6-minute read
keywords: paleontology

Reconstructing how dinosaurs behaved from just their fossilised bones might seem like science fiction but is very much science fact. Join me for a double review of two recent illustrated books. I will next review An Illustrated Guide to Dinosaur Feeding Biology but first up is Dinosaur Behavior: An Illustrated Guide. Here, veteran palaeontology professor Michael J. Benton joins forces with palaeoartist Bob Nicholls to do what it says on the tin: write a richly illustrated introductory book on dinosaur behaviour that is well-suited for novices.

Dinosaur Behavior: An Illustrated Guide, written by Michael J. Benton, illustrated by Bob Nicholls, published by Princeton University Press in October 2023 (hardback, 224 pages)

Dinosaur behaviour has come up previously on this blog. Locked in Time, also illustrated by Bob Nicholls, featured 50 vignettes of famous fossils, while Dinosaurs Without Bones was a chunky pop-science book. In Dinosaur Behavior, Benton takes the reader through five main topics: physiology (which sets the pace for everything else), locomotion, senses and intelligence, feeding, and social behaviour (which includes courtship, reproduction, parental care, and communication). One or several “forensics” boxes in each chapter introduce the basic gist of certain methods.

Reading through this book, it becomes abundantly clear that our understanding of dinosaur behaviour relies on two approaches. Though Benton does not mention it as explicitly as in his previous book The Dinosaurs Rediscovered, the first of these is new high-tech toys and tools. Examples include computed tomography (CT) scanners normally used in hospitals to make detailed X-ray scans of fossilised brains (so-called endocasts) and so determine brain anatomy, or finite element analysis normally used in engineering to model forces and stresses on jaws and teeth and so determine e.g. bite force. The second approach is “old-fashioned” comparative anatomy and ethology: it pays to have a good knowledge of natural history when you are a palaeontologist. One example is the histological study of fossil dinosaur bones. Cutting thin bone sections and examining these under a microscope shows that some dinosaurs closely resemble mammals and birds, supporting the idea that smaller species were endotherms (“warm-blooded”, i.e. generating their own body heat). Or take the microscopic study of melanosomes (pigment-containing organelles) in fossil feathers to determine colour in life. A final example is the comparison of footprints made by modern running birds with fossil tracks to determine things such as gait and running speed.

“it becomes abundantly clear that our understanding of dinosaur behaviour relies on two approaches […] new high-tech toys and tools [and] “old-fashioned” comparative anatomy and ethology.”

If you are well-versed in (popular) palaeontology, much of what is presented here will be familiar. Even so, I picked up interesting titbits. One example is a recent study of Psittacosaurus that describes a cloaca, the multipurpose orifice also seen in birds where the digestive, urinary, and reproductive tracts all open to the outside world. This suggests that dinosaur sex for at least some species was a matter of the appropriately named cloacal kiss rather than the brandishing of reptilian genitals. Other insights fell into the embarrassing “I should have known this” category. We tend to think of walking on two legs as something advanced because our mammalian ancestors walked on all fours, but for dinosaurs, it was the reverse; they started out bipedal and quadrupedality only evolved later in e.g. the large sauropods. Particularly interesting is the study by Kat Schroeder and colleagues who looked at fossil communities of theropods and noticed a so-called carnivore gap: there is a lack of medium-sized ones in the fossil record, even though there are medium-sized herbivores. One explanation could be that dinosaur eggs had an upper size limit, meaning that young carnivores hatched small and had an awful lot of growing to do. As they did, “they passed through a whole range of feeding modes, each step along the way acting like a different species” (p. 137), effectively plugging the ecological niche of medium-sized carnivores.

Despite the broad range of topics, there are some curious omissions. The chapter on feeding e.g. discusses jaws, teeth, and the use of isotopes to determine diet, but not microwear analysis of teeth. What I found most surprising is that Benton does not introduce the concept of trace fossils or ichnology, their study. Yet, examples such as trackways (some possibly showing long-distance migrations), coprolites (fossil poop), and nests are all discussed here. Another surprising omission is that the two-page bibliography does not include most studies mentioned in the text, even though it references other technical articles.

