The Tangled Tree: A Radical New History of Life
The stability of species represented the bedrock of natural history. It was taken for granted, and important, not just among clergy and pious lay people but scientists too. That all the varied forms of creatures on Earth had been fashioned by God, in special acts of creation, and are therefore immutable, was an article of faith to the Anglican scientific establishment of Darwin’s era. This tenet is known as the special-creation hypothesis, though at the time, it seemed less hypothesis than dogma. It had been embraced and supported by prominent naturalists and philosophers of the scientific culture within which Darwin had been educated at Cambridge. He was now home from his wildcat voyage, a youthful adventure with a bunch of rough English sailors, about which his stern father had been skeptical at the start. The experience had altered him—though not in the ways his father may have feared. He hadn’t become a drunk or a libertine. He didn’t curse like a bosun. Darwin’s wanderlust, satisfied physically, was now intellectual. He intended to investigate, very discreetly, a radical alternative to scientific orthodoxy: that the forms of living creatures weren’t eternally stable, as God had created them, but instead had changed over time, one into another—by some mechanism that Darwin didn’t yet understand.
It was a risky proposition. But he was twenty-seven years old and deeply changed by what he had seen and, in a quiet way, very gutsy.
So he had set himself up in the big city, with lodgings on Great Marlborough Street, a convenient location for his visits to the British Museum. This was just a few doors down from the house where his elder brother, Erasmus, had already settled. Darwin joined scientific clubs, the Geological Society, the Zoological Society, but had no job. Didn’t need one. The same formidable father who had first disapproved of the Beagle voyage—Dr. Robert Darwin, a wealthy physician up in the town of Shrewsbury—was now rather proud of his second son, the young naturalist well regarded within British scientific circles. Grumpy on the outside, generous within, Dr. Darwin had made supportive arrangements for both brothers. And Charles was single. He sauntered around London, he handled follow-up tasks on his specimens from the voyage, he worked on rewriting his Beagle diary into a travel book, and—very privately—he ruminated about that radical alternative to special creation. He read widely, scribbling facts and phrases into various notebooks. The “A” notebook was devoted to geology. The B notebook was first of a series on what, to himself only, he called “transmutation.” You can guess what that meant. Darwin had begun thinking his way toward a theory of evolution.
He opened the B notebook, in July 1837, with a few phrases alluding to a book titled Zoonomia; or the Laws of Organic Life, published decades earlier by his own grandfather, another Erasmus Darwin. Zoonomia was a medical treatise (Erasmus was a physician), but it contained some provocative musings that sounded vaguely evolutionary. All warm-blooded animals “have arisen from one living filament,” according to Zoonomia, and they possess “the faculty of continuing to improve” in ways that could be passed down across the generations, “world without end!” Improvement across generations? Heritable change throughout the history of the world? That was contrary to the special-creation hypothesis, but not too surprising from a gouty, libidinous freethinker and sometime poet such as old Erasmus. Darwin had read Zoonomia during his student days and shown little sign of giving his grandfather’s daring ideas much credit. But now, on revisiting, he took them as a point of departure. Page one, entry one, in the B notebook: his grandfather’s title, Zoonomia, followed by reading notes.
Then again, those wild suggestions didn’t lead anywhere. Erasmus Darwin had offered no material mechanism for “the faculty of continuing to improve,” and a material mechanism was what young Charles wanted, though he may not have fully realized that yet. As reflected in the B notebook, he now went from his grandfather’s work to other readings, other speculations and questions, jotting down clipped phrases, often in bad grammar and punctuation. He wasn’t writing to publish. These were messages to himself.
“Why is life short,” he asked, omitting the question mark in his haste. Why is reproduction so important? Why do animals of a given kind tend to be constant in form across an entire country but to differ at least slightly on separate islands? He remembered the giant tortoises on the Galápagos, where his stopover had lasted only thirty-five days but catalyzed an upheaval in his thinking. He remembered the mockingbirds too. And why had he seen two distinct kinds of “ostriches” (his label for big, flightless birds now known as rheas) on the Argentine Pampas, one living north of the Rio Negro, one south of it? Did creatures somehow become different when isolated? Put a pair of cats on an island, let them breed and inbreed there for generations, with a little pressure from enemies, and “who will dare say what result,” Darwin wrote. He dared. The descendants might come to look different from other cats, might they not? He wanted to understand why.
