There are plenty of popular palaeontology books that tell you everything we know about dinosaurs and several excellent examples have been reviewed here in the past. For the 500th review on this blog, I take the road less travelled. In The Future of Dinosaurs, English palaeontologist David Hone flips the script by asking what we do not know about dinosaurs. I have been meaning to review this book since it was first published in 2022. With the recent publication of his latest popular book on dinosaur behaviour, I decided to make time and read up on Hone’s work. First up, an exploration of our ignorance that is as much a celebration of all we have learned and how we have learned it.

The Future of Dinosaurs: What We Don’t Know, What We Can, and What We’ll Never Know, written by David Hone, published in Europe by Hodder & Stoughton^[1] in March 2022 (hardback, 252 pages)

This book seemingly flows from Hone’s efforts at science communication. In speaking to the general public, he has noticed the sometimes large disconnect between what palaeontologists know and what people think they know. Thus was planted the seed for a Rumsfeldian^[2] book about what is unknown, what is knowable, and what is unknowable in palaeontology. Fifteen chapters run the gamut of topics and chart the cutting edge of our ignorance and our understanding. Given the large number of popular dinosaur books, it is a refreshing take on the genre that is chock-full of interesting insights.

Take some examples of unknowns. What is the total number of species that ever evolved and how many can we still expect to find? That is unknown, though if you plot discovery rate against time you expect a curve that will eventually level off. Where we are on that curve? Who knows! The rate at which we name new species each year shows no signs of slowing down yet. Did sauropods have multi-chambered stomachs like today’s large herbivores? That is unknown, as soft-tissue preservation of this kind is exceedingly rare. All we have for now is a spectacular fossil of the small theropod Scipionyx where the guts were infilled with sediment, resulting in a remarkable cast of an otherwise unremarkable digestive tract.

Many things are knowable though, with questions often being an absence-of-evidence problem that can be resolved with more fossils and more research. Anatomical features such as the frills of ceratopsians are usually bent and broken during the long journey through time, but detailed scans and computer analyses allow us to undo such distortion virtually. It is still early days for this technique, but we have the fossils, so with time, effort, and experience, our knowledge is likely to improve. Were dinosaurs sexually dimorphic? That is, did males and females differ noticeably in size or appearance? Since most species are only known from one or a few specimens, we just do not have the required sample sizes yet, and where we do, we have not found such differences. However, given the large diversity of dinosaurs, there are bound to have been sexually dimorphic species, so with more fossils this question is ultimately answerable.

“[The] disconnect between what palaeontologists know and what people think they know […] planted the seed for a Rumsfeldian book about what is unknown, what is knowable, and what is unknowable in palaeontology.”

The unknowables are some of the most fascinating and frustrating questions discussed here. Are tooth marks on bones the result of predation or scavenging? Predators readily did both and the two are near-indistinguishable. “This is annoyingly trite as a revelation about their behaviour” (p. 183), but the data simply do not allow for firmer conclusions. The small asterisk is that some questions are not strictly unknowable, but would require such unlikely fossils that, for all intents and purposes, we can consider them unknowable. Others require completely new types of evidence that we had not previously conceived of. For example, the colour of dinosaurs was considered unanswerable until recently. The discovery of ancient biomolecules such as melanosomes changed that somewhat, as their shape corresponds to colour. “It is as if you went to a paint store and each colour was in a uniquely shaped pot” (p. 151), is how Hone memorably describes it. But he quickly tones down the enthusiasm: only certain colours are melanosome-based and sampling them is destructive, so palaeontologists try and limit it to spot sampling. Our knowledge is thus far less certain and complete than artist reconstructions suggest.

Despite my enthusiasm for Hone’s three-pronged approach, I do have some criticism. First is that the order of chapters feels somewhat haphazard, with the book jumping around between different topics and levels of organisation. Some closely related topics are scattered over the book with e.g. chapters on habitat and ecology separated by seven chapters. Others feel out of order. Why start with a chapter on extinction when, by your own admission, anatomy (now chapter 7) is the fundamental topic on which everything else in palaeontology is built? I would have reorganised the chapters for a more logical structure. Second is that the writing is sometimes not the most graceful. Take the book’s subtitle: to me, it cries out for a third “know”; leaving it out of the middle clause is just jarring. There are examples of long, run-on sentences expressing several ideas that should have been separated. Other sentences are awkwardly constructed, or switch from passive to active voice midway, with my brain expecting one thing but Hone writing something else. Now, I do not want to over-egg this: big picture, they are all minor blemishes on what is otherwise an entertaining and engaging book.

Next to Hone’s stated running theme of the disconnect between the public’s perception of our knowledge and our actual knowledge, some other fascinating themes emerge. One example is the nature of the fossil record and how it can mislead through both a glut and a dearth of fossils. Bonebeds are often significant sources of information due to the large number of bones preserved, but if they resulted from abnormal circumstances such as catastrophic floods, droughts, or volcanic eruptions, how representative are such assemblages? They risk giving “a very misleading picture of what lived locally and in what kind of numbers” (p. 194). At the other extreme are the low odds of preserving the statistical outliers such as the largest or oldest individuals. Similarly, newly emerging species often occur in low numbers and limited geographic areas, so the age of a species should be considered a minimum estimate. The odds are very much stacked against the earliest, if not the first, representatives of a species entering the fossil record, surviving to the present day, and then being discovered rather than lost to erosion.

“[…] some other fascinating themes emerge. One example is the nature of the fossil record and how it can mislead through both a glut and a dearth of fossils.”

Another fascinating theme that emerges is the impact of the huge size range of dinosaurs on their biology and ecology. Dinosaurs never evolved to be rodent-size small; as Elsa Panciroli discussed, mammals occupied that ecological niche. Furthermore, independent of their adult size, dinosaurs hatched from eggs as small juveniles and had to go through sometimes gargantuan growth spurts. Hone points out that juveniles of sauropods and large theropods simply could not have been eating the same food and playing the same ecological role as the adults. Fellow palaeontologist Michael J. Benton recently highlighted research that suggested carnivores passed through a range of feeding modes as they grew up, sequentially occupying different ecological niches and effectively acting as different species on their way to adulthood. Similar shifts might have also happened to metabolism, Hone suggests. Adult sauropods were so large that their surface-to-volume ratio meant they did not need to actively generate body heat to keep warm (a condition known as gigantothermy). But the same could not have been true of small juveniles; seeing their growth rates they likely were endotherms, actively generating body heat. If true, this would be remarkable, as we know of no living species that so radically change their physiology growing up.

Despite some minor flaws, The Future of Dinosaurs is both a fascinating exploration of our ignorance and a celebration of how much we have learned from sometimes very challenging source material. By tracing the contours of where our knowledge ends, Hone gives you an insider’s look into the discipline—from its storied history to today’s cutting-edge technologies. It seems almost prophetic that he writes that behaviour is the subject where we see the greatest disconnect between what palaeontologists know and what the public thinks they know: he tackles this topic in his recently published Uncovering Dinosaur Behavior to which I will turn next.

1. ↑ Note that in the USA, this book was published by Princeton University Press with the title How Fast Did T. rex Run?: Unsolved Questions from the Frontiers of Dinosaur Science.

2. ↑ I am, of course, referring to United States Secretary of Defense Donald Rumsfeld’s briefing where he popularised the notion of known knowns, known unknowns, and unknown unknowns.

Disclosure: The publisher provided a review copy of this book. The opinion expressed here is my own, however.

The Future of Dinosaurs

Other recommended books mentioned in this review:

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Whether you embrace the concept or think that we are on the cusp of it, the term “Sixth Extinction” serves as a useful shorthand to bring into focus the scale and tempo of recent and ongoing biodiversity loss. Famous victims such as the dodo, the thylacine, the passenger pigeon, or the great auk will immediately jump to mind, but they are just the tip of the iceberg of extinction. Few people will think of the St. Helena olive, the Bramble Cay melomys, or the Christmas Island forest skink. And therein lies the problem: behind the faceless statistics of loss lie numerous stories of unique evolutionary lineages that have been snuffed out. In this emotional gut punch of a book, author and journalist Tom Lathan takes the unconventional approach of examining ten species that have gone extinct since 2000, nine of which you will likely never have heard of. Lathan momentarily resurrects them to examine what led to their loss and speaks to the people who tried to save them.

