Worlds After Wallace

Anna-Sophie Springer, Etienne Turpin
Published 21 June 2016

Among the experts on Alfred Russel Wallace in the English-speaking world, Dr. George Beccaloni—a former curator of entomology at London’s Natural History Museum, and the Director of the Alfred Russel Wallace Correspondence Project—is perhaps the most compelling advocate for a reassessment of Wallace’s place in the history of science. His knowledge and excitement are contagious, and throughout our various visits, tours, and conversations, we became increasingly certain that our curatorial engagement with the legacy of Wallace was a necessary project to see through, despite numerous obstacles. During our research, we met with George in his office, while tending to the museum’s insect collection, at his home, and in Epping Forest (one of England’s oldest), to discuss the significance of Wallace’s collections and the legacy of his work today. What follows is an edited version of these various conversations, organized thematically (instead of chronologically) for readability. We are grateful to George for his generosity, mentorship, and good humor over the years. He has helped us grasp the nuances of Wallace’s thought, the importance of natural selection, and the amazing world of entomology.

A S
Given your expertise, it would be great if you would start off by providing a bit of context about Alfred Russel Wallace, The Malay Archipelago, and insect collecting. I would also be particularly interested in how, at the point when the theory of evolution was formulated, this transformation of knowledge changed the way that museums were ordered.  

G B
One of the predecessors of Enlightenment museum collections were the Renaissance cabinets of curiosities, which were just assemblages of interesting and strange objects. After the theory of natural selection was published and people started to accept that species had evolved from other species, displays became much more evolution-based. Wallace, as the co-discoverer of evolution by natural selection, was partly responsible for this. As the founder of evolutionary biogeography, Wallace was also responsible for another popular type of display, the faunal diorama, where animals of a particular region are shown together in one scene. All the taxidermy mammals of the Andes or the Himalayas, say, are placed together against a natural background showing some of the habitat. This method of display derives from the plates in his important book, The Geographical Distribution of Animals. [Fig. 03.]

A S
Let’s step back a bit: who was Wallace and where did he come from?

G B

The basic story is very well known. Wallace was born to a downwardly mobile, middle-class couple in Usk, England (now part of Wales) in 1823. He was educated in Hertford, to the north of London, and had to leave school when he was only fourteen. Charles Darwin left school much later, when he was sixteen, and then went on to two universities. After leaving school Wallace educated himself from books and also attended working men’s clubs. He became interested in natural history whilst working with his brother as a trainee land surveyor, travelling in the countryside of southern England and Wales. His first interest was botany, as he wanted to identify the plants he saw whilst out surveying. He bought his first books on the subject and realized that there was a whole science behind the classification of plants and animals. He formed a collection of pressed plants in order to remember which species he had seen before and more accurately identify them from the books that he read. He then got a job for a year as a teacher in Leicester. That’s when he met Henry Walter Bates, a keen beetle collector who got Wallace passionate about insects. Wallace then returned to Wales and started collecting beetles, moths, and butterflies.

E T
Was entomology a fairly common practice at the time?

G B
Yes, there were many entomologists at the time, and they published their records and observations in various specialist journals, just as they do today. However, entomologists formed a tiny proportion of the population, then as now. Most people think you’re weird when you tell them you collect beetles, and probably the same was true back then.

E T
Do you know how long this amateur scientific community of entomologists was working before Wallace’s time?

G B
The number of amateurs studying insects increased steadily from the mid-eighteenth century, and as a result the insects and other fauna of Britain were pretty well known by the time Wallace began collecting. By the 1850s natural history had also become very popular among the general public. A friend of mine, the writer and artist Errol Fuller, who is interested in the history of taxidermy, has said that everyone had to have a stuffed bird in their living room at that time. So, there was a greater appreciation of and interest in natural history, and a huge demand for showy foreign specimens to display domestically—butterflies on the wall, or a stuffed bird. However, Wallace’s market—the people who did serious scientific work on the collections he sent back from his expedition through the Malay Archipelago (1854–62)—was really just a handful of people. There were probably more amateurs doing the serious work of describing species in Britain than there are now, but that’s not the case everywhere. In Eastern Europe, for example, there are still many amateurs doing that sort of work.

A S
Did the majority of specimens that Wallace sent to Europe from the Archipelago end up in private or public collections?