Another important question: “Each species changes. does it progress.” Do the cats become better cats, or at least better cats for catting on that particular island? If so, how long would it take? How far would it go? What are the logical limits, if “every successive animal is branching upwards” and with “different types of organization improving,” new forms arising, old forms dying out? That one word, branching, was freighted with interesting implications: of directional growth, of divergence, of an arboreal form. And these questions Darwin asked himself, they applied not just to cats and ostriches but also to armadillos and sloths in Argentina, to marsupials in Australia, to those huge Galápagos tortoises, and to the wolflike Falkland Islands fox, all peculiar in certain ways, all unique to their isolated places, but recognizably similar to their correlatives—other cats and tortoises and foxes, etcetera—elsewhere. Darwin had seen a lot. He was an acutely observant and reflective young man. He sensed that he had seen patterns, not just particulars. It almost seemed, he wrote, that there was a “law of adaptation” at work.
All this and more, facts and speculations, crammed into the first twenty-one pages of notebook B. The pages are mostly undated, so we can’t know how many days or weeks passed in the opening burst of effort. Anyway, he didn’t yet have his theory. Big ideas were coming at him like diving owls. He needed some order as much as he needed the jumble of tantalizing clues. Maybe he needed a metaphor. Then, on the bottom of page 21, Darwin wrote: “organized beings represent a tree.”
2
We don’t know whether Darwin sat back after writing that statement and breathed deep with a new sense of clarity, but he might have. And he was entitled.
Then he scribbled on. The tree is “irregularly branched,” he told the B notebook, “some branches far more branched.” Each branch diverges into smaller branches, he wrote, and then twigs, “Hence Genera,” the next higher category above species, which would be the twiglets or terminal buds. Some buds die away without yielding further growth—species extinction, end of a line—while new buds appear, somehow. Although the very idea of extinction had once been problematic among naturalists and philosophers, doubted as a possibility or rejected outright on grounds that God’s acts of special creation couldn’t be undone, Darwin recognized that there’s “nothing stranger in death of species” than in death of an individual. In fact, extinction was not just natural but necessary, making space for new species as old ones die away. He wrote: “The tree of life should perhaps be called the coral of life, base of branches dead,” ancestral forms gone. Darwin knew something about coral, having seen reefs at Keeling Atoll in the eastern Indian Ocean and elsewhere during the Beagle voyage. They fascinated him; he concocted a theory of how reefs are formed; and in 1842, five years after this notebook entry, he would publish a book about coral reefs. Coral seemed apt—branching coral, not brain coral or table coral, was what he had in mind—because the lower limbs and base are lifeless calcitic skeleton, left behind like extinct forms of ancient lineages as the soft polyps advance upward like living species. But even he seems to have sensed that “the coral of life” didn’t have the same memorable ring. He drew a feeble pen sketch, on page 26 in the B notebook, of a three-branched coral of life, with dotted lines depicting the inanimate lower sections. And then he let the coral idea slide, abandoning that metaphor.
The tree of life was better. It was already a venerable notion in 1837, and Darwin could adapt it to his purposes as an evolutionary theorist—easier than inventing a new trope from scratch. Of course, to make that adaptation was to alter its meaning radically. Never mind, he took the step. Ten notebook pages along, he sketched a much livelier and more complex figure in bold strokes, with a trunk rising into four major limbs and several minor ones, each major limb diverging into clusters of branches, one branch within each cluster labeled A, B, C, D. The branches B and C were near neighbors in the treetop, within adjacent clusters, indicating close relationships among the creatures on those branches. The letter A was far away, on the opposite side of the tree’s crown, signaling a more distant relationship—but still a relationship. The letters were placeholders, meant to represent living species, or maybe genera. Felis, Canis, Vulpes, Gorilla. We don’t know exactly what he had in mind, and maybe it was nothing so specific. Anyway, this was a thunderous assertion, abstract but eloquent. You can look at the little sketch today, with its four labeled branches amid the limbs and the crown, and imagine the evolutionary divergence of all life from a common ancestor.