Lost Wonders: 10 Tales of Extinction from the 21st Century, written by Tom Lathan, published by Picador in November 2024 (hardback, 464 pages)

Of all the species featured in Lost Wonders, the one species and individual you are likely to have heard of is Lonesome George, the last Pinta Island tortoise, who received much media attention both in life and in death. Some others have been the subject of dedicated books—the po’ouli (a Hawai’ian bird), the Christmas Island pipistrelle (a bat), and Partula labrusca (a Polynesian tree snail)—though you would be forgiven for not knowing of them. The remainder are barely known outside of the small circle of conservationists who studied them. One comparable book is Errol Fuller’s Lost Animals which looked at the photographic record of 28 recently extinct animals, though it went into nowhere near as much detail as Lost Worlds does. Of the unfortunately long list Lathan obtained from the International Union for Conservation of Nature (IUCN), he has chosen a nicely balanced mix of species, including multiple snails, birds, mammals, reptiles, and a single fish and tree. Each chapter opens with a tastefully executed pencil-and-ink drawing by Lathan’s partner Claire Kohda. The geographic spread similarly includes organisms from around the globe (though no maps have been included, which would have been helpful). What unites these species is that they all lived, literally or functionally, on islands: environments that are particularly prone to both evolutionary experimentation and extinction. Indeed, the other factor that unites them all is that they are no longer with us.

Each of the ten chapters mixes several elements such that, despite most chapters being quite long at 30–40 pages, all are very engaging. Lathan introduces what we know about their biology and how the frequent paucity of information frustrated subsequent attempts at captive breeding (as was the case for the Malaysian microsnail Plectostoma sciaphilum). By telling the stories of their discovery and formal description, Lathan answers in the affirmative his question of whether naming a species “is itself a life-giving act” (p. 28). The tangible benefit is that we can formulate conservation plans, explaining how naming “can be the difference between life and death” (p. 29). Lost Wonders thus offers its own little rallying cry for taxonomy. Of course, species exist before we describe them, and his overview of their evolution is a potent reminder of this. It also highlights how, given enough time, organisms can reach remote islands and establish themselves there, despite the odds not being in their favour. The evolution of the Galápagos tortoises is outright fascinating. A genetic study concluded they share a common ancestor with the much smaller Chaco tortoise. You would think that the most parsimonious explanation is that the Galápagos tortoises evolved into giants after making it to the archipelago. However, the real kicker, which Lathan casually drops in a footnote, is the discovery of giant tortoise fossils on mainland South America, suggesting that their gigantism preceded their dispersal.

“there is both a long queue of species awaiting assessment by the underfunded IUCN and an even larger pool of “dark extinctions”: species that vanish before we even know of their existence.”

All of the above is relevant background information, but we are here for the stories of what went wrong. In the introduction, Lathan emphasizes just how incomplete our knowledge is: there is both a long queue of species awaiting assessment by the underfunded IUCN and an even larger pool of “dark extinctions” (p. 4): species that vanish before we even know of their existence. These stories are “a snapshot of extinction […] each a stand-in for other[s] that we will probably never know occurred” (p. 4). There is that importance of taxonomy again.

If extinctions of Pleistocene megafauna can reasonably be attributed to a mixture of human hunting and climate change, the fingerprint of more recent Holocene extinctions is clearly human. Lathan points out that our species is “one of the most potent agents of ecological destruction, regardless of time, place, or culture” (p. 135). For instance, the arrival of native Polynesians to Hawai’i already triggered a wave of extinction, such that the arrival of Europeans “was more like a passing of the baton in an ecocidal relay race” (p. 136). That said, in his next breath he immediately recognizes that European colonialism cranked up extinction to eleven—it is a good example of Lathan’s balanced reporting. What follows is the usual litany of rapacious resource extraction that destroys natural habitats and the accidental or purposeful introduction of invasive species. The two often work in tandem: the Christmas Island pipistrelle succumbed both to relentless phosphate mining and a suite of introduced predators, prominently the wolfsnake; the St. Helena olive had to contend with deforestation as the island was cleared for crops while numerous species escaped its botanic garden given its role as a staging post in the international trade in ornamental plants.

The strongest suit of Lost Wonders is the nearly 50 interviews with scientists, conservationists, hobbyists, and others whose first-hand experiences and frustrations imbue this work with much pathos. There are stories of species slipping through our hands as their habitats vanished (e.g. the Bramble Cay melomys, a rat, or the Alagoas foliage-gleaner, a bird); of ignored warnings, bureaucratic red tape, and apprehensive committees delaying meaningful action (e.g. the Polynesian tree snail P. labrusca or the Christmas Island pipistrelle); and of captive breeding efforts failing (e.g. the Mexican Catarina pupfish). By asking how the people involved experienced witnessing extinction, each chapter delivers an emotional gut punch that, I will not lie, sometimes brought me to tears. Some people still struggle talking about it, even a decade or more later, breaking down during their interviews. Others describe feelings of grief, depression, and loneliness, unable to truly share with others what they experienced. Lathan himself in his epilogue expresses his astonishment at “their capacity to articulate the profundity of what they had witnessed” (p. 351) and wonders out loud: “When a million years of evolution is extinguished right in front of you, what words suffice to describe this moment?” (p. 351).

“By asking how the people involved experienced witnessing extinction, each chapter delivers an emotional gut punch that, I will not lie, sometimes brought me to tears.”

Taking a step back to compose myself I do, however, have two points of criticism; or, if not criticism, two points I feel have been omitted. Had they been addressed, this book would have been a straight ten out of ten; that is, if I was in the habit of assigning a score to my reviews.

First, there is the proximate question of whether trying to save a species at all costs is always the best use of the limited time, money, and other resources available for conservation. Not everybody agrees it is, and e.g. Inheritors of the Earth provocatively argued that island species are evolutionary dead ends, vulnerable to invasion (I interviewed the author more in-depth back in 2017). Even if you disagree with that line of thinking, several stories here bring this question to the fore, such as the captive breeding attempts that failed because there was only one or a few same-sex individuals left. Terrestrial ecologist John Woinarski, who cared for the last Christmas Island forest skink, spoke of giving it palliative care. Park manager Rich Switzer sounds borderline traumatized caring for the last, ailing po’ouli, saying he would have euthanized it already had it been his own pet bird. Are resources better spent on populations that still stand a decent chance? What about the species that are now effectively marooned in captivity because their habitat has been destroyed? A counterargument could be made that, yes, these attempts are worthwhile because we learn how to improve our protocols, techniques, and technologies for the inevitable next extinction. This is driven home when reading the list of techniques that were brought to bear on the last St. Helena olive, including hand-pollination, propagation of cuttings, and tissue micropropagation. My point is that Lathan does not broach these questions here. I would have loved for him to wrestle with these and put them to his interviewees.

Second, there is the ultimate question of what it would take to turn the tide of extinction, of what such a world would look like. I judge him less harshly on this because, from my reading on biodiversity and environmental problems, very few authors seem willing to mention the root causes that got us here (overconsumption, overpopulation, capitalism, colonialism—all that good stuff), let alone provide a sensible stab at tackling these. Do I sound like a broken record? Undoubtedly. Are they huge, thorny, and uncomfortable topics? Yes, and that is exactly why such conversations are long overdue. His interviewees gave him several openings that he did not pursue. Take biologist Thor-Seng Liew who talks about the handbook he developed for the cement industry in Malaysia so they may avoid quarrying the most species-rich limestone outcrops. A pragmatic initiative for sure, but is this not just a stay of execution? Since when do companies turn around and say: “We have quarried enough now and thanks for your advice”? Similarly, when Professor Ana Laura Lara Rivera complains about the farmers who drain the lagoons where rare pupfish live, but also sympathizes with their need to put food on the table, that is a springboard for a conversation about the tension between poverty and conservation. This is another set of questions where both his views and those of his interviewees could have further enriched the book.

The above suggestions would have been cherries on the cake but, as it stands, the proverbial cake is both edible and rich. Lost Worlds is an incredibly moving book that tugs at the heartstrings and draws on an impressive number of interviews. These eyewitness stories are a powerful reminder that behind each reported extinction lies a tremendous amount of work, and the loss of a unique way of life on this planet.

Disclosure: The publisher provided a review copy of this book. The opinion expressed here is my own, however.

