Posts Tagged ‘Evolution’

The Galapagos Islands

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Wednesday, April 25th, 2012

Gene with a statue of Charles Darwin

The Galapagos, an archipelago over a thousand miles from the west coast of Ecuador, is a strange, contradictory place. Extensive lava formations and exotic plants contrast with beautiful, quiet, sandy beaches and harbors full of yachts. Snorkeling and scuba diving attract young adventurers. For others it’s a more mythic place – with huge land-tortoises, creepy black iguanas, misplaced penguins, stinky seals, porpoises, and schools of rays. Limited numbers of unique birds are a bonus.

Two “celebrities” dominate Galapagos publicity: Charlie Darwin and Lonesome George.

Lonesome George is a huge Galapagos Tortoise who is suspected to be the last surviving member of his subspecies and “the world’s rarest creature”. But he was in a dark corner of his zoo enclosure, as uninterested in me as he is, evidently, in sex. His near relatives crawl freely if slowly in meadows and cow pastures. They also engage in extremely slow-motion tortoise sex, complete with guttural wheezes and moans.

Large Tree-Finch © Renato Espinosa

Darwin visited the Galapagos Islands in 1835; there’s a large statue of him, splattered with guano from boobies and frigate birds, near where he first set foot. From his observations of four different kinds of mockingbirds on separate islands, he is said to have first devised his theory of evolution. All four mockers originated from an Ur-mockingbird couple and evolved differently.

More interesting are the 13 (or 14, depending on how you split them) finches that are now the islands’ main birding attractions. These “Darwin’s finches” also evolved, i.e., changed their shapes and behavior as necessity dictated. Ground Finches, Tree Finches, Warbler Finches, Cactus Finches, and Woodpecker Finches: they’re all just slightly different. Even though they’re as tame as chickens, coming fearlessly with arm’s reach, accurately identifying some of them drove me nuts!

Woodpecker Finch © Renato Espinosa

Fewer than 150 species are on the checklist for Galapagos birds. Of these, only 23 are endemic – birds you can’t see anywhere else. Put “Galapagos” in front of the following species names, and you’ll get some idea of the variety of endemics: Penguin, Dove, Hawk, Flycatcher, Mockingbird, Rail, Martin. All are stuck on the Galapagos and worth stalking and ticking.

But some of the birds you can see elsewhere (Yellow Warbler, Barn Owl and Short-eared Owls, for instance) are slightly different from their continental cousins. They’re stuck here too, and better off because of it. Unable to migrate, Yellow Warblers on Galapagos, for instance, are slightly bigger, more colorful, and more robust in their singing than the ones that expend vast amounts of energy getting to North America.

Barn Owl © Renato Espinosa

Perhaps they’ll continue to evolve. If so, I’ll be able to add Galapagos Yellow Warbler and a dozen or so other species to my life list some day – provided I live to be 10,000 years old. Wait; maybe global warming will make evolution speed up a bit.

Shaping Our Understanding: Part III

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Wednesday, September 7th, 2011

One interesting implication of the whales being related to the hippopotamus is that intelligence must have evolved several times. Most researchers would define the whales and dolphins as among the most intelligent of mammals, and very few would put the hippo high on that scale. Intelligence is extremely complex and is certainly not a single-gene trait (nobody knows how many genes are involved, and it is very likely that different genes are involved in different intelligent species). However, the intelligent mammals are not especially closely related to each other, and no matter how you define intelligence, you can’t group all the intelligent mammals together under a single ancestor. Very few people would argue that “intelligent mammals” doesn’t include at least include some carnivores (wolves are perhaps the best example, but many carnivores are intelligent), many cetaceans and many or most primates. Many people would also include the elephant and some other species among the intelligent mammals. Those species are scattered all over the mammals, and are more closely related to animals nobody would consider intelligent than they are to each other, strongly suggesting that their intelligence is separately evolved. We are actually probably less closely related to dolphin and dog than we are to mice and rats (and dolphins and dogs aren’t close to each other, either).

Other animals, entirely apart from mammals, are also often considered intelligent. The farther one gets from the type of intelligence we are most familiar with, the tool using, collaborative, highly communicative intelligence of the primates, the harder it becomes for humans to define intelligence. It is easy for a human to see that an ape or a monkey is smart, because they are smart in fundamentally the same ways that we are. A dog or a dolphin is a little harder to see as “like us”, but there is still enough commonality between the ways humans, dogs and dolphins think about things that we can devise intelligence tests that work across that barrier, and, perhaps more importantly, we can see them (and they us) as kindred, intelligent spirits.