“dinosaur sex for at least some species was a matter of the appropriately named cloacal kiss rather than the brandishing of reptilian genitals.”

Dinosaur Behavior is mostly very suitable for readers with little to no background in palaeontology. Benton explains even basic terminology (physiology, cannibalism) as he goes, though there is the occasional curveball. One example is the morphospace diagram showing a principal component analysis on page 131, which, I hope those with a background in statistics will agree, is a rather abstract way of visualizing data that requires a bit more explanation than is given here. Though the book is published by Princeton University Press, it has been produced by UniPress Books who, as I explained in a footnote to another review, can be considered the spiritual successor to popular science publisher Ivy Press. What this means is that information is accessibly presented in bite-sized sections on one or several page spreads, with long sections further divided using subheadings. The downside is that this restricts how thoroughly topics can be explored. Leafing through e.g. Naish & Barrett’s Dinosaurs: How They Lived and Evolved shows more nuance in its chapter on behaviour.

Finally, I have to mention the excellent colour and black-and-white artwork by Bob Nicholls that livens up the text. I loved the drawing of courtship in Confusiusornis on pages 168–169. Despite the overlap in topic, this is all-new artwork compared to Locked in Time. Other diagrams have all been carefully designed or redrawn, using colours where appropriate. The only design element that did not work for me was the choice of sans-serif font which made e.g. the letters a and o hard to tell apart.

Overall, this is a handsomely illustrated book that offers an accessible introduction suitable for novices and possibly even curious high-school pupils. Since it is December, I will add the words “gift suggestion” to that. Admittedly, serious palaeontology buffs might find the contents here somewhat superficial. So what if you wanted to go deeper on one of these topics, what would that look like? I will next turn to An Illustrated Guide to Dinosaur Feeding Biology to find out. [Edit: As of November 2024, the logical follow-up I would recommend is David Hone’s Uncovering Dinosaur Behavior.]

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

Dinosaur Behavior

Other recommended books mentioned in this review:

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Though I could not paint or draw a dinosaur if my life depended on it, I never tire of reading about palaeoart. What a treat, then, that Crowood Press revisits this subject with this book by US palaeoartist Emily Willoughby. Very much a resource for those already familiar with basic art techniques, it counsels the reader on what goes into making believable and memorable palaeoart. Featuring foremost Willoughby’s favourite subject, feathered dinosaurs, the book also doubles up as a beautiful portfolio of her artwork, showcasing her mastery of a wide range of media.

Drawing and Painting Dinosaurs: Using Art and Science to Bring the Past to Life, written by Emily Willoughby, published by Crowood Press in October 2021 (paperback, 176 pages)

Willoughby juggles her palaeoart with a passion for bird photography and a research career in behavioural genetics. Her work has graced scientific articles, museum exhibitions, magazines, newspapers, websites, and, of course, books (e.g. in Dinosaur Art II). To me, a standout feature of Drawing and Painting Dinosaurs is her philosophy. First, palaeoart is not just educational but is a restoration of an individual animal’s life, “a sort of homage to the dead” (p. 8). The artist has an obligation to render an accurate depiction, not just to the public, but also to the fossil that was once alive. “It is our unique responsibility to make sure that these creatures are not forgotten to the dust of time” (p. 19). Second, palaeontology and palaeoartistry are intimately entwined, each having the potential to advance the other field.

Drawing and Painting Dinosaurs is loosely divided into two parts, both of which are informed by this dedication to realistic and lifelike artwork. The first half covers general principles, while the second half applies these to four groups of dinosaurs. The whole is liberally illustrated with over 250 images, many unique to this book. Willoughby masters a range of techniques: oil paint, gouache, graphite, digital media, watercolour, pencil, and ink. I especially like the graphite works and the family trees, and I love how many of her works are impressionistic rather than photorealistic. The art portfolio is worth the price of admission alone. Admirably, she even includes a few comparisons that show how artwork can quickly become outdated and includes the sobering lesson that “in all probability, and with few exceptions, all palaeoart will one day be shown to be inaccurate as the science of palaeontology marches ever forward. And that is okay” (p. 50).