Lost Wonders

Other recommended books mentioned in this review:

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Can the environmental and wildlife conservation movements learn from the (distant) past? This turns out to be a fraught question, with many practitioners preferring to preach pragmatism over nostalgia. Journalist and writer Sophie Yeo agrees that there is no turning back time, but this is no reason to ignore history. In Nature’s Ghosts, she mixes several parts reportage with one part nature writing to both criticize different conservation approaches and showcase some really interesting research. Though centred on the UK, she also discusses projects and problems in Europe and the USA, and the book was deservedly shortlisted for the 2024 Wainwright Prize for Writing on Global Conservation^[1].

Nature’s Ghosts: The World We Lost and How to Bring It Back, written by Sophie Yeo, published by HarperNorth (a HarperCollins imprint) in May 2024 (hardback, 327 pages)

What unites Yeo’s criticism of current conservation practices is how they ignore the past. For example, the British approach of designating areas with particularly valuable habitat, flora, or fauna as sites of special scientific interest (SSSIs) aims to keep these frozen in time. Yeo counters that by resisting the natural dynamism of ecosystems, “these sites, like Peter Pan, may never grow up” (p. 64). EU subsidies as part of the Common Agricultural Policy aimed at enhancing biodiversity in practice have had the opposite effect, as discussed for Romania. They are disbursed on the condition that farmers abandon traditional practices, even though these are far more interwoven with the natural world than the centralised rules imposed by the EU. Fortress conservation, the designation of areas as nature preserves or national parks, has been frequently combined with the forced eviction of Indigenous people from their lands. A recent searing critique exposed its colonialist roots as it pertains especially (though not exclusively) to Africa. It replaces the actual past with a mythical past of virgin wilderness that never existed in the first place. Yeo adds that: “conservation should have never become about displacing people from the land. The wilderness is not a blank slate: until recently, it was our home, our larder, our library” (p. 150).

Yeo is more positive about the track record of rewilding though here the problem of baselines crops up. “The natural world has never experienced one perfect moment of Eden: it has always been a series of novel ecosystems” (p. 66). With nature forever in flux, where do you plant your yardstick? She is critical of the feted Knepp Estate: “whether the land amounts to a genuine approximation of wilderness or a glorified farm remains hotly debated” (pp. 54-55). She feels conflicted by the Carrifran Wildwood project in Scotland that has accelerated its rewilding goals by gathering tree seeds from all over Scotland and Northern England. She likes what she sees but also considers it an example of intense micromanagement: “In many ways, it couldn’t have been a grander monument to human intervention” (p. 72). The Finnish Snowchange Project has the more nuanced aim of integrating humans into the landscape, in this case by supporting traditional small-scale ice fishing. Project leader Tero Mustonen argues that rewilding becomes powerful once you integrate cultural knowledge into it.

Of course, Yeo acknowledges more than once, we cannot turn back time. There are good practical and philosophical reasons for conservationists to be pragmatic and future-oriented. However, “that does not mean that we should treat the past as irrelevant” (p. 5). So, how is the past relevant to conservation today? The second half of Nature’s Ghosts features fascinating research of which I can only discuss a few examples.

“Yeo is more positive about […] rewilding though here the problem of baselines crops up. “The natural world has never experienced one perfect moment of Eden: it has always been a series of novel ecosystems” (p. 66).”

First, the extinction of Pleistocene megafauna. To me, it has become uncontroversial middle ground in this ongoing debate to implicate both climate change and humans. Novel is the notion that humans effectively replaced megafauna as a keystone species, with a new suite of plants and animals becoming dependent on our role in ecosystems (think of the archaeological evidence for forest gardens). The displacement of traditional societies through colonialism has disrupted such relationships, such that rewilding should “reinstate not only the lost ecological functions of megafauna but also people” (p. 136). This is what Mustonen’s Snowchange Project aims for.

Second, shifting baseline syndrome. Originally developed by Daniel Pauly in the context of fisheries, this has become another staple when talking about lessons from the past. Novel is the idea that, with each passing generation, we have not just collectively forgotten how abundant the natural world was, but also how animals used to behave. Ecologists mistakenly assume that an animal’s current habitat is both its preferred habitat and the limits of its tolerance, with conservation boiling down to “wherever an animal lives now, we need more of that habitat”. One example given here is the wild bison in Poland’s Bialowieza Forest. These are not a relic species but refugees: “survivors that have been backed into a corner, making do with what little remains” (p. 170). Indeed, one South African ecologist studying them was puzzled as to “why these European people have cows in the forest” (p. 174). The fact that many plants and animals are more tolerant than we give them credit for is a feeble ray of hope in the ongoing maelstrom of extinction.

Third is how time matters both more and less than we think. On the one hand is the long shadow cast by past human activities. The advent of remote sensing technology in archaeology has e.g. revealed how presumed old-growth forest in France hides the ruins of Roman farms. Amazingly, the soil and plant communities still differ as a result of fertilizer application two millennia ago. One shudders to think what that means for the future legacy of today’s environmental footprint. Will future fossils indeed be technofossils? On the other hand is temporal dispersal. Yeo discusses the many ponds, pools, and puddles that once dotted Britain’s landscape but were backfilled over time. When such ghost ponds are excavated again, their old bottoms contain spores, eggs, and seeds that remain viable centuries later. Many plants and invertebrates can rely on dormancy as a survival strategy to bridge time. Step aside deextinction! This “is scratching back the surface to reveal the real thing” (p. 230).

“we have not just collectively forgotten how abundant the natural world was, but also how animals used to behave.”

Nature’s Ghosts flows over with thought-provoking ideas and Yeo is quick to step in and prevent them from being misinterpreted or hijacked. No, the traditional agricultural practices in Romania or Wales are not blueprints for feeding the world. No, the Finnish Snowchange Project is not “arguing for the revival of a Stone Age lifestyle in North Karelia” (p. 138). No, discussing traditional foraging is “not advocating for the return of a pre-agrarian society” (p. 144), etc. She is well aware that conservation debates have been toxic at times. But it does beg the question, if these are not solutions, then what is?

I have two minor frustrations with Nature’s Ghosts. First, Yeo is reluctant to name the elephant in the room, even if she comes close. When discussing traditional farming methods, she remarks how “fossil fuels enabled us to exceed earth’s natural carrying capacity” (p. 104). When discussing biodiversity loss, she concludes that “so long as humans remain dominant, nature can never return to its primeval abundance” (p. 181) and that “wildlife cannot be restored to prehistoric levels without massive reductions in the human population: the earth will only support so much life” (p. 169). The dreaded O-word is studiously avoided, possibly for fear of getting tarred and feathered. Second, her solution. By no longer seeing landscapes as something sacred (for which she thanks Christianity) we paved the way for today’s relentless extraction. “The trouble is not that the old stories have faded. It is that no new magic has emerged to take their place” (p. 255). Rather than revitalising old beliefs or rituals whose meaning has been lost, she thinks we each need to rewild ourselves and find new symbolic connections with the natural world. If you are of the pragmatic bend that might sound a bit wishy-washy. To my surprise, I actually agree with her. Somewhat. I agree that we are not moved by facts but by stories, so, yes, we do need a new narrative. However, I am rather thinking along the lines of Eileen Crist’s call for an ecological civilization that does not see nature as a mere larder, or Carl Safina’s call to abandon the Platonic dualism that has allowed us to think of ourselves as apart from nature. Abandoning the currently dominant fairytale of endless economic growth will mean a wholesale revision of our socio-economic system, likely involving a period of degrowth towards a steady-state economy. Tall order? Absolutely. Do I expect Yeo to here write the book-length treatment that such big topics need? No, but acknowledging them would have been nice.

Those niggles notwithstanding, Nature’s Ghosts is intriguing, discussing some of the fascinating insights gleaned from studying the past. Yeo has read up on the literature and gone to great lengths to speak to researchers and conservationists first-hand. Given how she combines journalistic flair and evocative writing to explain the relevance of this research, it is easy to see why the Wainwright Prize jury was impressed.

1. ↑ For all my fellow non-UK readers, the Wainwright Prize is an annual UK literary prize that is named in honour of British author, illustrator, and hillwalker Alfred Wainwright (1907–1991). It was first awarded in 2014, split into two categories (UK nature writing and writing on global conservation) in 2020, and saw a third category for children’s books added in 2022. Various sponsors have affixed their names to the prize over the years, resulting most recently in the James Cropper Wainwright Prize.

Disclosure: The publisher provided a review copy of this book. The opinion expressed here is my own, however.