As we get farther away from ourselves on the evolutionary tree, this becomes harder. Some birds are highly intelligent, yet the mind of the raven is much harder for humans to penetrate than that of the chimpanzee, the dog, or even the entirely aquatic dolphin. Parrots, with their capacity for vocal mimicry, make it easier for us to see them as similar to us – they can speak our languages, and, it sometimes appears, know what they’re saying. It appears to many researchers that the mimic capability of the parrots goes beyond being a biological “tape recorder” to beginning to understand a few phrases of human language. Even so, the way birds think and emote is exotic to us in a way that similar features of mammals are not.

Yet another step away from us is the intelligence of the molluscs. By many definitions, octopus and squid are highly intelligent organisms, yet their form of intelligence is so alien to us that we share far less with them than we do even with the birds. While some scientists have said that an octopus is roughly as intelligent as a house cat, how can we ever know? A cat shares many experiences with us, even if it is a wild animal like a bobcat that has never lived with people. A bobcat lives fundamentally in our world, and, like most intelligent mammals and birds, takes care of its young and teaches them much of what they need to know. This touchstone, behavior that is learned from parents, older relatives and even non-relatives, is one thread that connects the intelligent mammals and birds – all of them (unlike many less intelligent mammals and birds) have long periods of parental care, and all seem to have some form of teaching and learning. A mouse is born knowing how to be a mouse, but a wolf isn’t born knowing how to be a wolf, or a raven how to be a raven.

The octopus is so alien to this world that it breeds once in its lifetime, lays eggs and then dies before the eggs ever hatch. Octopi DO seem to learn, but they learn strictly through experiment, because there is no older generation around to teach them. They are also almost entirely solitary, and communicate none of the knowledge each individual gains back to the group in any way. Humans, of course, are the exact opposite, with an intelligence based almost entirely on building on knowledge of past generations. Since the invention of writing, the human capacity to build and store knowledge has far outstripped that of any other species, but there are other examples of similar phenomena. Orca (so-called killer) whales have perhaps the most sophisticated nonhuman culture, with food preferences, vocal dialects and behaviors that are entirely learned, and that go back many generations in what appears to be a rich trove of lore that is completely different between populations. In some places, two populations of orcas can live alongside each other and barely interact – they’re biologically the same species, but have grown far enough apart culturally that they don’t interbreed, compete for prey, or even really interact (despite interactions within each population that are among the most complex of all creatures’). Compare this to the octopus, whose breeding system means that, by definition, no culture can exist. Each generation is starting over from scratch.

Maybe even more alien to us than the intelligence of the octopus is the intelligence of social insects. Is an ant “smart”? Not by any conventional definition – it is a fairly simple creature with a small number of entirely instinctual behaviors. An individual ant is no more or less intelligent than an individual mosquito by most definitions, yet an ant colony is capable of numerous behaviors we would characterize as intelligent. Some ant colonies can farm, others are capable of organized warfare, and simulated ant colonies are even used to determine efficient ways to route information. No individual member of the group has any intelligence to speak of, but the group as a whole behaves intelligently.

Intelligence is just one example of the power of evolution. Since evolutionary processes are essentially random, yet selective pressures produce highly successful organisms (if a given combination of traits is detrimental to survival, there are enough mutations around that something else will replace it). On the other hand, because the process has a lot of randomness, many different “solutions” to the same problem are likely to appear. It would be very difficult to fit a brain that was conventionally intelligent in an organism the size of an ant, but a colonial intelligence gives ants an advantage over solitary insects in many circumstances. There is almost certainly no relationship between the “intelligence” of social insects and that of birds or mammals, yet they convey many of the same advantages.

If you have enjoyed my series on taxonomy and evolution, some longer works worth reading are:

Anything by the great biologist and natural history essayist Stephen Jay Gould – numerous collections of essays, many on topics I’ve touched on.

On the Origin of Species by Charles Darwin – yes, it was written in 1859, and the language is not that of modern science, but it is amazing to see how well the Origin has stood up as a work of science, not just for its historical importance.