“palaeoart is not just educational but is a restoration of an individual animal’s life, “a sort of homage to the dead“”

Doing background research is of paramount importance to palaeoart and, uniquely, Willoughby offers readers a guide to parsing the scientific literature. What kinds of academic papers are published in the field of palaeontology? And what kind of information of relevance to palaeoart can you distil from a description of a new dinosaur, a classification paper, or a biomechanical study? The other starting point is the fossils themselves. Willoughby reminds us how much palaeoartists owe museum preparators, whose efforts to piece back together incomplete and deformed fragments are often overlooked. Third, context is everything. By studying the geologic formation in which a fossil is found you can prevent anachronisms and populate your artwork with plants, insects, and other dinosaurs that were alive at the same time as your focal animal.

Having laid out these general principles, Willoughby then applies this to particular groups. She does this most thoroughly for her favourite subject, the dromaeosaurs: small to mid-sized feathered theropod dinosaurs such as Deinonychus and Velociraptor. Thanks to exquisitely preserved fossils from China, such as the Jehol biota, feathered dinosaurs are no longer controversial. Given their evolutionary relationship, the skeletons and general anatomy of modern birds is a good guide, though there are key differences to keep in mind. Similarly, the colouration of modern birds can function as an inspiration, but Willoughby warns against copying specific patterns, that can “turn into insufferable memes in palaeoart” (p. 89). Here, too, understanding the underlying biology is important. Blue is rare in nature, usually resulting from (micro)morphological structures, while greens or reds result from pigments obtained through diet, so might not be appropriate for the dinosaur you are drawing. And serious thought needs to be given to the behaviour you wish to depict.

“Willoughby explains how to come to a plausible reconstruction using the principles of phylogenetic bracketing and parsimony”

The topic of feathers is of such interest to Willoughby that she returns to their evolution in the last chapter. Their structure ranged from hollow, cylinder-like protofeathers, to messy feathers with filamentous barbs that gave a shaggy appearance, to, ultimately, neatly interlocking flight feathers. The process, meanwhile, was a “non-linear path to aerodynamic feathers and powered flight” (p. 166), complete with evolutionary dead ends, convergent evolution, and secondary loss of flight. This chapter gives an overview of what we know about feather evolution and how it maps onto the dinosaur family tree. Willoughby explains how to come to a plausible reconstruction using the principles of phylogenetic bracketing (looking at an animal’s closest relatives and the traits they did or did not have) and parsimony (minimising the number of assumptions).

So far, I am mightily impressed and it should be obvious that this book is invaluable to both aspiring and seasoned palaeoartists. That said, there are a few caveats to keep in mind. This book strikes me as a conversational guide to creating palaeoart first, and a hands-on manual second. There are some useful practical tips sprinkled throughout the book (e.g. tracing over images of bones, making simple 3D models, using masking fluid to block part of a painting), while step-by-step examples mostly show you “how to draw dinosaur X from basic shapes” or show digital artwork in various stages. Furthermore, despite the title, this book does not give equal airtime to all groups but focuses on feathered dinosaurs. There is advice for drawing large carnivores, represented by the Tyrannosauroid family, and sauropods, but only brief introductions to the biology of other large herbivores. Simultaneously, the title undersells the book a bit: search for it online and you will find many generic guides which this book patently is not! I guess “The Palaeoartist’s Handbook” was already taken. Speaking of which…

“Willoughby explains how to come to a plausible reconstruction using the principles of phylogenetic bracketing and parsimony”

I have so far ignored the Triceratops in the room, namely Mark Witton’s 2018 book that was Crowood Press’s first venture into palaeoart. How do the two compare? Both stress thorough background research and originality in subject and composition, but Willoughby, who contributed illustrations to Witton’s book, has been careful to avoid too much overlap. Witton uses many anatomical diagrams to discuss soft-tissue reconstruction in detail and has valuable advice on palaeoart as a paid profession, but he did not cover the actual process of drawing on which Willoughby has some tips. If you already purchased The Palaeoartist’s Handbook then Drawing and Painting Dinosaurs is a very welcome companion. If you have neither you will want to get both.