G B
Probably less than fifty percent were purchased directly by the British Museum. Wallace mostly collected insect and bird specimens, and he shipped them to his agent, Samuel Stephens, in London. Stephens had rooms near the old British Museum (the natural history collections that we have here in South Kensington used to be in Bloomsbury, in what’s now the British Museum). When new shipments came in, Stephens would let the scientists in  the museum know, and they would come to pick out all the things they thought were interesting or new. The rest of the material was then sold to keen amateurs such as William Wilson Saunders. Saunders would take all of Wallace’s smaller orders of insects, whereas the beetles went to certain specialists on the different groups. For instance, Francis P. Pascoe got the longhorn beetles. Stephens often kept some specimens aside for a certain collector. Then there was the general public, who had very little knowledge of natural history but wanted really showy specimens— brightly colored parrots or hummingbirds or whatever—to decorate their homes. We don’t know what proportion of specimens went to the third group of people because typically the original labels were removed. Even if you went through collections of old Victorian taxidermy today (and there are many such collections), you wouldn’t know if they were Wallace specimens or if they were collected by someone else.

E T
But there was a certain accounting procedure, was there not? Everything had to pass through Stephens, who would have had some form of master list to track payments owed to Wallace, no?

G B
Wallace kept rough records of how many specimens and species he collected on each island and shipped back to Stephens. His notebook detailing his consignments to Stephens is in the Linnean Society library. Unfortunately, however, Stephens’s records do not survive.

A S
Are there any shipping papers or transportation documentations available?

G B
None that were issued by the actual shipping agents, at least none that anybody has ever found. Maybe Stephens had lists but they don’t survive at all, so we only have fragmentary information, and we don’t even know exactly where most of Wallace’s specimens are now. We have a fairly good idea which museums have Wallace specimens in their collections, but we generally don’t have lists of the specimens they have. Although I’m pretty sure that there must be thousands of Wallace specimens in the Paris museum, there’s no list of them and no way of easily finding them. This is also true in the Natural History Museum, because our specimens haven’t been individually databased, and won’t be for a very long time, if ever—the collection is just too huge! We have about 25,000,000 insect specimens; although we don’t have a record of what Wallace specimens we have, I have estimated that we must have roughly seventy percent of everything he collected. Our museum not only purchased Wallace’s specimens directly from Stephens, but many others came in collections formed by entomologists which were purchased, donated, or bequeathed to the Museum when the collectors died. The Oxford Museum of Natural History has the second biggest collection of Wallace’s specimens, mostly insects. Sadly, in the whole of the Malay Archipelago there are only two Wallace specimens—a dung beetle in the Sarawak Museum in Malaysia and a drab little bird in the natural history museum in Singapore.

A S
So what first made you interested in Wallace?

G B
When I was doing my Ph.D. on the evolution of mimicry in butterflies from South America, I became interested in theories of animal coloration—for warnings, camouflage, sexual selection, and so on. I realized that it was Wallace who proposed the majority of these theories. I hadn’t really heard of him before, nor did I know that he was the co-discoverer of natural selection, so I started to read a bit more about him. I was reading James Marchant’s Alfred Russel Wallace: Letters and Reminiscences, which says that Wallace was buried in Broadstone, Dorset, on a windswept hill. On the first outing that I had with my wife-to-be, we happened to be camping in that area of Dorset and I had just read this, so we ended up going to find Wallace’s grave. After some searching, we eventually discovered it behind a huge conifer; it was marked by a strange monument, which looks a bit like a phallus on a stone base. I decided to find out who owned it because I felt it was a shame that it was in such bad condition—you had to climb inside the tree in order to see the name plaque, and the roots were tipping it over. I contacted the cemetery and they said that Wallace’s grandsons still owned the grave—I hadn’t realized that any of his grandsons were still alive. I managed to find the address of his grandson Richard and wrote to him, saying that I’d seen their grandfather’s grave was in a sorry state. He wrote back saying something like, “Yes, it’s a great shame. We do our best, but we’re 72 73 getting kind of old and we go there once a year to clear shrubs and brambles from the base.” As Wallace is one of the greatest figures in the natural sciences, at least in biology, I decided that his grave should be restored. I started the Wallace Memorial Fund in order to raise the money to do this and extend the lease on the plot. I’d discovered that the lease only had another fourteen years to run, after which time they would use the plot for another burial and dispose of the monument. I sent articles to various places telling them about the fund and that I was looking for donors. Within a pretty short time we had over 100 donors from all around the world, which enabled us to restore the monument, cut down the tree that was pushing it over, put up a new bronze plaque explaining who Wallace was, and extend the lease on the plot. By this stage I was in touch with various people in places where Wallace had lived and they were really interested in participating; working with them, we set up other monuments.