Shaping Our Understanding Part II

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Thursday, August 25th, 2011

In modern biological thinking, a valid clade (any grouping larger than a species, be it a genus, family, order, class, phylum, kingdom or some variation) is any group of an ancestor and all of its descendants. The ancestor can be chosen arbitrarily, but there are two rules. First, a clade cannot consist of some of the descendants of a given ancestor, with others excluded. Second, a clade cannot consist of descendants of two or more ancestors, unless they are the descendants of another ancestor further back (and the clade includes all descendants of that ancestor).

As an example, the dog family seems to be a valid clade according to current research. It contains dogs, coyotes, wolves, foxes and several other animals that are less familiar to us. There was a “proto-dog” that was the most recent common ancestor of all of these animals, that was not also the ancestor of any other animal. Within this clade, there is a smaller clade (equally valid) that consists of the genus Canis (domestic dogs, wolves and the coyote). The common ancestor of this clade is more recent than the ancestor of the larger family clade, and is an ancestor of dogs, wolves and the coyote, but NOT foxes or jackals. Since a clade can be defined from any ancestor, both are equally valid, and the smaller clade, corresponding to the Linnaean genus Canis, nests completely within the larger one, which corresponds to the Linnaean family Canidae.

Until the recent arrival of molecular techniques, scientists going back to Linnaeus had classified species based on appearance, although often basing classifications on highly technical details that required a microscope to see. To a lessor extent, behavior, especially among animals with complex behaviors, has influenced classification. Molecular techniques permit us to read the underlying genetic code directly, instead of making our best guess based on physical appearance and similar factors.

The dogs are a good example of a place where the molecular classification agrees with the means of classification used before the development of molecular techniques. Linnaeus was the first to describe both the family Canidae, and the genus Canis in their modern sense, both of which, as we’ve seen, are valid molecular clades.
In other cases, though, the genetic code gives us some surprising results – organisms that appear related turn out not to be, or those that appear very different are actually relatives. Examples of surprising clades are appearing throughout life as molecular data become more available for a wide range of species.
To demonstrate the effects of molecular taxonomy, I’ll choose a couple of favorite examples – a class, and an order within it, that make a great deal of sense from their appearance, but turn out to be partial clades (and the additional members to complete the clade are very surprising), and an order that fits in a surprising spot on the tree of life.

The clades with surprising members are the reptiles, and the lizards within them. There’s no such thing, biologically, as a reptile, and this is a recent discovery. If you think about “reptile”, most of us can come up with a large number of animals that fit the bill. Lizards, snakes and turtles all come to mind quickly, and with only a little more thought, we can add crocodilians and dinosaurs. No valid clade joins all of these animals without a significant number of surprising “extra members”. The lizards and snakes share a common ancestor and form a valid clade, there is another clade composed of turtles, and another with alligators and crocodiles. Another clade holds all the dinosaurs, but also contains all birds. The dinosaur/bird clade joins with the crocodile clade, then reaching for a more ancient ancestor, this clade joins with the lizard/snake clade. The turtles are less closely related to any of these animals than any of them are to each other. When we think about core reptiles, without having to include birds (which the average person doesn’t consider to be reptiles), all we get are lizards and snakes. Any attempt to add the crocodiles pulls both dinosaurs and birds along with them, and the turtles are so distantly related to anything else that any clade containing both turtles and lizards also contains crocodiles, snakes, dinosaurs and birds.

Within this broad group of reptiles, there are two surprises. One is that the birds are quite deeply nested within the dinosaurs. Birds and the classic carnivorous dinosaurs like Velociraptor and Tyrannosaurus rex are more closely related to each other than either one is to plant-eating dinosaurs such as Triceratops. The next time you see a Robin, realize that, in a real sense, you’re looking Velociraptor in the eye (OK – maybe it’s easier to see with a Bald Eagle).

The other surprise in the reptiles is that you can’t get a clean lizard clade. There are several clades composed entirely of what would commonly be called lizards, but any attempt to find a common ancestor that unifies them also includes the snake clade. All snakes do form a clade (all snakes are more closely related to all other snakes than any snake is to any lizard), but some “lizards” are more closely related to snakes than they are to other “lizards”, making a lizard clade that doesn’t also include the snakes impossible to define.

