In his new book, Darwin Comes to Town, evolutionary biologist Menno Schilthuizen explores how urbanisation is dramatically changing the way animals are evolving to their new-found life in the city, and that it is happening far quicker than Charles Darwin could ever imagine.
But it is not only now that we can see the rapid change evolution, the process of natural selection was observed as far back as the Industrial Revolution. Here is an extract from the book, about Albert Brydges Farn, a lepidopterist who first observed the effects of ‘industrial melanism’ in the annulet moth, long before it was more famously seen in the peppered moth.
Albert Brydges Farn was born in 1841. His entry in The Aurelian Legacy, a sort of Who’s Who for British collectors of Lepidoptera (butterflies and moths), describes him as ‘an all-round naturalist’, ‘a man of vigour, courage, and rather boisterous good humour’. It also calls him ‘a sportsman’, which in those days did not imply he would be seen jogging along country lanes or playing rugby with the village lads, but rather referred to his reputation for potting at bats with a .22 rifle and having bagged a legendary thirty snipe with thirty consecutive shots on Lord Walsingham’s Estate. Farn, clearly, liked to kill stuff.
Most of what he killed, though, were butterflies and moths, which he pinned, mounted, labelled, identified, and organized with great precision. When, in 1921, he died, he left what many considered the finest private lepidopterological collection in Britain at the time. Sadly, the collection was auctioned off piecemeal, and bits of it ended up in different places. Some, says Adam Hart of the University of Gloucestershire, are now ‘in the bowels of the Natural History Museum’ in London. Though he does not know for sure, Hart likes to think that among them are some specimens of the Annulet moth (Charissa obscurata), which Farn collected near Lewes in the 1870s.
It’s a rather drab species, the Annulet, paling in comparison to some of Farn’s prize possessions, like the spectacular Purple Emperor (Apatura iris) that he caught in South Wales. Or the rows and rows of pinned European Map butterfly (Araschnia levana), a pretty black-orange-white species from continental Europe, illegally introduced into the Forest of Dean in 1912 only to be single-handedly exterminated by Farn, who disapproved of all exotic species, pretty or not. Despite its humdrum appearance, though, the Annulet is Farn’s claim to fame. A fame, however, that came 130 years late.
In 2009, Hart, a professor of science communication, was visiting the Gloucester City Museum and Art Gallery in preparation for a class. ‘I was going through some backroom stuff to look for teaching specimens,’ he says. There, he came across a print-out of a letter dated 18 November 1878. The letter had been written by Farn, and the print-out was there because the museum owns a copy of an annotated book that had once belonged to Farn, and the librarian had taken an interest in him. The reason that this particular letter survives and had even been transcribed and put online is, however, not because of the author, but because of the addressee: Charles Darwin.
By 1878, the ageing Darwin was one of the most famous scientists in England. A new generation had grown up since On the Origin of Species had been published, and his name as Mr Evolution was firmly established. Colleagues from all over the world corresponded with him and Darwin kept a meticulous administration of the letters he received and sent – not for social but for scientific reasons. What his correspondents conveyed to him was crucial for his work. As the archivists of the Darwin Correspondence Project at Cambridge University (where much of Darwin’s library is kept) evocatively explain, ‘[h]e went back over some letters again and again as he worked on different subjects, scrawling on them in different coloured pencil, and cut them up so that he could file the pieces with relevant notes or stick them into his experiment book. Letters were dissected like specimens, every useful bit of information sucked out of them and then reincarnated in his publications.’
As far as we know, Albert Farn wrote to Darwin only once. The letter survived in Darwin’s library, the Darwin Correspondence Project duly transcribed it, placed the text online and it was a print-out of this text that Hart found lying around. It is only a brief note, and it seems that Darwin never did anything with it or replied to it.
My dear Sir,
The belief that I am about to relate something which may be
of interest to you, must be my excuse for troubling you with a
Perhaps among the whole of the British Lepidoptera, no
species varies more, according to the locality in which it is
found, than does [the Annulet moth]. They are almost black
on the New Forest peat; grey on limestone; almost white on the
chalk near Lewes; and brown on clay, and on the red soil of
Do these variations point to the ‘survival of the fittest’? I
It was, therefore with some surprise that I took specimens
as dark as any of those in the New Forest on a chalk slope; and
I have pondered for a solution. Can this be it?
It is a curious fact, in connexion with these dark specimens,
that for the last quarter of a century the chalk slope, on
which they occur, has been swept by volumes of black smoke
from some lime-kilns situated at the bottom: the herbage,
although growing luxuriantly, is blackened by it.
I am told, too, that the very light specimens are now much
less common at Lewes than formerly, and that, for some few
years, lime-kilns have been in use there.
These are the facts I desire to bring to your notice.
I am, Dear Sir, Yours very faithfully,
A. B. Farn
‘I must admit it was a bit of a eureka moment,’ says Hart.