A S
Tell us more about how Wallace collected and identified his 125,660 specimens?

G B
There are actually people who know very little about natural history who think that Wallace had an easy time, that he sort of just picked out the insects that flew into his hair as he wandered around. In reality, how Wallace went about collecting was an incredibly skilled practice that very few people in the world could do as well as him, even today. He was primarily gathering specimens for his private collections. He always made that very clear, both in his work in the Amazon and in the Malay Archipelago. He was very interested in geographic distribution from early on, and wanted specimens of many species of insects and birds so that he could study them back in Britain. Whenever he collected a species for the first time, he would keep the first specimen or two for his private collection, and only when he had duplicate specimens would he sell them. We know he didn’t collect many of the same species because it wouldn’t have made sense financially. He must have had an incredibly good memory, and not having a camera he had to remember each and every species he collected so he wouldn’t collect them over and over again. He was able to identify most of the bird species he collected using a book he had with him: Lucien Bonaparte’s Conspectus. [Fig. 02.] Rather incredibly, this book has no pictures in it, only brief Latin descriptions of the birds. Even today a top bird specialist would find it incredibly difficult to use a book like that to identify species in the field. But Wallace obviously had a remarkable grasp of the distinguishing characteristics of birds, and using the brief descriptions in this book, he was able to visualize exactly what the species looked like. Even if you have a modern bird book with photographs or illustrations, it is difficult enough to determine what you’ve seen. Yet we know that Wallace accurately identified many birds and realized which were yet unnamed species. He named and described a lot of the new species himself, and sent off others he believed were new with instructions for Stephens. Often Stephens would then contact the bird people at the museum and they would buy and name them. For insects, all he had was a book that described the known species of two families of butterflies: Pieridae and Papilionidae. It was in French and had no illustrations, yet as with his bird book he was able to identify most of the butterfly species he collected. With all the other insects, he memorized what they looked like when he collected them. I have a good memory for that too, and can remember nearly all of the insects I have ever seen—the interesting ones at least! Because Wallace had a photographic memory, he could remember all the species of insects from each island without having to assign scientific names to them. Since most of the insects he was collecting didn’t yet have scientific names anyway, Wallace would just need to know whether he had them yet or not. He assigned a number to each of the species he collected in a particular place and listed the numbers in his collecting notebooks, sometimes with a few notes about the species—two of these notebooks are in our museum here, and one in the Linnean Society’s library.

E T
How would you describe Wallace’s reliance on local knowledge of the species he was collecting? In a way, he was completely out on his own, with one or two books to guide him; so, did he depend on knowledge from local inhabitants on the islands?

G B
I would say that, truly, avifauna distinctions and major patterns are simply not discernable to the average layperson in the archipelago. Wallace’s assistant, Ali, was about as far from a scientist as one could get. So, understanding the newness of a species is the result of knowledge of the descriptions themselves. There’s a much more complicated relationship between the scientific descriptions and the species collected than might first appear to certain historians of science. Some tend to be rather politically correct these days and say that local assistants deserve so much of the credit because they really understood the animals and they are key in the whole process. John van Wyhe even says in his book Dispelling the Darkness that maybe Wallace got his inspiration for the Wallace Line by staying at a local person’s house on Lombok.¹ In reality, the local people really don’t have a clue about major biological patterns like that. So no, it was only Wallace, with a very broad picture of the whole situation, who would have seen the significance in species breaks and continuums across the islands of the region. He knew that the cockatoo was centered in Australia and that a few could be found in Lombok, but no further west. Who else would have been able to draw that kind of conclusion?