Moving over to the mammals (which are a valid clade with no surprising members), the whales have long been a bit of a mystery. Where do they fit within the mammals? Are they even a single clade, or are the baleen whales not all that closely related to the toothed whales? Prior to the arrival of molecular evolutionary biology, the whales had been placed all over the mammals, generally together but occasionally separated. The molecular evidence shows that the closest living relative of the whales (which are a single clade) is the hippopotamus.

Shaping Our Understanding

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Wednesday, August 17th, 2011


Last week’s entry covered how scientific names work and describe a given organism’s place in the evolutionary framework. This week, I’ll build both back and forward on that, going from the origins of the system to what modern molecular genetic research has told us about how life fits together (there are a few surprises)! We’ll look briefly at the careers of several biologists, from the 18th century to the present day, who have shaped our understanding of life.
We actually need to start more than a century before Darwin published On the Origin of Species, with Swedish physician and botanist Carolus Linnaeus (sometimes written as Carl Linnaeus or Carl von Linne). Linnaeus’ great contribution to biology was his two books Systema Naturae (1st edition 1735, but the influential edition is the 10th, of 1758), and Species Plantarum (1753), which introduced the binomial (genus and species) method for naming animals and plants, respectively. Linnaeus is also responsible for a significant portion of the system of names above the genus level, although some of those levels are later additions.
In addition to having invented the system of nomenclature we use today, Linnaeus named approximately 7,700 species of plants and 4,400 species of animals. Many of his names are still in use today, including such familiar organisms as the Harbor Seal (Phoca vitulina), the Red Maple (Acer rubrum) and even ourselves (Homo sapiens) – in 1959, his successors honored Linnaeus by designating him as the lectotype for humans – the formal description of human beings is “the animal species that includes Linnaeus”.
While Linnaeus was a great field biologist, leading expeditions throughout Sweden for most of his life, he rarely left Sweden, and never during the period when he classified thousands of species. Most of his classifications were based on material his students sent him from around the world. Students of Linnaeus traveled to an amazing variety of places (several sailed with Captain Cook, for example) and sent back specimens for Linnaeus to classify and name. He also came amazingly close to evolution without ever understanding it, and, in fact steadfastly denied the possibility that any sort of evolution could exist. Looked at through an evolutionary lens, it seems amazing that anyone could have completed this brilliant work of classification without becoming aware that organisms were related to each other. Linnaeus placed humans, monkeys and apes together in the Primates, while always maintaining that each species was separately created in an unrelated event, and that his system was merely a method of organization – he was famous for saying “God created, Linnaeus organized”. Linnaeus came amazingly close to evolution several times without actually seeing it – in addition to correctly grouping humans with other primates for the first time, he was also the first to recognize that both bats and whales are mammals. He seems in all those cases to have been acutely aware of biological relationships, yet unable to make the mental leap from separate creation to an evolutionary paradigm that would permit these species to actually BE related.
After Linnaeus designed the modern system of biological classification and hugely expanded the number of known species, many scientists began to wonder whether each species was, in fact, a separate creation. While evolutionary ideas were beginning to be discussed by the beginning of the 19th century, nobody had worked out a system by which evolutionary change could occur. It was thought that since the descendants of any given species were always of the same species, the species were immutable. The chicken and egg problem is the classic example of this – a chicken is what lays an egg, but is also what hatches from one, so how does a non-chicken ever lay a chicken-producing egg? The answer to this was uncertain until the middle of the 19th century, and was the fundamental impediment to a workable theory of evolution.
In 1831, a young naturalist named Charles Darwin set off on a long sea voyage aboard HMS Beagle, a survey ship of the Royal Navy. Darwin was actually aboard primarily as a social companion to Captain FitzRoy, because English captains of the time were of a much higher social class than their crews, and forbidden to socialize with them. Darwin’s secondary function aboard the Beagle was to keep notes on geology and natural history, a position in which he excelled, and which would later bring him fame. As Darwin traveled around the world observing the plants and animals, he was able to see relationships between them – the data point that was directly in front of Linnaeus and many other earlier biologists, but that none had satisfactorily explained. It took him over 20 years from the first rough sketches of the idea of evolution to the publication of On the Origin of Species, a timeframe that seems less surprising when you consider that the Origin is so beautifully thought out and written that it is still referred to today, 152 years after its first publication in 1859.
Starting while he was still aboard the Beagle on the five-year voyage, and refining his ideas over many years at home in England, Darwin developed the theory of evolution by natural selection. Different members of a species are not identical – they possess slight, but important variations. According to Darwin, and to 150 years of biology since he published On the Origin of Species, individuals which have variations favorable to survival will be more likely to survive to reproductive age, and to leave offspring behind, which will carry the (initially randomly occurring) variation. This view of species as essentially mutable – individuals vary, and those which have variations which benefit their survival will leave more offspring, enhancing their contribution to the next generation – is diametrically opposite to Linnaeus’ view that each species is perfect and unchanging, created by God for a specific purpose.
In Darwin’s view, species diverge from a common ancestor when any of several circumstances occur. Two populations of the same species could become separated (one moves to a new habitat), where different variations lead to better survival. The variations are present in some members of both populations, but one form leads to better survival in one location, while a different form is advantageous in the other. Eventually, in the absence of contact between the populations, they might diverge into separate species. Alternatively, a change in environment for part of a population might mean that a different set of variations was most successful, leading to a distinct population that may eventually diverge far enough to form a new species. A third possibility is that species could diverge through competition with each other, rather than through environmental change. For example, a bird with a mid-sized beak could evolve towards a small-beaked and a large-beaked species even in the absence of environmental segregation, if the smallest-beaked birds could take advantage of one food source, and the largest-beaked ones another, while a bird with a mid-sized beak was a disadvantage in exploiting either source. All of these possibilities have been confirmed by research over the last 150 years.