Nature’s Ghosts

Other recommended books mentioned in this review:

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Stephen Jay Gould (1941–2002) was a well-known evolutionary biologist, palaeontologist, and science populariser. Amongst his many achievements stand the 300 popular essays that appeared from 1974 to 2001 in the magazine Natural History, published by the American Museum of Natural History. Many of these were collected in bestselling volumes that have been reprinted repeatedly. To celebrate this legacy of essays, his friends and close colleagues Bruce S. Lieberman and Niles Eldredge, themselves evolutionary biologists and palaeontologists of considerable renown, here present thirteen of their own essays that do exactly what the subtitle promises. They entertain as often as they intrigue in a collection that draws serious and, looking at the chapter titles, sometimes not-so-serious connections between macroevolution and palaeontology on the one hand, and popular culture, philosophy, and the history of science on the other. To my shame, I have to admit that I have never read Gould’s essays or his many books (while having several on my shelves). Macroevolutionaries convinced me that this gap in my knowledge needs closing.

Macroevolutionaries: Reflections on Natural History, Paleontology, and Stephen Jay Gould, written by Bruce S. Lieberman and Niles Eldredge, published by Columbia University Press in September 2024 (hardback, 216 pages)

Despite being a towering intellectual figure, or perhaps because of it, there is no book-length biography of Gould as far as I know. However, over the years, books have reflected on his intellectual life, his political views, his humanism, and (still forthcoming) his criticism of biases in science. Macroevolutionaries is similarly not a biography but adds to this growing body of work engaging with Gould’s views. Furthermore, the book is also not an uncritical celebration of Gould. Though they respected him greatly and will defend him where they feel he is no longer given the credit he is due, “that doesn’t mean that we agreed with him on each and every last thing” (p. 40). They rather candidly write that he sometimes went solo and modified ideas cooked up with others, in the process “getting more than his share of the credit and attention” (p. 7).

This characterisation of Macroevolutionaries as neither biography nor hagiography also extends to the essays themselves. Not all of them invoke Gould, instead standing on their own two feet. The essay Expanding Evolution: Organisms and Species, the Soma, and the Technosphere explores how the logic underlying evolution by natural selection can be expanded to e.g. cancer cells (somatic evolution) and material culture (technospheric evolution). Eldredge here draws on his passion for musical instruments by exploring “convergent evolution” between trumpets and cornets, a by now similar-looking instrument. In Darwin in the Galápagos: Running the Beagle Tape Backward, Eldredge provides an answer to an intriguing thought experiment a historian once put to him: would Darwin have come up with the same ideas had his trip on the Beagle happened in reverse, with the Galápagos being the first port of call? Eldredge thinks not and highlights key observations Darwin made during his trip that changed his initial views on transmutation: “there were things Darwin needed to see first to comprehend the significance of what he would see in the Galápagos in August 1835” (p. 147).

“This characterisation of Macroevolutionaries as neither biography nor hagiography also extends to the essays themselves. Not all of them invoke Gould, instead standing on their own two feet.

Other essays do engage explicitly with Gould’s ideas. A recurrent touchstone is the theory of punctuated equilibria that Gould and Eldredge developed in a 1972 paper (drawing on earlier work by Eldredge): the idea that speciation in the fossil record alternates between long periods of stasis and periods of abrupt change. This contrasted with Darwin’s original view of slow and steady change. How do such long periods of evolutionary stasis come about? In their essay, the authors point out how Gould initially argued for the importance of developmental constraints, though he later downplayed their role. Punctuated equilibria is such an influential idea that Belknap Press posthumously reprinted the long chapter on it in Gould’s magnum opus as a separate book in 2007, notably only with Gould’s name on it; Eldredge wrote about the conception of the idea separately in his 1985 Time Frames.

Gould is similarly famous for his thought experiment of “replaying the tape of life” and asking whether the outcome would be the same (an idea that has come up here in previous reviews). He answered “no” and in the process made a case for the importance of contingencies in the history of life. What I did not realise, and the authors here clarify, is how Gould’s views on the relative importance of contingency and natural laws swung back and forth over the course of his career. One more example of engagement with Gould’s ideas will have to do. When Is a Raptor a Parrot? The Curious Case of the American Kestrel starts as an essay about bird phylogeny before pivoting to Gould & Lewontin’s famous spandrels paper and Gould & Vrba’s exaptation paper. In these, Gould and colleagues argued against adaptationist explanations for every single trait an organism shows, while proposing a new term (exaptation instead of adaptation) for traits that initially evolved in one context or were neutral, and were later coopted in another context.

“A second strong component in this collection is the history of science; an area that is relevant as it also interested Gould.

A second strong component in this collection is the history of science; an area that is relevant as it also interested Gould. In several essays, the intellectual movers and shakers that influenced Darwin come up, notably Giambattista Brocchi, Georges Cuvier, and Jean-Baptiste Lamarck. Brocchi had some ideas about sudden evolutionary transitions that lined up nicely with punctuated equilibria. Cuvier championed catastrophism and species extinction^[1]. And Lamarck was at loggerheads with Cuvier by arguing for slow and gradual change of species. The authors contend that he, rather than Lyell, should be credited for influencing Darwin’s gradualist take on evolution. Elsewhere, Eldredge has at length sought a rehabilitation of Lamarck and here reiterates this. “We grew up hearing the same sort of smack about Lamarck that is still au courant [but] Lamarck was Darwin’s hero, and not the Darwinian anti-hero we were taught to believe” (pp. 57–58). Fascinating, too, was the essay Of Cultural Nationalism, Hamlet, and the Cloaca Universalis: Why Citation is the Best Policy that wonders out loud why Charles Lyell is venerated as the father of geology when in his time he had a reputation for blatant plagiarism.

These are but some of the major themes in these intriguing essays. Take it from me that there are others—it’s a worthwhile collection. Overall, the essays are very readable and often rather informal in tone: the authors are on first-name terms with many people and Gould is praised as “bloody brilliant” (p. 13), “we respect the dude a lot” (p. 40), and (together with Lewontin) given “mad props and kudos” (p. 176) for the audacious title of their spandrels paper. Occasionally, the authors swerve into rather verbose writing that had me reach for the dictionary. Similarly, the significance of certain references to US-specific lore and history (e.g. stuntman Super Dave or journalist Walter Winchell), while briefly explained, might be lost on an international audience.

Readers who miss Gould’s essays or are enthralled by his ideas are warmly recommended to pick up a copy of Macroevolutionaries. Lieberman & Eldredge celebrate his legacy while writing fine essays that can stand on their own two feet. For me, this collection additionally highlighted a gap in my knowledge (and on my shelves) that I intend to fill while reminding me that I have unread books by both Gould and Eldredge that I should carve out time for.

1. ↑ Mind you, mass extinctions were not fashionable yet: one essay highlights the role of Gould’s mentor Norman Newell in making this idea more palatable.

Disclosure: The publisher provided a review copy of this book. The opinion expressed here is my own, however.

Macroevolutionaries

Other recommended books mentioned in this review:

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Gould’s collected Essays On Natural History:

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What a killer title. Rarely have I seen three snappy words so effectively capture the essence of a concept in biology. What concept is that? Zoogeochemistry. Many scientists have convincingly made the case that it is the small things that run the world. Though it is undeniably true that e.g. microbes and insects have shaped our planet, and continue to do so, it would be a mistake to think that larger animals are just along for the ride. I was stoked the moment the announcement for this book dropped and conservation biologist and marine ecologist Joe Roman did not disappoint. Eat, Poop, Die is fun and fascinating, while always keeping one eye firmly on the facts and complexities of ecology. Is it too soon to start earmarking titles for this year’s top 5? I think not.

Eat, Poop, Die: How Animals Make Our World, written by Joe Roman, published in Europe by Profile Books in April 2024 (hardback, 277 pages)

Given that this concept “is the beating heart of this book” (p. 214), what is zoogeochemistry? Physical forces shape the living world and living forces can shape the physical world right back—you may well have heard of biogeochemistry. Traditionally, research has focused on bottom-up interactions, revealing how e.g. microbes influence the climate and fungi and invertebrates help form soils. The phrase “ecosystem services” is often bandied about in this context, referring to how organisms keep our planet liveable by fertilizing soils, filtering water, or absorbing carbon dioxide from the atmosphere. Increasingly, however, it is becoming clear that larger animals similarly have the power to shape ecosystems, influencing the flow of chemical elements at large scales. Hence zoogeochemistry. How do animals do this? The title of the book says it all: rotting carcasses, deposited faeces, “and other unglamorous forces” (p. 52) can collectively move nutrients around ecosystems and influence the climate.