Meanwhile, as I said, Ali was completely illiterate. He had no scientific background, no knowledge about the wider issues. It’s a bit like saying that Darwin’s gardener deserves a share of the credit for Darwin’s great work on carnivorous plants because the gardener helped to grow the plants. Ali went out and shot birds. Most of the time, he didn’t know what the significance of the target was. Even if he shot the first Wallace standardwing and thought it was new—and it would have been new to him—it could have been discovered by somebody 100 years earlier. It was Wallace who had the specialist knowledge, and that’s what counts—not just collecting the stuff.

E T
Maybe this leads us towards geography. What is the importance of Wallace’s work on the biogeographical distribution of species?

G B
Well, rhinos and tigers are on one side of the Wallace Line, while marsupials, cockatoos, and birds-of-paradise are on the other. As Wallace said in one of his early papers, when you look at two islands like New Guinea and Borneo, they seem very similar. They are similar in their climate, and their forests look the same on the face of things, and yet in Borneo you have monkeys and in New Guinea you have tree kangaroos and marsupials, but no monkeys. This was Wallace’s key argument against Charles Lyell’s idea regarding “centres of creation,” which held that over geological time, as climates changed, God would create species fitted for the new environment. For Lyell, if the climate changed from a desert to a rain forest, then God would create a whole bunch of monkeys. But this theory wouldn’t explain why everything to the west has monkeys and to the east there’s marsupials. Same climate, east and west. So why would God create different organisms to live in trees and eat leaves in such similar places? Why not just create monkeys everywhere?

A S
Wallace not only argued with Lyell over biogeographical distribution; he also envisioned a mode of display for the museum that would show a continental evolutionary panorama of the species. Did he also have a certain curatorial agenda, so to speak?

G B
Well, he did apply for a position as the director of the British Natural History Museum, which was to be at Bethnal Green. The collection used to be at the British Museum with all the archeology, but they wanted to make a new museum. The curious thing is that Wallace sent Richard Owen drawings of how to arrange the ideal collection of natural history, and the ideas in them are strikingly similar to the way our museum is actually designed. You can almost imagine Owen actually taking the ideas directly from Wallace. You have to look at the drawings because they’re incredibly similar.

E T
Do you have the original correspondence?

G B
No, I don’t think so. This was before the museum was built. Wallace wrote a paper about the design of natural history museums.² He was the first to suggest that animals from one particular place or habitat should all be displayed together in order to give a sense of the fauna in that area. Museums like ours and the Powell-Cotton Museum, which still has the best formal dioramas in Britain, obviously took up this idea. I think the American Museum of Natural History has the best formal dioramas anywhere in the world. Unfortunately, there are no formal dioramas in our museum anymore. There used to be some in what’s known as the Rowland Ward Pavilion—Rowland Ward, the taxidermy company, produced these dioramas free of charge for the museum with the agreement that they would always be there on display. However, about 10 years ago or so, the museum broke this agreement and destroyed them.

E T
Why? Aren’t they of some historical value?

G B
They were just too old-fashioned. The museum needed space for storing old wooden cabinets and things. There was a beautiful display of a scene on the African Plains with a giraffe and giant sable antelope and then one of the Congo forests with other animals. It’s a shame they were destroyed. As for the formal diorama idea, I don’t know if Wallace ever published another paper on it or whether it was just present in his book The Geographical Distribution of Animals. The plates sort of show the animals of one place illustrated together.

A S
This brings me to another question about curatorial thinking, about the importance of commemorating Wallace. Why is it important for you to bring the memory of Wallace into the grand narratives (especially of Darwin) already dominant in the space?

G B
I think the current story about the theory of natural selection is fatally flawed, and is just a kind of fairytale. Darwin has been central while all the other people have been forgotten. After all, Wallace was the co-discoverer of the theory, as he published the paper with Darwin 14 or 15 months before On the Origin of Species was published. So, in my view, he deserves half the credit, but not only that! He and Darwin almost exclusively, together, laid the foundations of modern evolutionary biology and all the other add-on theories in the early days, like understanding animal colouration in an evolutionary context, biogeography, etc. You know, if you think about the lasting scientific achievements of some of the prominent biologists of the nineteenth century like Haeckel and Huxley, you can’t really come up with anything. No major theoretical ideas that they developed have lasted to this day, whereas both Wallace and Darwin made really major contributions that still endure.

A S
So there’s a concern for historical accuracy, but what about the elements of stories that can be told differently through Wallace than Darwin. For example, Wallace is sometimes called the father of conservationism. He was very outspoken on certain issues which we’d now call conservation, which seem quite relevant for contemporary purposes.