All said, this book is fantastic value for money and it is a real treat to have another established palaeoartist take the time to write a coherent overview of their methodology. Finally, if you want to know more about the process of making this book, Witton published an interview with Willoughby on his blog while the good folk over at Palaeocast interviewed her for an episode of their podcast.

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

Drawing and Painting Dinosaurs

Other recommended books mentioned in this review:

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After three previous books in this format on fossils, rocks, and dinosaurs, geologist and palaeontologist Donald R. Prothero here tackles the story of evolution in 25 notable discoveries. More so than the previous trio, this book tries to be a servant to two masters, resulting in a mixed bag.

The Story of Evolution in 25 Discoveries: The Evidence and the People Who Found It, written by Donald R. Prothero, published by Columbia University Press in December 2020 (hardback, 360 pages)

Prothero has organised The Story of Evolution in 25 Discoveries in a logical fashion. After convincing the reader that the universe and our planet are, indeed, really old, he considers some of Darwin’s lines of evidence for evolution, followed by several great transitions in evolution as revealed by the fossil record, more recent evidence from genetics and molecular biology, and, of course, evidence for the evolution of humans.

As in his previous books, Prothero manages to dig up some remarkable stories. For example, Darwin initially mistook the finches on the Galápagos for wrens, blackbirds, and other species. Only when he handed them to the famous ornithologist John Gould for a second opinion did it become clear that these were all finch species adapted to local conditions on the different islands. It later fell to others such as Peter and Rosemary Grant to do the long-term studies that elevated them to the icon of evolution they have become. Meanwhile, Othniel Charles Marsh’s monograph on primitive birds that still had teeth was unexpectedly branded a waste of taxpayer’s money when US congress was looking for excuses to cut funding to the US Geological Survey in the 1890s.

In many places, Prothero is careful and balanced in his coverage. He highlights the contribution of the historically overlooked Alfred Russel Wallace who independently hit on the idea of natural selection after Darwin had already been labouring on it for decades. And while Ernst Haeckel was accused of fraud over his famous drawings showing the embryonic development of different vertebrates, Prothero explains how there is a kernel of truth to Haeckel’s claim that ontogeny recapitulates phylogeny, even if “he may have been a bit overzealous in his drawings” (p. 69). On Jean-Baptiste de Lamarck and his ideas, Prothero clarifies how there was much more to him than the caricature of the “guy who got evolution wrong” (p. 198) that he became.

“In many places, Prothero is careful and balanced in his coverage […] he clarifies how there was much more to Jean-Baptiste de Lamarck […] than the caricature of the “guy who got evolution wrong””

There are classic topics such as convergent evolution, the evolution of the eye, and Lynn Margulis and her theory of endosymbiosis. The relatively young branch of evolutionary developmental biology and the discovery of Hox genes showed that, actually, yes, nature does make leaps and does not always result in slow and gradual changes. Throughout, Prothero repeatedly reminds you that the evolutionary relationships between organisms are like a bush, and not a linear progression from primitive to more advanced creatures. He explains that evolution does not always result in perfect adaptations—they only have to be good enough to help in producing the next generation. And he points out that natural selection can only ever work with the material at hand, resulting in many jury-rigged contrivances, including in humans.