By visiting and interviewing fellow ecologists, and mixing in some of his own research, Roman introduces you to a variety of very interesting studies that are now uncovering these processes. He starts simply enough with the volcanic island Surtsey which appeared above the waves off the coast of Iceland in 1963. It has offered a unique opportunity for ecologists to study how a newly formed island gets colonized by plants and animals, with birds and seals enriching the initially scrappy plant communities through faeces, urine, carcasses, and eggshells. In the oceans, zooplankton move carbon and other elements from the surface to the deep , forming one component of the so-called biological pump. But whales, because of their sheer size, have the potential to counteract this. By bringing nutrients back from the deep, where they feed, to the surface, where they release nutrient-rich faecal plumes, they form what Roman calls a whale pump. (This happens to be his speciality, so expect a fair bit of lyrical waxing about whale poop.) In the Maasai Mara, wildebeest that die in river crossings provide an annual nutrient pulse, on top of the steady and intense input of hippos pooping in their pools. Coral-crunching parrotfish poop out the sand that covers the beaches of many tropical holiday destinations, and when periodical cicadas die in their trillions, they provide enormous but irregular nutrient pulses that have barely been studied. Roman appropriately concludes the book with James Estes’s work on sea otters in Alaska that has become the poster child for the concept of trophic cascades. In the process, he provides the interesting backstory of how the United States Atomic Energy Commission left a legacy of both atomic test sites and translocated otters that would repopulate the Pacific coast of the USA, a backstory Estes did not really detail in Serendipity. These and other examples form a delectable list of study systems, each more interesting and unbelievable than the last. No wonder Roman is often asked: “How did we overlook this for so long?”

“rotting carcasses, deposited faeces, “and other unglamorous forces” (p. 52) can collectively move nutrients around ecosystems and influence the climate.”

What I particularly appreciated is that for some of the better-studied systems, Roman gets the nuances right. The story of wolf reintroduction in Yellowstone National Park suppressing elk overgrazing and leading to a restored ecosystem has almost become a meme in ecology. The details, however, are a bit more complicated than the idea that you can just sprinkle wolves over a broken ecosystem like some sort of magical fairy dust. Similarly, the notion that bears indirectly fertilize trees by littering the forest with salmon carcasses, which Suzanne Simard approvingly mentioned in Finding the Mother Tree, has a few detractors. Roman spends time in the field with Daniel Schindler who thinks that poorly executed science and overinterpreted stable-isotope data have created a good story. One of his students adds that, though the cycling of nutrients between different animals in this system is not in question, the idea that it also fertilizes trees with nitrogen is considered more shaky by some. When it comes to the Pleistocene extinction of megafauna, Roman again hits the nail on the head by acknowledging the debate about the relative contributions of human hunting and a changing climate.

Eat, Poop, Die avoids becoming an endless showcase of stories about animals moving nutrients around ecosystems in said fashion. Chapter 5 (Chicken Planet) could have easily concluded the book but instead sits smack in the middle, punctuating the flow nicely. These sombre crescendos partially answer Roman’s earlier question of why we have overlooked the role of large animals for so long: because many have been driven to near-extinction by humans. The consequences of this have been studied in quite some detail for whaling, which resulted in impoverished marine ecosystems. But similar landscape changes resulted from the North American fur trade that saw the slaughter of bison, beavers, and otters. Humans have since become the prime mover of nutrients, with the mining of guano islands one sordid chapter in that history. The Haber–Bosch process might nowadays provide plentiful nitrogen (at a large environmental cost), but phosphorus still has to be mined. Future shortage is one of those creeping problems that is often overlooked in environmental discourse.

“[The book presents] a delectable list of study systems, each more interesting and unbelievable than the last. […] I particularly appreciated that for some of the better-studied systems, Roman gets the nuances right.”

Twice, Roman ruminates on whether rewilding our planet could help restore the nutrient flows that we have disrupted. It certainly has a better track record than geoengineering proposals. He is honest enough to highlight the downsides of large animals again roaming the land (from beavers flooding properties to otters hunting large clams that are commercially fished) and acknowledges that certain scenes from the past would cause social upheaval today. Achieving E.O. Wilson’s vision of setting aside half the planet for nature is “a long shot, especially if people are still eating meat on the bone and the population is increasing from eight billion to nine billion or ten” (p. 236). The most outspoken he gets is when he concludes: “Look, there’s no way around it. […] we’re going to have to consume less, consume better, if we’re going to rewild the world. A managed retreat is the only reasonable approach” (p. 238). I agree, and I cannot but marvel at how he is quietly radical here, almost sneaking in under the radar the three proverbial f-bombs (“veganism”, “degrowth”, and “overpopulation”) that often spark heated discussions. Though I could slate Roman for not developing this much further, I recognize that these are complex topics that could easily fill several books, and it would take us too far away from the main theme of this book.

Eat, Poop, Die is an absolute stonker that captivates from start to finish. The topic of zoogeochemistry is an incredibly fascinating development in ecological research and Roman recounts it here with flair.

Disclosure: The publisher provided a review copy of this book. The opinion expressed here is my own, however.

Eat, Poop, Die

Other recommended books mentioned in this review:

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In this review, I am revisiting the spectacular diversity of marine reptiles that flourished in the planet’s oceans and waterways during the time of the dinosaurs. After having gone without popular titles on the subject for almost two decades since Richard Ellis’s Sea Dragons in 2005, suddenly we have three. Last year April-May I reviewed Ancient Sea Reptiles and The Princeton Field Guide to Mesozoic Sea Reptiles, and mentioned that this book was in the works. Ocean Life in the Time of Dinosaurs was originally published in French in 2021 as La Mer au Temps des Dinosaures by Belin/Humensis and has been translated into English by Mark Epstein. Technically speaking that makes it the first of this recent crop, though the English translation was only published in November 2023, after the aforementioned two works. It brings together four French palaeontologists and one natural history illustrator for a graphics-heavy introduction. So, what is in this book, and how does it compare?

Ocean Life in the Time of Dinosaurs, written by Nathalie Bardet, Alexandra Houssaye, Stéphane Jouve, and Peggy Vincent, illustrated by Alain Bénéteau, published by Princeton University Press in November 2023 (hardback, 208 pages)

Ocean Life in the Time of Dinosaurs breaks down into two halves. The obligatory first short chapter introduces the state of the world during the Mesozoic Era 252–66 million years (Ma) ago, specifically the position of the continents (the palaeogeography) and the various extinction crises by which we divide this time span. After this, the first half is a very long chapter 2 that discusses all the major and minor groups: the “big three” (ichthyosaurs, plesiosaurs, and mosasaurs); the groups with survivors today such as the crocodylomorphs and sea turtles; and lesser-known groups such as hupehsuchians and thalattosaurs. The second half of the book consists of five chronological chapters that help you put all this diversity into some sort of logical order. This starts with life’s first coy attempts at reptiles-returning-to-the-sea in the Paleozoic Era, the main event of the Mesozoic in three chapters (the Triassic, Jurassic, and Cretaceous), and the Cenozoic Era in which the survivors of the K–Pg extinction continued and sometimes thrived.

The book’s second half stood out to me for two reasons. First, it helps prevent the same rookie mistake that is often made regarding dinosaurs: they did not all live at the same time. This may sound incredibly obvious and yet is easily and frequently forgotten. Where marine reptiles are concerned, a good example of this is that the ichthyosaurs evolved ~252 Ma ago and went extinct ~90 Ma ago, while the mosasaurs evolved ~100 Ma ago and went extinct 66 Ma ago at the K–Pg boundary, the two groups thus overlapping for “only” 10 million years. Turtles and crocodylomorphs survived the K–Pg extinction and positively flourished, though some groups subsequently went extinct and left no living descendants, such as the dryosaurids (a crocodylomorph lineage) going extinct ~40 Ma ago. The second reason I liked this chronological section is that it is largely told through the lens of key fossil localities around the globe (here amongst others Monte San Giorgio, Holzmaden, and the Oxford Clay). Though their names are often familiar and each of these deposits offers a unique view of a certain ecosystem at a certain time, they rarely get much attention themselves. The authors here provide just that little bit of extra information on their geography and stratigraphy, the history of their discovery and exploitation, and the palaeoenvironment that can be deduced from them.