G B
Personally, I think that Wallace’s role as an environmentalist has been a bit exaggerated. He didn’t really write that much about it. And yet, what he did write was very powerful and it was probably far ahead of its time. For people like Darwin, on the other hand, it was wonderful that all the natural habitats were going to be replaced by monocultures; he thought that was progress. Wallace sometimes thought like that, but he realized that there would be a major loss of scholarship if all these species were destroyed by development. He was also passionate about the giant redwoods and their destruction in America. He met the pioneer of American conservationism, John Muir, and Wallace was ahead of his time in that respect, but he didn’t really focus his work on environmentalism or conservationism. I suppose that back then it was far less of a problem; it wasn’t nearly as serious as it is today.

A S
We are eager to know whether the species Wallace collected could still be found today, or, if one could currently come up with the theory of natural selection based on available specimens, given the habitat loss one encounters in certain parts of Indonesia?

G B
Actually, very few of the species of insects and birds that Wallace collected in Southeast Asia are known to be extinct. I can’t think of any, in fact. Many of them are probably much more rare than they were, but if you had all the official permissions you could still make collections like Wallace’s today. But it would be impossible now to travel from island to island shooting every bird you wanted. Anyway, collecting birds is not done very often these days. So, the birds are still there, and you could go to Halmahera and kill a whole lot of Wallace standardwings if you had permission. But that wouldn’t be a very responsible thing to do given how rare they are. With insects, it’s a different picture because, in general, you can’t collect enough of one particular species to damage the population. At least it’s pretty difficult to do so. But entomologists are still very active in Indonesia, observing habitats and collecting different kinds of species, just like Wallace did.

E T
So, even though larger mammals are extinct or rare, there’s still a wide array of living evidence for natural selection?

G B
Sure, it’s just that the political situation has changed. It would actually be impossible today to just travel wherever you like and collect what you wanted. So, say for example that the Wallace Line hadn’t yet been discovered. There are two ways you could discover it today: either by doing all the collecting yourself, or by reading enough about the patterns that others have discovered. You might be able to work it out just by reading about the distributions of various groups.

E T
Are such categories changing as a result of new forms of DNA analysis?

G B
Not as much as you might think. It tends to be that DNA studies confirm what expert taxonomists have always thought, or at least that’s the overriding trend. A good example is some work that I did on cockroaches and termites; I initiated a big DNA study of these critters, and we finally proved that termites are actually, truly cockroaches that have evolved to be highly social. This was actually first proposed in the 1930s using obscure morphological characteristics like the structure of the gizzard and different protozoa in the guts of termites and cockroaches—very technical, anatomical things. This is the cockroach tree. [Fig. 06] Rather than being an outlying relative, termites arise from within the tree. Since then, other DNA and RNA studies have confirmed our findings. Anyway, all of this is to say that we reinforced what expert morphologists realized a hundred years ago.

E T
What about mimicry? This idea has been very important for discussions of natural selection; have these discussions been changed by more recent DNA or RNA studies?

G B
Well no, actually. Colour patterning and mimicry were the first great tests of the natural selection theory. If you look at the early papers on the topic, they used this example because it was clear—one species has evolved to look like another species because one’s tasty and one’s nasty, or whatever. All of the arguments were centered around these visual examples, and mimicry was of great interest. Wallace proposed a lot of the ideas that are still valid today about animal colours and polymorphic mimicry in butterflies, where one species has females that look like members of several different species living in the same habitat. In the swallowtail butterfly, for example, males will all look the same and they wouldn’t be mimetic, but the females have different morphs and each of those discrete morphs mimics a different species of poisonous butterfly. Wallace was the first person to explain this.

A S
Can you explain the significance of this?  

G B
Well, there aren’t discrete morphological types of human. We’re mixtures of our parents; whereas with butterflies, there’s one kind of male with, say, black wings, and then five different female colour patterns, each of which has evolved to look like a different species of poisonous butterfly. When they breed, you always get the same males and this array of different female patterns. That is what we call polymorphic mimicry. Wallace discovered this in the Southeast Asian swallowtail butterfly, Papilio memnon, but it became more famous when something similar was discovered in a habitat of swallowtails called Papilio daedalus in Africa. It was then studied for a hundred years and still, to this day, researchers are trying to work out how the different mimetic morphs actually arose, and the genetics of this process. Wallace must have gotten a batch of eggs, reared the caterpillars, and saw that they produced black males and five different types of females; he realized that all these females he thought were different were actually of the same species.