Now, Prothero is also a noted sceptic. A good deal of this book has the secondary aim of showing that creationism is an utterly flawed idea and that the evidence for evolution reveals no traces of intelligent design whatsoever. The thing is, he already did this exercise in Evolution: What the Fossils Say and Why It Matters, so does it need repeating? He spends no fewer than seven chapters here on examples of transitional fossils that provide a detailed picture of fish leaving the water, whales returning to it, birds evolving from dinosaurs, giraffes evolving longer necks and elephants longer trunks, horses losing their toes and snakes their legs, and turtles acquiring shells. You almost get the feeling that he just cannot help himself.

I have few gripes with the topics that Prothero chose to include here, but I felt somewhat disappointed by all the topics he left out as a consequence of this secondary mission. Two notable omissions are the process of domestication, even though Darwin used numerous examples of it in On the Origin of Species and then wrote a separate book about it. The same is true for Darwin’s other big idea: there is no mention of sexual selection, sperm competition, or mate choice.

“[…] this book has the secondary aim of showing that creationism is an utterly flawed idea […] but I felt somewhat disappointed by all the topics he left out as a consequence […]”

Beyond these, there is little on speciation and biodiversity, the formation of higher taxa, or the difficulty with species concepts. Epigenetics is mentioned, but not by name. The textbook example of the peppered moth is included, but there is no further discussion on camouflage, mimicry, or warning signals. Richard Dawkins has to make do with two brief mentions, but there is nothing about the different levels of selection, whether selfish genes or group selection. And I am sure Prothero could have beautifully explained the difference between the modern and extended syntheses.

The focus on convincing the reader of the evidence against design raises the question of who this book is written for. For evolutionary biologists like myself, Prothero is preaching to the choir, while creationists are unlikely to pick this book up. The best this might achieve is to remind biologists of the evidence for evolution if we ever find ourselves debating creationists.

Prothero is a fantastic science communicator, and I really enjoy this format of 25 vignettes by which to examine the many facets of a topic. The material that he did choose to include is written with verve and balance. In my opinion, however, the dual motive underlying The Story of Evolution in 25 Discoveries means that he left out many relevant topics and has written a book of narrower focus than the title might suggest.

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

The Story of Evolution in 25 Discoveries

Other recommended books mentioned in this review:

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When I recently reviewed The Real Planet of the Apes, I casually wrote how that book dealt with the evolution of Old Work monkeys and apes, ignoring New World monkeys which went off on their own evolutionary experiment in South America. But that did leave me wondering. Those New World monkeys, what did they get up to then? Here, primatologist Alfred L. Rosenberger provides a comprehensive and incredibly accessible book that showed these monkeys to be far more fascinating than I imagined.

New World Monkeys: The Evolutionary Odyssey, written by Alfred L. Rosenberger, published by Princeton University Press in September 2020 (hardback, 350 pages)

Most people are probably not very familiar with these monkeys. Technically known as platyrrhines, they are predominantly arboreal (i.e. living in trees), small to medium-sized primates. You might know the insanely loud howler monkeys from nature documentaries. Perhaps you have heard of capuchin monkeys or spider monkeys. But you could be forgiven for not having heard of marmosets and tamarins, or the even more obscurely named titis, sakis, and uacaris. A total of 16 genera are recognized, but outside of the scientific literature and abovementioned technical books, these monkeys are not all that well known. And that is a shame as, from an evolutionary perspective, this is a unique group.

Now, before Rosenberger gets to this, it helps to better know these monkeys. Accompanied by many excellent illustrations and photos, the first half of New World Monkeys is dedicated to their ecology, behaviour, and morphology. Topics covered include their diet and dentition; locomotion and the anatomy of hands, feet, and prehensile tails; but also brain size and shape; and their social organization and ways of communicating via sight, sound, and smell.