“The authors tell the chronological section largely through the lens of key fossil localities that, though familiar-sounding to many readers, rarely get much attention themselves”

Ocean Life in the Time of Dinosaurs is richly illustrated in full colour with photos, diagrams, and palaeoart by Alain Bénéteau, including single and double-page spreads. There are several cladograms mapped onto timelines, with the simplified phylogeny of crocodylomorphs on page 69 particularly useful in visualizing the uncertain placement of thalattosuchians. Drawings show unique anatomical adaptations, explaining e.g. the evolution of turtle shells. The text is regularly interspersed with boxes discussing notable species or concepts such as proposed forms of swimming or adaptation of bones to life underwater. In short, the visual presentation of this book is outstanding.

I normally prefer to review each book on its own merits, but given that we now have two richly illustrated introductory books, there is no avoiding the mosasaur in the room. How does Ocean Life in the Time of Dinosaurs stack up against Darren Naish’s Ancient Sea Reptiles? As I also observed about his Dinopedia, Naish is particularly interested in phylogeny and species diversity. Whereas the discussion of the different groups here takes up 55 pages in chapter 2, Naish does this in 132 pages and six chapters. He goes into more detail on phylogenetic conundrums and for most groups discusses more species. What the French quartet here adds are the five chronological chapters, extending their coverage of evolutionary events to before and after the Mesozoic. As mentioned, they also give more detail on key fossil sites whereas Naish briefly mentions some of these in his chapter 1. My impression is that palaeontology buffs will want to get both books, despite the inevitable overlap. If, however, you are looking to buy just one book then Ocean Life in the Time of Dinosaurs is the most entry-level of the two, while Ancient Sea Reptiles provides more detail (and in that scenario would be my book of choice). My original observation regarding Greg Paul’s The Princeton Field Guide to Mesozoic Sea Reptiles, that it is more of a reference work to be consulted after either of these books, still holds.

Disclosure: The publisher provided a review copy of this book. The opinion expressed here is my own, however.

Ocean Life in the Time of Dinosaurs

Other recommended books mentioned in this review:

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Of all the insects that have a PR problem, cockroaches must rank very high. That, however, did not stop German entomologist, journalist, and filmmaker Frank Nischk from spending a year-long internship studying them. In this book, he regales the reader with stories of his time in the lab and the field studying first cockroaches and later crickets. A light and breezy read despite the serious undercurrent of biodiversity decline, Of Cockroaches and Crickets turned out to be an entertaining read.

Of Cockroaches and Crickets: Learning to Love Creatures That Skitter and Jump, written by Frank Nischk, published by Greystone Books in March 2023 (hardback, 214 pages)

This book was originally published in German in 2020 as Die fabelhafte Welt der fiesen Tiere by Ludwig Buchverlag and has been translated into English by Jane Billinghurst who frequently works with Greystone Books. Carl Safina contributes a short foreword that cracked me up and immediately set the tone. The book is effectively a memoir of Nischk’s early years studying for his undergraduate and doctorate degrees in the mid-nineties, told in 18 short chapters in two parts. His subsequent career pivot to documentary filmmaking only receives passing mention.

Given Nischk’s concern about biodiversity decline, and his desire to communicate to a broad audience why insects are fascinating and important, there is an irony to his undergraduate internship. He spent a year in the lab of Martin Dambach studying the aggregation behaviour of the German cockroach, Blattella germanica. By day, large groups of them bed down on their own excrement, likely attracted by pheromones released by the faeces. The irony? Nischk’s internship was funded by biotechnology giant Bayer which was hoping to isolate the chemicals responsible for putting the cockroaches in sleep mode to develop a pheromone-based cockroach trap: “the exterminator’s holy grail” (p. 25).

For his subsequent doctoral studies, Nischk got his conservation priorities in order. Staying with Dambach, he turned to crickets and spent time in Ecuador recording their songs. Next to discovering species new to science, this is his entry into the fascinating field of soundscape ecology or ecoacoustics. A small cadre of ecologists has been recording soundscapes of natural habitats. Bernie Krause (not mentioned here) is one particularly well-known example. By comparing recordings made years or decades apart they have shown how natural soundscapes are changing and often disappearing due to human encroachment. Others are hoping to train software to analyze recordings and identify species by their calls. If scaled up, the dream is to have passive acoustic monitoring stations in biodiversity hotspots around the globe.

“For his subsequent doctoral studies, Nischk […] spent time in Ecuador recording [cricket] songs. Next to discovering species new to science, this is his entry into the fascinating field of soundscape ecology.”

This backbone of his research is livened up with personal anecdotes and interesting asides. A friend’s call about a wasp infestation in her kitchen drawer is an excuse to introduce the 18th-century French entomologist Jean-Henri Fabre who was one of the first European naturalists to systematically collect and study butterflies, beetles, and wasps. Getting stung by a bullet ant in the rainforest of Ecuador leads to an aside about the late entomologist Justin O. Schmidt, the man who got stung for science (and wrote a fine book about it too). Tracking down a particularly loud cricket in Ecuador is the starting point for an unusual case where entomologists helped to defuse international political tensions between the USA and Cuba (this story has a surprising twist that I will not spoil here). A botched attempt to eradicate cockroaches that escape his experimental setup backfires most spectacularly, while fieldwork in the tropics is always fodder for amusing cultural misunderstandings and sober reflections. There is a nice mix here that never dwells on any one topic too long and makes for a book that is hard to put down.

The third and final part is, perhaps surprisingly, comparatively the weakest of the book. In four chapters Nischk muses on the biodiversity crisis, particularly the still poorly understood decline of insects, and discusses examples of individuals and organisations who are creating and protecting wildlife habitat. Probably most interesting are the little-known grassroots initiatives in Ecuador that are undertaken by villagers and farmers turning to ecotourism. But is this really the answer? Or does it merely perpetuate the idea that nature can only be protected if it has monetary value? You will not find a critical or comprehensive analysis of wildlife conservation here. There is also an odd focus on projects in the USA, e.g. the High Line in New York, the Xerces Society, Joan Maloof’s Old-Growth Forest Network, and the Yellowstone to Yukon Conservation Initiative. I wonder if this was added for the English translation. There is no mention of e.g. the European Natura 2000 network of protected areas or E.O. Wilson’s bold call to protect half the planet, and only passing mention of the German environmental organisation NABU or the practice of rewilding. Putting aside such nitpicking, none of this takes away from his genuine concern about the ongoing loss of biodiversity nor from his conclusion that the key to protecting species is protecting their habitat.

Overall, Of Cockroaches and Crickets is an amusing and light read that I devoured in a day. Nischk offers a nicely balanced blend of interesting natural history, amusing personal stories, and captivating scientific research. Whether it is flies, wasps, or rats, we need more books that celebrate those species we all too readily dismiss as pests.

Disclosure: The publisher provided a review copy of this book. The opinion expressed here is my own, however.

Of Cockroaches and Crickets

Other recommended books mentioned in this review:

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When it comes to popular science books, some of the books I admire most are the smallest ones. It takes great skill to capture the essence of a subject into a short book, steer clear of well-trodden ground, and contribute something novel that will educate and enthuse your reader. Palaeontologist and science communicator Dean Lomax here collects ten short essays on dinosaurs, convincingly showing that good things come in small packages.

Dinosaurs – 10 Things You Should Know: 230 Million Years for People Short on Time, written by Dean R. Lomax, published by Seven Dials (an Orion Books imprint) in September 2021 (hardback, 120 pages)

This small hardback is part of the 10 Things You Should Know series by Seven Dials, a popular science series that has so far covered physics, astronomy, mathematics, and neurology. Entry-level popular science books on dinosaurs, especially those aimed at children, often fail to go beyond factoids such as who was the biggest or who had the strongest bite. Sure, “these are cool topics to study, but they are not necessarily at the top of the research list for palaeontologists. Some of the most important research areas to explore focus on the bigger pictures of evolution” (p. 114). Thus, Lomax here makes pop science subservient to his primary goal of covering some of the hottest and most relevant topics grabbing the attention of professional palaeontologists today—from food, sex, and death; to deep time, biogeography, and the limits of our knowledge. And though he discusses fossil evidence of behaviour, the subject of his previous book Locked in Time, he avoids the easy option of rehashing material from that book.