Darwin’s 1837 sketch, redrawn by Patricia J. Wynne.
Just above the sketch, as though gesturing toward it bashfully, Darwin wrote: “I think.”
3
Darwin didn’t invent that phrase, “the tree of life,” nor originate its iconic use, though he put it to new purpose in his theory. Like so many other metaphors embedded deep in our thinking, it came down murkily, modified and reechoed, from early versions in Aristotle and the Bible. (Why do these things always go back to Aristotle? Well, that’s why he’s Aristotle.) In the Bible, it’s a grand bookend motif, invoked in Genesis 3 just as Adam and Eve are booted out of the Garden, and reappearing at the end of Revelation, on the very last page of the King James version—excellent placement for a launch into Western culture. There in Revelation 22, verses 1–2, the authorial prophet describes his ecstatic vision of the “water of life,” flowing out like a pure river from the throne of God, and beside which grows “the tree of life,” bearing fruit every month, plus leaves “for the healing of the nations.” This tree possibly represents Christ, supplying his leafy and fruity blessings to the world; or maybe it’s grace, or the Church. The passage is opaque, and differences in translations (one tree or many?) have confused things further. The point here is simply that the “tree of life” is an ancient poetic image, a resonant phrase, variously construable, with a long presence in Western thought.
In Aristotle’s History of Animals, written during the fourth century BCE, the tree of life is not yet a tree. It’s more like a ladder of nature or—as later Latinized from his Greek—a scala naturae. According to Aristotle, the diversity of the natural world “proceeds” from lifeless things such as earth and fire to living creatures such as animals “little by little,” in a progression so incremental that it’s impossible to draw absolute lines between one form and another. This idea remained useful throughout the Middle Ages and beyond, turning up in woodcuts during the sixteenth century as a Great Chain of Being or a Ladder of Ascent and Descent of the Intellect, which typically rose step-by-step from inanimate substances such as stone or water, to plants and then beasts, then humans, then angels, and finally to God. By that point it was a “Stairway to Heaven,” almost five centuries before Led Zeppelin.
The Swiss naturalist Charles Bonnet reverted to this linear, stair-step model as late as 1745, even while other Enlightenment thinkers and artists were allowing images of nature’s diversity to burgeon sideways with limbs and branches. Bonnet’s treatise on insects, published that year, included a foldout diagram of his “Idea of a Scale of Natural Beings,” arranged in vertical ascent from fire, air, and water, through earth and various minerals, upward to mushrooms, lichens, plants, and then sea anemones, followed by tapeworms and snails and slugs, upward further to fish and then flying fish in particular, and then birds, above which came bats and flying squirrels, then four-legged mammals, monkeys, apes, and lastly man. See the logic? Flying fish are superior to other fish because they fly; bats and squirrels exist on a higher level than birds because bats and squirrels are mammals; orangutans and humans are the best of mammals, and humans are more best than anybody. Bonnet made his living as a lawyer but much preferred studying insects and plants. He was a lifelong citizen of the Republic of Geneva, his French ancestors having been chased out of France by religious persecution, and so maybe it’s no accident that his ladder diagram culminated in people, not God.
The other notable absence from Bonnet’s scale of natural beings, besides God, are microbes. He paid no attention to microorganisms, although the pioneering Dutch microscopist Antoni van Leeuwenhoek had discovered the existence of bacteria, protozoans, and other tiny “animalcules” about seventy years earlier. We all know Leeuwenhoek’s name from our reading in high school of Paul de Kruif’s Microbe Hunters (a terrible book full of concocted dialogue and bogus detail, but an influential doorway to the subject) or other storybook histories of science, though we might not remember that Leeuwenhoek was a draper in Delft who started making his own magnifying lenses in order to better inspect the thread-count of textiles. Then he turned the lenses onto other materials, out of sheer curiosity, and made astonishing discoveries: he found menageries of tiny creatures living in lake water, in rain water, in water from drain pipes, even in scrapings of crud from his own teeth.