The platyrrhines are a diverse bunch with some remarkable specialisations. In the family Cebidae we find the smallest members, some of whom, the Marmosets and Pygmy Marmosets, have teeth specialized for gouging the bark of gum trees and feeding on the gum that is released in response. In the family Pitheciidae we find the only nocturnal member, the Owl Monkeys, which have concomitant morphological adaptations such as enlarged eyes. In both this and the closely related Titi Monkeys, individuals have the adorable habit of twining their tails when e.g. socializing or sleeping. The family Atelidae is home to species with exceptionally prehensile tails whose underside ends in a pad with a fingertip-like surface. The Muriquis and the aptly-named Spider Monkeys use them as a fifth limb in locomotion, as demonstrated by a striking photo of a Black-faced Spider Monkey on plate 13. Here we also find the well-known Howler Monkeys, whose skull is heavily modified to support the exceptionally loud vocal organs in their throat and neck.

“[…] the whole radiation has been remarkably stable for at least 20 million years. Today’s New World monkeys are virtually unchanged from their ancestors […] This stands in sharp contrast to the evolutionary history of Old World monkeys […]”

Despite these differences, platyrrhines are closely related and form what is called an adaptive radiation. Just like the textbook example of Darwin’s finches, many members have evolved unique adaptations and ways of living to minimise competition and maximise resource partitioning. Two ideas feature prominently in this book to explain how platyrrhines have evolved and what makes this adaptive radiation both so diverse and so interesting.

One idea is what Rosenberger calls the Ecophylogenetics Hypothesis. If I have understood him correctly, this combines information on a species’s ecology and phylogeny, its evolutionary relationships. It can offer hypotheses on how ecological interactions have evolved, but it also recognizes that ecological adaptations are shaped and constrained by evolutionary relatedness. For the platyrrhines, taxonomically related members are also ecologically similar. To quote Rosenberger: “[…] phylogenetic relatedness literally breeds resemblance in form, ecology, and behavior” (p. 96) and “Each of the major taxonomic groups that we define phylogenetically is also an ecological unit […]” (p. 97).

The other idea that makes the platyrrhines so interesting is dubbed the Long-Lineage Hypothesis. An extensive chapter on the fossil record documents how the whole radiation has been remarkably stable for at least 20 million years. Today’s New World monkeys are virtually unchanged from their ancestors, living the same lifestyles and occupying the same ecological niches. Some fossils have even been classified in the same genus as their living counterparts. This stands in sharp contrast to the evolutionary history of Old World monkeys where there has been a constant churn, whole groups of primates evolving and going extinct with time.

“Without ever losing academic rigour or intellectual depth, Rosenberger quietly proves himself to be a natural-born teacher and storyteller […]”

What stands out, especially when Rosenberger starts talking taxonomy and evolution, is how well-written and accessible the material here is. He takes his time to enlighten you on the history, utility, and inner workings of zoological nomenclature, making the observation that “names can reflect evolutionary hypotheses“. Here, finally, I read clear explanations of terms such as incertae sedis (of uncertain taxonomic placement), monotypic genera (a genus consisting of only a single species), or neotypes (a replacement type specimen). Similarly, there are carefully wrapped lessons on how science is done—on the distinction between scenarios and hypotheses, or how parsimony and explanatory efficiency are important when formulating hypotheses. Without ever losing academic rigour or intellectual depth, Rosenberger quietly proves himself to be a natural-born teacher and storyteller, seamlessly blending in the occasional amusing anecdote.

A final two short chapters conclude the book. One draws on the very interesting question of biogeography, i.e. on how platyrrhine ancestors ended up in South America, which was long an island continent. Rosenberger convincingly argues against the popular notion of monkeys crossing the Atlantic on rafts of vegetation* and in favour of more gradual overland dispersal. The other chapter highlights their conservation plight as much of their tropical forest habitat has been destroyed by humans.

With New World Monkeys, Rosenberger draws on his 50+ years of professional experience to authoritatively synthesize a large body of literature. As such, this book is invaluable to primatologists and evolutionary biologists and should be the first port of call for anyone wanting to find out more about the origins, evolution, and behaviour of these South and Central American primates.