Some of these essays do an excellent job of using familiar tropes to explain an important concept. As the first essay clarifies, Tyrannosaurus rex and Stegosaurus never crossed paths. Lomax gently dispels the popular media portrayal of dinosaurs all existing simultaneously, in the process introducing you to deep time. Stegosaurus was already a fossil by the time T. rex evolved. And while early dinosaurs roamed the supercontinent of Pangaea, the drifting apart of the continents meant that later species evolved in separate environments. In making this observation, this essay introduces concepts such as plate tectonics, the supercontinent cycle, and biogeography.

“[…] what I appreciated most in this book: Lomax repeatedly stresses how science is a dynamic process, not a static monolith of knowledge.”

Lomax is also not shy of cracking open the door to reveal the complexities and subtleties once you delve deeper. Having discussed how you define a dinosaur and introduced the division between saurischians and ornithischians, he mentions the 2017 study that proposed shaking up the dinosaur family tree, and how this is being tested and debated. This is what I appreciated most in this book: Lomax repeatedly stresses how science is a dynamic process, not a static monolith of knowledge. Point in case are feathers in dinosaurs and the growing realisation that birds might be related to…; no, they might actually be the descendants of…; no, wait, they are dinosaurs. What I did not realise is that Thomas Henry Huxley already advocated this idea shortly after the publication of Darwin’s On the Origin of Species, but that it was downplayed and ignored until John Ostrom revived it in the 1970s.

Another strong suit of this book is that it shows that you can be both brief and accurate in popular science. When introducing Darwin’s theory of natural selection he avoids that old chestnut of “survival of the fittest” and correctly characterises it as a stochastic process where “those animals with features better adapted to their environment have a higher chance of surviving and reproducing” (p. 98). And he acknowledges that Alfred Russel Wallace arrived at this idea around the same time as Darwin did. Thus, in just two sentences, he hits the nail on the head regarding two aspects of the theory of evolution that frequently get mischaracterised. Similarly, when discussing the idea of bringing back mammoths via a process known as de-extinction, he writes: “if this were to be successful, the resulting creation would never be an actual mammoth, but some oddball hybrid crossed with a modern elephant” (p. 116). And with that single sentence, he lays bare a major philosophical stumbling block of this controversial idea.

As befits a popular science book, there are plenty of interesting titbits (Why did Michael Crichton depict Velociraptor the way he did in Jurassic Park?), fascinating questions (Will we ever find all dinosaurs that ever existed?), and a healthy dose of humour. Entertaining and remarkably educational, this short book is perfect for both novice and seasoned palaeontology fans. Since we have entered December, I am going to earmark this book as a recommended stocking filler.

Disclosure: The publisher provided a review copy of this book. The opinion expressed here is my own, however.

Dinosaurs – 10 Things You Should Know

Other recommended books mentioned in this review:

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Seven Dial’s 10 Things You Should Know series:

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In preparation for Andy Secher’s new book Travels with Trilobites I decided to first reach back in time to read Richard Fortey’s 1999 book Trilobite! as a warm-up exercise. Why? For no other reason than that Fortey’s autobiography A Curious Boy impressed me so much that I bought several of his earlier books and I need an excuse to read them. This, then, is the first of a two-part dive into the world of that most enigmatic extinct creature: the trilobite.

Trilobite! Eyewitness to Evolution, written by Richard Fortey, published by Flamingo (a HarperCollins imprint) in March 2001 (paperback, 269 pages)

Fortey’s first encounter with a trilobite fossil at the age of 14 led to a life-long career as a palaeontologist and trilobite specialist at the London Natural History Museum, from which he retired in 2006. As a group, trilobites were tremendously successful, surviving for almost 270 million years from the early Cambrian (521 million years ago) to their extinction at the end of the Permian (252 million years ago), spanning nearly all of the Palaeozoic era. Looking through their eyes (which is something you can literally do), there are numerous fascinating facets to their story, and Fortey expertly reveals some of these here.

Take their curious biology. Their segmented forms are instantly recognisable, but look closer and you will see a “mixture of strangeness and familiarity […] The trilobites are lodged in this betwixt and between category, familiar as arthropods, yet strange in all their particularities” (p. 207). The fossils we find are frequently only (fragments of) cast-off shells. Trilobites went through numerous moults as they grew, just as crustaceans and insects do today, and thus were “veritable fossil factories” (p. 34). Their soft undersides were preserved only very rarely so we initially had no idea what their legs looked like. That answer came at the beginning of the 20th century from rock strata in New York State, which preserved exquisite detail in pyrite, and from the German Hunsrück Slate, where x-ray photos revealed limbs. It seems they were branching structures (biramous limbs in biological lingo) that split into a walking appendage and a second appendage that possibly held gill-like filaments involved in breathing.

“Their segmented forms are instantly recognisable, but look closer and you will see a “mixture of strangeness and familiarity”“

Consider, next, their growth. Starting as a disc-shaped larva, individuals would develop a split between the head and tail, with the tail-end budding off thoracic segments as the larva grew. When the species-specific number of thoracic segments was reached, the individual would simply grow larger with each subsequent moult without adding further segments. Finally, consider the eyes. Compound eyes like those of insects, yes, but uniquely made out of pure, transparent calcite. Fortey details how they worked and how the orientation of the individual hexagonal lenses has been used to reconstruct a trilobite’s field of vision. Some species with bulging eyes were able to see in almost all directions at once. And, as mentioned above, some researchers have even looked and taken photos through fossilised eyes, that is how well these structures preserve. Remarkably, some species with larger lenses even solved the problem of chromatic aberration by incorporating a thin, bowl-shaped layer of magnesium in the lens, nature pre-empting the very solution that scholars such as Huygens and Descartes would propose in the 16th century. For Fortey, they are a clear example of the tempting but mistaken belief of life’s story as one of linear progress. How would you compare the trilobite eye against that of the dragonfly or the wasp? “Who is to calibrate progress, who to legislate on the unit of improvement?” (p. 103).

But trilobites also enlighten other aspects of evolution, palaeontology, and geology. They were studied by Victorian geologists such as Sedgwick and Murchison who defined and named the geological periods we still use today. Part of Fortey’s work has been to use the distribution of species through time and space—their palaeobiogeography—as one line of evidence towards the larger project of reconstructing maps of our planet through time. As an example, the trilobite faunas found on the eastern and western sides of Newfoundland differ radically (Newfoundland was once split in two), while those of western Newfoundland match those recovered in Scotland (the two were once connected). Lists of trilobite species thus “provide the raw ingredients for maps of distribution: these, in their turn, describe the boundaries of former continents” (p. 195). The picture he restores shows how “around each ancient plate the continental shelves were stacked in order, carrying tier after tier of different trilobites, each minding their own particular business. […] trilobites dissected their world into throngs of niches: this is how they became the ‘beetles of the Palaeozoic’” (pp. 205–206).

“And then there is the matter of the Cambrian Explosion, which saw the sudden appearance of a wild diversity of arthropod forms. […] But how much of an explosion was this really?”

Other big-ticket evolutionary topics trilobites have shone a light on is that of punctuated equilibrium, the idea that evolution proceeds in fits and starts. Niles Eldredge would later popularise this idea together with Stephen Jay Gould, but it was his work on trilobites that provided the first spark. He saw periods of rapid species turnover followed by long periods of stasis which contrasted with the then-dominant idea of gradualism, of slow and steady evolutionary change (as Fortey shows, there are examples of this as well in the trilobite fossil record). And then there is the matter of the Cambrian Explosion, which saw the sudden appearance of a wild diversity of arthropod forms. “In popular accounts it became an ancient moment of madness, a magnificent evolutionary Mardi Gras, when a parade as bizarre as could have been devised by a surrealist on speed would be permitted for a geological day” (p. 125). But how much of an explosion was this really? Fortey has been critical of it and has drawn up family trees of the Burgess Shale invertebrates (made famous by Gould), showing high degrees of relatedness; we may have over-egged the peculiarities. And where are their Precambrian ancestors? One interesting idea he floats here is that trilobites appear suddenly in the fossil record thanks to an increase in size. Earlier, smaller forms were possibly not secreting shells, muscle support being less of an issue at tiny sizes; “the ‘explosion’ was a dramatic appearance of characters that had been rehearsing out of sight for more than a hundred million years” (p. 133).