Darwin built on Linnaeus’ classification of species, bringing relatedness to Linnaeus’s order. A Linnaean view would say “these two species in the same genus are similar, therefore they belong next to each other, like library books on a shelf”. Darwin found the mechanism to say “they’re not just similar, they’re related to each other and share an ancestor, and that’s why they’re next to each other”.
Darwin’s great insights were the importance of subtle variations and time. Until shortly before Darwin, the Earth was believed to be only a few thousand years old, leaving time for a fairly limited number of generations of most species. Geologists in the half-century before Darwin were revising the age of the Earth steadily upward, giving substantially more time for change to occur – in Darwin’s day, there wasn’t an agreed-upon age of the Earth, but estimates were from hundreds of thousands of years to hundreds of millions or more (the currently accepted estimate is slightly over 4.5 billion years).
An older Earth gave time for subtle variations to accumulate – a non-chicken didn’t have to lay an egg from which a chicken hatched. Over many generations, a near-chicken could lay an egg that produced a slightly nearer-chicken, and, even today, a chicken doesn’t lay an egg that produces exactly the same chicken (as a matter of fact, from the standards of what Linnaeus first named Gallus gallus (a chicken), some of what shows up on today’s large poultry farms may be perilously close to being non-chicken, a different species).
While Darwin was formulating his grand idea, there was no concept of genetics in the intellectual discourse at all. People had long known that children tended to look like their parents (and that children who looked exactly like the wandering peddler who passed through town nine months before their birth indicated something had gone on). Nobody had provided any mechanism for why this was true, however.
At almost exactly the same time Darwin was working out the mechanisms of evolution, an Austrian monk named Gregor Mendel was studying heredity. While Darwin was studying the world as it is on a broad scale, gathering ideas from a round-the-world voyage, Mendel was the great small-scale experimentalist. He planted thousands of pea plants over several years, keeping track of the parents of each plant, and of the characteristics of the offspring. Mendel worked out how characteristics are passed from generation to generation.
Mendel was the first to discover that every organism receives one copy of each gene from each parent, and that it is random which of a parent’s two copies a given offspring receives. He also developed the concepts of dominant and recessive traits – a dominant trait appears if you get at least one copy of its gene (from either parent), while a recessive trait is only visible if each parent contributes a copy of its gene (if one parent contributes the dominant version of the gene, and the other the recessive, the offspring will have the dominant appearance, not a blend of the two). Mendel had worked out most of the basic math of inheritance, at least for traits that are controlled by a single gene (many complex traits are influenced by several genes, which makes the situation more difficult).
Unfortunately, and unlike Darwin, who was one of the best-known scientists of the 19th century within his lifetime, Mendel’s work was largely forgotten until well after his death. He presented his work as primarily about hybridization, without going into the much larger implications of inheritance in general. As a monk, and later abbot of his monastery, he was not able to give his scientific work the wide publicity it deserved. Other scientists dismissed Mendel as a monk breeding peas in a garden, not realizing until many years after his death that he had discovered the first principles of the modern science of genetics.
Mendel’s genetics was promptly forgotten about for 35 years, as Darwin’s evolution raised firestorms of criticism, largely from a religious establishment that did not want to hear that humans are close relatives of apes, preferring instead to view humans as direct descendants of God. By Darwin’s death in 1882, the battle was essentially over, at least in scientific circles. Just about every biologist had accepted that Darwin was right – debate would continue, and still does, on the precise mechanisms of evolution, but Darwin’s basic idea of evolution through natural selection was, and still is, essentially unchallenged. Many religious scholars also came to accept Darwin’s ideas relatively quickly – he was buried at Westminster Abbey, with many of the senior clergy of Britain in attendance.
Mendel’s ideas would take much longer to achieve general acceptance – his great paper was published in an obscure location, and discussed at a local natural history society. Shortly afterwards, his work faded into obscurity, where it remained for 35 years. Around the turn of the 20th century, several biologists began to independently work out the laws of heredity, and, in the process rediscovered Mendel’s writings. It was quickly accepted that Mendel had, in fact, pioneered this work in the mid-19th century.
Through the first half of the 20th century, the thrust of evolutionary thought was applying Mendelian genetics to the study of Darwinian evolution, leading by the middle of the century to the understanding of evolution that, on a basic level, is current today.
Mendel’s work only said that some material was passed from parent to child that carried genetic information – it made no assumption as to what that material could be. That would have to wait for nearly a full century, until James Watson, Francis Crick and colleagues worked out the structure of DNA in the early 1950s. The structure of DNA (and with it, how genetic information was passed on a molecular level) caused another revolution in our broader-scale understanding of the relationships between organisms. Building on the original work on the structure of DNA, recent biologists have been able to decode significant portions of the genetic code, presenting a clearer picture of evolutionary relationships among species. As we have come to understand the genetic basis of relationships between species, the emphasis of taxonomy has shifted from groups of physically apparent relatives (how Linnaeus constructed his classifications nearly 300 years ago) to groups of genetically closest relatives, called clades. Clades can be nested within each other, just as Linnaeus’ genera nested within families, families within orders, and so on. Clades of roughly the same extent as Linnaean groups are still called by the Linnaean name, so a Linnaean family that more or less encompassed a molecularly defined clade would still be called a family.