A S
How does this relate to sexual selection?

G B
Thinking about the underlying purpose of animal colouration and display, the modern theory of sexual selection actually has more to do with Wallace’s ideas than Darwin’s. Everyone says that Darwin came up with the theory of sexual selection and Wallace rejected it, but if they knew enough about the modern theory, it’s actually quite the reverse!  

A S
Can you parse these two theories?

G B
Darwin’s theory says that the females of a species have an appreciation of beauty and that they pick the most beautiful males to mate with because they deemed them to be beautiful. Wallace couldn’t imagine that a butterfly would have an aesthetic sense. Why would a tiny insect brain be able to judge beauty in this way? He couldn’t understand how female butterflies could choose more beautiful males, so he argued against Darwin’s idea, which suggested in essence that these creatures knew what beauty was and chose it for its own sake. Wallace’s idea was that the plumage, for example, had some other function. It was basically the most vigorous males who were able to produce the best plumage, which was a sign of vigor and health. So, by choosing the best plumage the females were choosing the healthiest males. Or, in the case of antelopes, say, it would be the males with the biggest horns who were chosen by the females because they knew their offspring would have those characteristics. That’s what the modern theory of sexual selection is all about, which follows from Wallace’s “better genes” argument for selection, as opposed to the aesthetic sense idea from Darwin. In sexual selection, for Wallace, beauty is an index or register of health and vigor, not an aesthetic quality chosen for its own sake.

A S
To conclude, can you summarize what distinguishes Wallace from his contemporaries? Why is he so special in the history of science?

G B
He was poor and had no infrastructure. I think these things allowed him to develop a much deeper understanding of nature because he was so closely engaged with and attentive to the local culture and customs. When people they think of Wallace—if they think of Wallace at all—they think of the Wallace Line, or maybe even the co-discovery of natural selection, but his legacy goes far beyond that. His contributions to biology are very important, much more so than most of the other people in his day, such as T.H. Huxley or Charles Lyell, even. Only Darwin made similar contributions.

A S
What about Ernst Mayr?

G B
People will often suggest that Ernst Mayr came up with the biological species concept, but he actually took it from Wallace, who was the first to claim that species should be defined as interbreeding groups that  are reproductively separate from other such groups. There’s nothing in On the Origin of Species that actually explains what a species is. So, even though it’s a book about their origins, Darwin never defines what he means by species.

Wallace’s contributions to biology went far beyond co-discovering the theory of natural selection, upon which the modern study of evolution is based.³ Unlike Darwin, he rejected Lamarckism. In fact, he was the first natural scientist to reject it and was, in fact, the first Neo-Darwinian. Wallace devised the first modern species concept, a slightly modified version of what would later become known as the biological species concept. Interestingly, although many people think of sexual selection as Darwin’s theory, Wallace’s  argument about selection is regarded by many today as more plausible than Darwin’s belief that mates are chosen based on aesthetic grounds, as I said earlier. The Great American Interchange, when South America and North America connected via Central America and the animals of the north moved down, and vice versa—that was also Wallace’s idea. There were other things, too, like recognition marks in animals; a scientific paper about colour patterns in monkeys recently reinvigorated Wallace’s concept of recognition marks. He was also the founder of astrobiology, he came up with the first plausible evolutionary idea of aging and death, and he was first to propose mimicry in birds and polymorphism in butterflies. There was also the Wallace Line, of course.

A S
Do you think there is a way that the study of Wallace could contribute to the current discussion of the Anthropocene?

1 John van Wyhe, Dispelling the Darkness: Voyage in the Malay Archipelago and the Discovery of Evolution by Wallace and Darwin (Singapore: Scientific World, 2013).

2 See Alfred Russel Wallace, “Museums for the People,” The Alfred Russel Wallace Page, http://people.wku.edu/charles.smith/wallace/S143.htm.

3 Charles H. Smith and George Beccaloni, eds., Natural Selection and Beyond: The Intellectual Legacy of Alfred Russel Wallace (Oxford: Oxford University Press, 2010).