions in the scorched and alternately flooded plains of Caraccas, will not in a wild state extend beyond a certain degree of latitude into the intermediate country of Paraguay; this is owing to a certain fly depositing its eggs on the navels of the foals: as, however, man with a little care can rear horses in a tame state abundantly in Paraguay, the problem of its extinction is probably complicated by the greater exposure of the wild horse to occasional famine from the droughts, to the attacks of the jaguar and other such evils. In the Falkland Islands the check to the increase of the wild horse is said to be loss of the sucking foals[337], from the stallions compelling the mares to travel across bogs and rocks in search of food: if the pasture on these islands decreased a little, the horse, perhaps, would cease to exist in a wild state, not from the absolute want of food, but from the impatience of the stallions urging the mares to travel whilst the foals were too young.

[333] This corresponds approximately to Origin, Ed. i. p. 317, vi. p. 458.

[334] The case of Trigonia, a great Secondary genus of shells surviving in a single species in the Australian seas, is given as an example in the Origin, Ed. i. p. 321, vi. p. 463.

[335] This point, on which the author laid much stress, is discussed in the Origin, Ed. i. p. 319, vi. p. 461.

[336] Origin, Ed. i. p. 72, vi. p. 89.

[337] This case does not occur in the Origin, Ed.

From our more intimate acquaintance with domestic animals, we cannot conceive their extinction without some glaring agency; we forget that they would undoubtedly in a state of nature (where other animals are ready to fill up their place) be acted on in some part of their lives by a destroying agency, keeping their numbers on an average constant. If the common ox was known only as a wild S. African species, we should feel no surprise at hearing that it was a very rare species; and this rarity would be a stage towards its extinction. Even in man, so infinitely better known than any other inhabitant of this world, how impossible it has been found, without statistical calculations, to judge of the proportions of births and deaths, of the duration of life, and of the increase and decrease of population; and still less of the causes of such changes: and yet, as has so often been repeated, decrease in numbers or rarity seems to be the high-road to extinction. To marvel at the extermination of a species appears to me to be the same thing as to know that illness is the road to death,--to look at illness as an ordinary event, nevertheless to conclude, when the sick man dies, that his death has been caused by some unknown and violent agency[338].

[338] An almost identical sentence occurs in the Origin, Ed. i. p. 320, vi. p. 462.

In a future part of this work we shall show that, as a general rule, groups of allied species[339] gradually appear and disappear, one after the other, on the face of the earth, like the individuals of the same species: and we shall then endeavour to show the probable cause of this remarkable fact.

[339] Origin, Ed. i. p. 316, vi. p. 457.


Charles Darwin

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