Leeuwenhoek’s revelatory observations of microbial life were reported in the journal of the Royal Society of London and became famous in scientific circles throughout Europe, but Charles Bonnet wasn’t interested enough in those “very wee animals” to fit them into his rising scale—not even where they might dismissively have been slotted, somewhere between asbestos and truffles. That omission presages a lasting discomfort with placing microbes on the ladder of life or, harder still, arranging their diverse forms on the tree—and it’s a discomfort to which I’ll return, because it became acute in 1977.
The linear approach to depicting life’s diversity was on the way out, notwithstanding Charles Bonnet’s scale of nature, and being replaced by its more complicated and dimensional successor, the tree. By the late eighteenth century and the start of the nineteenth, natural philosophers (we’d call them scientists, but that word didn’t yet exist) tried to classify and arrange living creatures into distinct groups and subgroups, reflecting their similarities and differences and some sort of organizing schema. The linear alignment, in order of what passed for increasing sublimity, the ladder raised toward God, was no longer satisfactory. There had been a knowledge explosion in Europe since the great age of sailing explorations began—knowledge of diverse animals, plants, and other creatures from all over the world—and scholars wanted to set that explosive abundance of new facts within hierarchical categories so that it could be easily accessed and used.
This wasn’t evolutionary thinking; it was just data management. The knowledge would fill volumes (one man alone, the German naturalist Alexander von Humboldt, published a thirty-volume account of his travels in South America), making all the more necessary an overview, an organizing principle, that could be apprehended at a glance: an illustration. But the illustrators now needed two dimensions, not one, and so the ladder turned into a trunk, and the trunk sprouted limbs, and the limbs diverged into branches. This offered more scope, sideways as well as up and down, for arranging the varied abundance of known creatures.
The tree of life was an old symbol by then, an old phrase, dating back at least to those mentions in Genesis and Revelation. The tree had also served as a model for family histories—the genealogical tree or pedigree of a German duke, for instance. Now the secularized tree became useful for organizing biology. Among the first to embrace this convention was another Frenchman, Augustin Augier, who wrote in 1801 that “a figure like a genealogical tree appears to be the most proper to grasp the order and gradation” of what concerned Augier: the diversity of plants.
Augier was an obscure citizen of the French Republic, living in Lyon, working on botany part-time; his real profession was unknown, his biographical details lost, even to a historian of Lyonnais botanists writing a hundred years later. Augier disappeared. But he left behind a book, a little octavo volume, in which he proposed a new classification of plants, “according to the order that Nature appears to have followed.” That is, a “natural order,” as opposed to an artificial classification system based merely on human whim or convenience. And in the book was a figure representing that system: his arbre botanique (botanical tree). Its trunk and limbs look almost as orderly and stiff as a menorah, but its sideways branching and copious leafing suggest a rife multiplicity of plant forms.
Again, this didn’t imply any heretical ideas about origins. Augier was no evolutionist before his time. His natural order wasn’t meant to suggest that all plants had descended from common ancestors by some sort of material process of transformation. God was their maker, shaping the varied forms individually: “It appears, and one can hardly doubt it, that the Creator, when making flowers, followed certain proportions and progressions in the number of their different parts.” Augier’s contribution, as he saw it, was discovering those proportions and progressions—design principles that had satisfied the Deity’s neat sense of pattern—and using them after the fact to organize botanical knowledge into a tidy system.
Augier wasn’t the first naturalist to hanker for a natural order of nature’s diversity. Aristotle had classified animals as “bloodless” and “blooded.” In the first century of our era a Greek physician named Dioscorides, attached to the Roman army, gathered lore on more than five hundred kinds of plants, arranging them in a compendium mainly on the basis of their medicinal, edible, and perfumatory uses. That book, in various reprints and translations, served as a trusted botany text for the next fifteen hundred years. Toward the end of its run, around the time of the Renaissance, as people traveled more widely and paid closer attention to the empirical details of nature, old Dioscorides gave way to newer illustrated herbals. These were essentially field guides to botany, graced with better illustrations based on improvements in drawing and woodcut techniques, but still organized for convenience of use, not natural order. In the sixteenth century, Leonhart Fuchs produced one of those books, an herbal cataloging hundreds of plants, beautifully illustrated and arranged in alphabetical order. Two centuries later, the great systematizer Carl Linnaeus described a genus of plants with purplish red flowers, naming it Fuchsia in honor of Leonhart Fuchs (and hence we got also the color, fuchsia). Linnaeus himself, a Swede who traveled widely as a young man and then took up a professorial life in Uppsala, emerged from this herbalist tradition but went beyond it.