* One mechanism that Rosenberger does not mention is that tsunamis could be behind transoceanic rafting, as argued in a recent Science paper. This looked at marine species in particular and I doubt it would make much of a difference for terrestrial species. Most of the objections Rosenberger gives would still apply.

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

New World Monkeys

Other recommended books mentioned in this review:

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When thinking of human ancestors, the name “Lucy” will likely come to mind. But a dedicated team of scientists spent decades labouring on the discovery of a species more than a million years older still, at 4.4 million years of age. Nicknamed “Ardi” and classified as Ardipithecus ramidus, it was finally revealed to the world in 2009. For a full decade, journalist Kermit Pattison immersed himself in the story of Ardi’s discovery to bring to life both the science and the scientists. The resulting Fossil Men is an incredibly well-researched book that tells the definitive insider’s story of how one of the most divisive fossils in palaeoanthropology was discovered by one of its most divisive characters: Tim White.

Fossil Men: The Quest for the Oldest Skeleton and the Origins of Humankind, written by Kermit Pattison, published by William Morrow & Co. in December 2020 (hardback, 534 pages)

Fossil Men stands out for its brutally honest portrayal of the main protagonist, Tim White, something for which Pattison has had the full cooperation of White, his co-workers, and his many adversaries. From the very first pages, he is unsparingly described as a relentless perfectionist with “a razor intellect, hair-trigger bullshit detector, short temper, long list of discoveries, and longer list of enemies” (p. 2). He will take as long as he darn well needs to make sure his findings are beyond bulletproof. And while many loathe being at the receiving end of his withering criticism—”enemies not only resented him; they fucking hated him” (p. 71)—many also admit that he excels at what he does: “Tim White is brutal. He’s a real scientist. His literature will stay forever” (p. 4).

Pattison follows White’s story of fossil discovery in Ethiopia from its start in 1981. Against a backdrop of bureaucracy, corruption, and civil wars, White organised annual expeditions that “he micromanaged without apology“, while fellow anthropologist Bruce Latimer marvels that he is “without question the best field worker there is […] His science, logistics, and efficiency are phenomenal” (p. 127). What stands out, and endeared him to this reader, is how he insists on local capacity building, employing and training numerous Ethiopians, and only rarely recruiting foreigners, the Japanese Gen Suwa being one of these notable exceptions.

As a journalist who has published in the New York Times, GQ, and other outlets, it comes as no surprise that Pattison engagingly portrays the human interest story. He has extensively interviewed the people around White, including Ethiopian collaborators such as Berhane Asfaw, Alemu Ademassu, and Yohannes Haile-Selassie, or the creationist-turned-palaeoanthropologist Owen Lovejoy. But he also speaks to White’s many adversaries, including Ian Tattersall, Lucy team members Jon Kalb and the celebrity-loving Don Johanson, as well as members of the world-famous Leakey dynasty.

“Fossil Men stands out for its brutally honest portrayal of the main protagonist, Tim White”

What positively surprised me, however, was how thoroughly and accurately Pattison delves into the biological details. Whether it is related species such as Ardipithecus kadabba, the anatomy of the foot and spine, or the genetical wonderland of ancient DNA, evolutionary developmental biology, or comparative genomics—he puts to good use the more than 500 pages that have been crammed into Fossil Men. White’s obsessive attention to detail seems to have rubbed off on Pattison, and in the acknowledgements he admits the enormity of writing this book: “Nobody in their right mind takes on a project like this” (p. 423). The bibliography reveals just how deeply he has ventured: next to books and peer-reviewed papers, Pattison has consulted unpublished manuscripts, interviews, grant proposals, video material, oral histories, court files, and archives of private correspondence.

Over the years, White’s team unearths fragmentary remains and puzzles together the creature that will become known as Ardi. However, much to the frustration of peers and grant agencies, White refuses to go public until he has every detail nailed down. This is not a popular strategy and the briefly-featured Michel Brunet was similarly scolded in Ancient Bones for holding back important material. After an initial 1994 announcement in Nature, White and collaborators labour for a full 15 years in strict secrecy. Pattison here exclusively reports the many twists, turns, and realisations during this long period.