‘This letter had been lying around for so long but no one had realised its significance!’ That significance, as Hart pointed out to the evolutionary biology community in a 2010 article in Current Biology, was of course that Farn’s observation may have been the first recorded case of ongoing natural selection. What Farn was suggesting was that the light-coloured Annulets, originally well camouflaged on the pale limestone, had now become sitting ducks against the soot-blackened background, and were being picked off by birds and other predators. Meanwhile, a genetic mutant with dark wings had appeared and had been ‘naturally selected’ because it did not stand out as much as its pale ancestors. If Farn was correct, it would be the very first observation of evolution in action. As Farn rightly anticipated, Darwin should have been thrilled. So why did he ignore Farn’s letter?
Of course, it is possible that Darwin just couldn’t be bothered on that 18th of November 1878. Maybe he was tending to his orchids, or playing with his grandchildren, or laid up with one of his fits of general malaise. But of course we prefer to see a deeper significance in his lack of response. If it means anything, my guess would be that Darwin underestimated the power of his own discovery, natural selection, and that he found it hard to imagine that its work could be observed on the timescale of years or decades. After all, in Chapter IV of On the Origin of Species, he wrote, ‘We see nothing of these slow changes in progress, until the hand of time has marked the long lapse of ages’.
In the preceding pages of his great book, Darwin had laid out the foundations of his theory in four easy, steadfast steps. One – there is variation: in many (sometimes near-imperceptible) ways, each individual is different from the next one. Two – this variation is heritable: offspring resemble their parents. Three – there is surplus: most offspring do not survive. Four – there is selection: survival is not random but favours those who are best suited to the world they live in. To Darwin’s mind – and to everyone since who has fully grasped the enormity of this insight – natural selection is a law of nature. As Darwin wrote, ‘natural selection is daily and hourly scrutinizing, throughout the world, every variation, even the slightest; rejecting that which is bad, preserving and adding up all that is good.’
And yet, despite that ‘daily and hourly’, Darwin did not actually believe that natural selection could be observed in real time. Maybe this was because he lacked the mathematical prowess to calculate exactly how long it would take for natural selection to do its thing. It lasted until the 1920s for mathematical biologists like J.B.S. Haldane and Ronald Fisher to do this. With Darwin’s theory cast in algebraic formulas, it became possible to see whether his pessimism was well founded or not.
As it turned out, it was not. Darwin’s mistake probably was that he imagined natural selection to be a linear process. He may have thought to himself, let’s imagine a population of 100,000 pale-winged moths. Then, a black-winged mutant appears that enjoys a teeny-weeny advantage. Say, an imperceptible 1 per cent – meaning that for every 100 black-winged moths born, surviving and reproducing, there would be 99 pale-winged ones. It’s that small a difference. So how long would it take for all those 100,000 white-wings with one black-winged mutant thrown in, to evolve into a fully black-winged moth population and all pale-winged gone? Forever, right? Wrong – it takes only about a few hundred generations.
That is because natural selection is not a linear process. In the beginning, when the black-wings are still rare, they increase only very slowly, one moth at a time. But when the frequency of black-wings has gone up to a few per cent, the process speeds up, because all those thousands of black-winged moths enjoy the same advantage, and pour their joint offspring into the total gene pool, which becomes duskier by the day.
You can see this for yourself by doing an online simulation. Radford University, for example, has a website where you can enter the population size, the advantage of a mutant (the socalled selection coefficient), and the start-off frequency of the mutant, and you can just watch a virtual population evolve in a nice, S-shaped curve. Play around with the settings and you’ll see that it doesn’t make much difference whether your moth population is 10,000 or 100,000 or even 1 million winged souls: in all cases they evolve into a black-winged moth in less than a thousand generations, while enjoying only a 1 per cent advantage. Make that selection coefficient 5 per cent and it all happens in just 200 generations. For some moth species, 200 generations is less than a century. So, at least in theory, even very weak natural selection can have dramatic effects before the hand of time has begun to mark any longish lapses.
It seems Darwin never really entertained the notion that such evolutionary agility was a possibility. Although . . . In the first four editions of On the Origin of Species, he still writes emphatically, ‘I do believe that natural selection will always act very slowly’. But in the fifth edition, published ten years after the first, he changed ‘always’ for ‘generally’, so he may have begun to doubt that natural selection is so tardy a process. Be that as it may, Darwin missed a trick by not picking up on Farn’s tip, and it was left to the next generation to reveal the breakneck evolution of ‘industrial melanism’. Not in the Annulet moth, but in Biston betularia. The ‘peppered moth’, as it is called, is literally a textbook example of urban evolution, and you’ve probably heard about it in school. But there have been so many recent twists and turns to the story that I hope you’ll forgive me for re-telling it.
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