Winter Reading: Remarkable Creatures (Tracy Chevalier)

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Friday, February 18th, 2011

Now that the weather here is a bit more wintry (California wintry – wind and rain), I’ve been catching up on my long reading list. Last week I finished Remarkable Creatures (by Tracy Chevalier, $15, Penguin Group, 2010), an account of the friendship between two women who played significant but largely forgotten roles in the development of evolutionary theory. Though it lacks some of the social or personal nuance of much literature written about nineteenth-century Britain, this book is like Jane Austen for science and nature nerds.

Mary Anning was born in 1799 in the British town of Lyme Regis, a tiny hamlet on the coast of Dorset, and struck by lightning as an infant. As she grows up, it becomes clear that she is odd, made different by luck or lightning. Her unique ability to find fossils on the wild coastal beaches of Lyme keeps her destitute family from debtor’s prison, but she doesn’t realize the significance of the creatures she unearths until Elizabeth Philpot, a spinster with a passion for natural history, moves from London to Lyme Regis with her sisters. Through a series of encounters on the nearby Jurassic cliffs, Elizabeth becomes a mentor for Mary, introducing her to the work of Georges Cuvier and other contemporary natural historians. As their discoveries of ammonites, “ichies” (ichthyosaurs), and “plesies” (plesiosaurs) challenge the dogma of the Church, Mary and Elizabeth confront challenges both social and ideological and begin to make their mark on science, a stratum of society where neither women nor radical ideas are welcome.

Tracy Chevalier, who has vivified numerous other characters marginalized by history (e.g., Girl with a Pearl Earring), animates Anning, Philpot, and their social and natural milieux in a way that’s both engaging and edifying. While she certainly takes some creative liberties (and that’s her right and duty as a historical novelist), Remarkable Creatures is valuable recognition for two figures that have been largely obscured from the annals of scientific history.