Augier’s Arbre Botanique, 1801.
Linnaeus’s Systema Naturae, as first published in 1735, was a unique and peculiar thing: a big folio volume of barely more than a dozen pages, like a coffee-table atlas, in which he outlined a classification system for all the members of what he considered the three kingdoms of nature: plants, animals, and minerals. Notwithstanding the inclusion of minerals, what matters to us is how Linnaeus viewed the kingdoms of life.
His treatment of animals, presented on one double-page spread, was organized into six columns, each topped with a name for one of his classes: Quadrupedia, Aves, Amphibia, Pisces, Insecta, Vermes. Quadrupedia was divided into several four-limbed orders, including Anthropomorpha (mainly primates), Ferae (doggish forms such as wolves and foxes, plus cat forms such as lions and leopards, in addition to bears), and others. His Amphibia encompassed reptiles as well as amphibians, and his Vermes was a catchall group containing not just worms, leeches, and flukes but also slugs, sea cucumbers, starfish, barnacles, and other sea animals. He divided each order further into genera (with some recognizable names such as Leo, Ursus, Hippopotamus, and Homo), and each genus into species. Apart from the six classes, Linnaeus also gave a half column to what he called Paradoxa: a wild-card assemblage of mythic chimeras and befuddling but real creatures, including the unicorn, the satyr, the phoenix, the dragon, and a certain giant tadpole (Pseudis paradoxa, under its modern label) that, strangely, paradoxically, shrinks during metamorphosis into a much smaller frog. Across the top of the chart ran large letters: CAROLI LINNAEI REGNUM ANIMALE. His animal kingdom. It was a provisional effort, grand in scope, integrated, but not especially original, to make sense of faunal diversity based on what was known and believed at the time. Then again, animals weren’t Linnaeus’s specialty.
Plants were. His classification of the vegetable kingdom was more innovative, more comprehensive, and more orderly. It became known as the “sexual system” because he recognized that flowers are sexual structures, and he used their male and female organs—their stamens and pistils, those delicate little stems sticking up to present and receive pollen—for characterizing his groups. Linnaeus defined twenty-three classes, into which he placed all the flowering plants, based on the number, size, and arrangement of their stamens. Then he broke each class into orders, based on their pistils. To the classes, he gave names such as Monandria, Diandria, and Triandria (one husband, two husbands, three husbands), and, within each class, ordinal names such as Monogynia, Digynia, and Tryginia (numbers of wives, yes, you get the idea), thereby evoking all sorts of polygamous and polyandrous ménages that must have caused lewd smirks and disapproving scowls among his contemporaries. A plant of the Monogynia order within the Tetrandria class, for instance: one wife with four husbands. Linnaeus himself seems to have enjoyed the sexy subtext. And it didn’t prevent his botanical schema from becoming the accepted system of plant classification throughout Europe.
Our man Augustin Augier, coming along a half century later with his botanical tree of classification, seems to have seen himself challenging Linnaeus’s overly neat sexual system. “Stamen number is a striking character,” Augier conceded, but “not when it comes to the examination of plants”—that is, not always unambiguous and therefore not reliable as a basis for organizing the great jumble of botanical life. He nodded respectfully to Linnaeus—also to the French botanist Joseph Pitton de Tournefort, who had sorted plants into roughly seven hundred genera based on their flowers, their fruits, and other bits of their anatomy—and offered his own system, using multiple characters for different levels of sorting and to resolve the ambiguities and fine gradations. “This figure, which I call a botanical tree, shows the agreements which the different series of plants maintain amongst each other, although detaching themselves from the trunk; just as a genealogical tree shows the order in which different branches of the same family came from the stem to which they owe their origin.” All discrete, yet all connected: bits of the same tree.