Finally, the sometimes remarkable lives of individual scientists are wrapped up in the story of trilobites. Fortey mixes in some autobiographical elements (leaving plenty for later memoirs) but mostly highlights the lives of others. There is the tragic story of Rudolf Kaufmann, executed in World War II, who came to the same insights on punctuated equilibrium 40 years before Niles Eldredge (as Eldredge has acknowledged in his publications). There is the intriguing story of 19th-century French geologist and palaeontologist Jacques Deprat who build up a formidable reputation up to World War I, to then fall from grace when doubts arose about the identity of some supposed Chinese trilobite specimens. After being evicted from the Société Géologique de France in 1920, he took on a new identity and resurfaced as a successful novelist, a fact not revealed until his death in 1935. And Fortey shared offices with the Cambridge scholars that studied the Cambrian Explosion. He analyses in some detail the fall-out between Simon Conway Morris and his erstwhile mentor Gould over what exactly the invertebrates of the Burgess Shale tell us about the evolutionary history of life.

This multifaceted story is driven along by Fortey’s writing, which I found to be as captivating when he was writing in 1999 as when writing A Curious Boy in 2021. The book is complemented by black-and-white photography of notable species. One or two of the inline photos are poor-quality monochromes, but especially the plate sections contain some particularly fine photography. However, if it is visual splendour you want, we need to turn to Travels with Trilobites, which I will review next.

Trilobite!

Other recommended books mentioned in this review:

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This is the second of a two-part dive into the world of pterosaurs, following on from my review of Mark Witton’s 2013 book Pterosaurs. Almost a decade later, the well-known independent palaeontologist and palaeoartist Gregory S. Paul has written and illustrated The Princeton Field Guide to Pterosaurs. Admittedly, a field guide to extinct creatures sounds contradictory. Really, this is an illustrated guide for the palaeo-enthusiast in which Paul’s signature skeletal reconstructions take centre stage.

The Princeton Field Guide to Pterosaurs, written by Gregory S. Paul, published by Princeton University Press in June 2022 (hardback, 184 pages)

This is Paul’s third Princeton Field Guide to prehistoric life. Paul wrote and illustrated The Princeton Field Guide to Dinosaurs in 2010, followed by a second edition in 2016 (which is when Donald Prothero wrote The Princeton Field Guide to Prehistoric Mammals) and a third edition in 2024. A fourth book was published in October 2023 with The Princeton Field Guide to Mesozoic Sea Reptiles which completed Paul’s informal trilogy covering all the major reptile groups of the Mesozoic. Finally, The Princeton Field Guide to Predatory Dinosaurs has been announced for November 2024. All these guides follow a unified style. Indeed, when comparing the dinosaur guide with this book, I noticed that it closely follows the same format and layout, down to very similar wording in some sections.

The Princeton Field Guide to Pterosaurs is divided into two halves: an introduction, and the group and species accounts. The introduction opens with the history of pterosaur research, describes what kind of reptile pterosaurs were and who they were related to, and gives some basics on dating fossils. The problem Witton flagged up in 2013 holds a decade on: clear transitional fossils showing how pterosaurs became airborne are still lacking. Paul then introduces the world in which pterosaurs evolved and I liked how he effectively zooms out to provide more context. What was the world like in the Triassic? What other organisms lived in it? How did all this interact throughout the approximately 162 million-year lifespan of the pterosaurs? And what creatures exploited their ecological niche after they went extinct?

Most of the introduction is dedicated to pterosaur biology, giving an overview of anatomy, locomotion and flight, pneumatic skeletons, integument and colouration, physiology, sensory biology, diseases and injuries, behaviour, and metabolism and growth. Much of the material here agrees with what Witton wrote in 2013, but Paul provides a more thorough introduction to locomotion and especially flight. Where Witton covered flight in eight pages, Paul lavishes 22 pages on the topic, going into quite some technical depth by making thorough comparisons to birds, bats, and aircraft, and drawing on aerodynamics and aeronautics. Given the way he references classic fighter planes, I got the impression he has at least an abiding interest in aeroplanes. Another noticeable difference is that Paul is bolder in his assertions and opinions as to what pterosaurs did or were capable of. For example, since reptiles and birds have full-colour vision extending into the UV range, “pterosaurs almost certainly did as well” (p. 70). I noticed the same in the species accounts, where data is presented in bullet-point statements, leaving little room to indicate how robust or speculative ideas are about e.g. lifestyles and foraging strategies of different species. A final section describes where fossils are typically found and how many species there might have been. Paul also engages in a bit of speculative zoology, imagining a spot of time travel to go on a pterosaur safari, and what might have happened had pterosaurs not gone extinct.

“Paul lavishes 22 pages on the topic [of flight], going into quite some technical depth by making thorough comparisons to birds, bats, and aircraft, and drawing on aerodynamics and aeronautics. “

The second half of the book discusses groups and species. Where Witton made taxonomy the leitmotif of his book, picking a phylogeny and dedicating a chapter to each major group, Paul refuses to put his nickel down. He recognizes the division between rhamphorhynchoids (a name he admits is obsolete but that he prefers over “nonpterodactyloid pterosaurs”) and pterodactyloids, divided into basal and derived forms. But since there is no consensus and phylogenetic studies are too conflicting for his liking, he shows no family tree. He does arrange species in a phylogenetic order, with derived groups nested inside basal clades, but he uses “a degree of personal choice and judgment in arranging the groups“, some of which reflect “my considered opinion, but most are arbitrary choices among a large array of competing research results” (p. 91). A second difference with Witton’s approach is that Paul writes the species entries as if this were a real field guide, with telegram-style entries describing size and weight estimates, known fossil remains, anatomy, geological age, location and geological formation of fossils, habitat, lifestyle, and other notes. This is the reference section of the book, collecting facts without a guiding narrative.

A stand-out feature of this book is the illustrations, specifically the skeletal reconstructions. Paul’s signature style of white bones on the black outline of a body is well-known and one he has been practising for decades. Rather than full skeletons, most show only the bones that have been found for a species, positioned in the right place. Where sufficient fossil material is available, full skeletons are shown. Some of these, Paul mentions, are composites. For a few species there is even enough material to show hatchlings, juveniles, or immature individuals. Paul criticizes the quadrupedal launch pose used by Witton and others for skeletal reconstructions as visually awkward and obscuring wing shape and proportions. He instead shows animals taking off with the wing at the top of an upstroke. I actually agree with Paul’s choice of pose: it looks more familiar, resembling a large bird taking off. For a select few species, there are muscle studies, which he admits are speculative but based on sound anatomical principles, as well as life reconstructions. Lastly, there are some palaeoart pieces showing animals in their natural environment. I admit I was not fond of all of these; some show a pencil-drawn animal placed on a photographic background, which is visually rather jarring. Notably absent are photos of fossils.

“A stand-out feature of this book is the illustrations, specifically the skeletal reconstructions. Paul’s signature style of white bones on the black outline of a body […] show only the bones that have been found for a species, positioned in the right place.”

So, what has happened in the world of pterosaur research since Witton’s 2013 book? Paul makes no particular point of highlighting significant advances, and without an expert background, the different approach of the two books makes it quite hard to determine this. Two studies that caught my attention were the evidence for pterosaurs eating cephalopods (quite logical) and a study on istiodactylid hyoid bones (the bone to which the tongue attaches). Obviously, Paul references new papers describing new species: China has continued to be a goldmine; Witton was involved in the 2017 description of the giant azhdarchid Hatzegopteryx thambema; and I found the 2014 description of Ikandraco avatar equally interesting. The last one is mentioned as a potential skim-feeder, a foraging strategy Witton has strongly rejected before. Maybe this time it is for real?

To wrap up, how do the two books compare? Pterosaurs is more narrative-driven and Witton is more circumspect in his claims; The Princeton Field Guide to Pterosaurs is more data-dense and Paul is bolder in his claims. If you are completely new to pterosaurs, I would actually recommend that you start with the former; I found its overview of the different groups more accessible. Then, if you want to read deeper into the details or need a reference work, the latter is a good follow-up; the two books complement each other nicely that way. If you already have Witton’s book, I think it is worthwhile getting Paul’s book. Next to being up to date, The Princeton Field Guide to Pterosaurs offers a different, sometimes more thorough take, and much more factual detail.

Disclosure: The publisher provided a review copy of this book. The opinion expressed here is my own, however.

The Princeton Field Guide to Pterosaurs

Other recommended books mentioned in this review:

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