The crescendo of Fossil Men comes with the big reveal of Ardi in a 2009 special issue of Science. As expected, the popular press laps it up while the academic establishment is initially more resistant. Pattison again excels at showcasing the range of opinions. Reading the objections it is easy to see how White’s arguments go against the grain. Specifically, A. ramidus was initially considered to be the most chimp-like ancestor known so far. However, the more White’s team looked at the anatomy of the feet, pelvis, skull, and other body parts, the more they argued that human ancestors never went through a stage resembling modern apes. The long-held paradigm that modern apes are good models of our human ancestors was declared dead. Instead, compared to Ardi, humans are the ones retaining primitive anatomical aspects, while modern African apes are considered evolutionarily more derived. It would require several instances of convergent evolution amongst apes for this to happen, and is thus considered less parsimonious by evolutionary biologists, but that does not mean it is impossible.

“Ardi, it seems, has everyone flummoxed, proving to be “a simian-human combination that nobody had predicted“”

Ardi, it seems, has everyone flummoxed, proving to be “a simian-human combination that nobody had predicted” (p. 353). Refreshingly, some opponents have come around to White’s arguments after they have been given the opportunity to inspect the fossils for themselves. Another supporter is David Begun who dedicated a section to A. ramidus in The Real Planet of the Apes, discussing the configuration of the big toe. He agrees with White that “meticulous collection of data should come before interpretation“, and highlights how the assumption that hominin ancestors by definition cannot have a grasping big toe imposes restrictions on how to interpret new and contradictory evidence such as Ardi.

Arguments over Ardi’s status continue unabated. While writing this review, New Scientist reported on a study in Science Advances that analysed Ardi’s hands, arguing that they are more chimp-like and adapted for swinging from branches (suspensory locomotion). Unsurprisingly, White has responded to this in his usual gruff fashion: “This is another failed resurrection of the antiquated notion that living chimpanzees are good models for our ancestors“.

I admit that I was initially mildly concerned that the publisher was overselling Fossil Men by calling it a scientific detective story. However, it did not take long for me to become completely engrossed by Pattison’s portrayal of these scientists, and to be in awe of the skill with which he tackles numerous complex biological topics. This is a chunky book, but it is a page-turner that you will not regret.

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

Fossil Men

Other recommended books mentioned in this review:

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The history of human evolution has become firmly wedded to the Out of Africa hypothesis: the idea that we evolved in Africa and from there spread around the world. Back in 2015, palaeoanthropologist David R. Begun gave the proverbial tree of life a firm shake with The Real Planet of the Apes, making the case that the picture is a bit more complicated than that. Providing an incredibly well-written overview of the deep evolutionary history of great apes and humans, an interesting picture emerges of species moving into and out of Africa over time. Some reviewers hailed it as provocative—but is it really?

The Real Planet of the Apes: A New Story of Human Origins, written by David R. Begun, published by Princeton University Press in October 2015 (hardback, 262 pages)

In case you are wondering why I am only now reviewing a book published in 2015: I had just started reading the foreword of the recently published Ancient Bones when it occurred to me that its spiel—our deep evolutionary history lies in Europe rather than Africa—sounded familiar, as did the name of its writer, David R. Begun. Where did I read about this before? One glance at my bookshelves confirmed that, yes, Begun wrote The Real Planet of the Apes which I bought some years ago but had not yet read. Better late than never.

With this book, Begun takes the long view, surveying the deep evolutionary past of the ancestors of great apes and humans, i.e. the period from approximately 35 to 7 million years ago (mya). His starting point is the split between the ancestors of New World monkeys (which went off on their own evolutionary experiment in South America), and the ancestors of Old World monkeys and hominoids (humans, lesser and great apes, plus all their ancestors). Begun would some years later contribute a very readable 10-page summary of archaic ape evolution to Gurche’s Lost Anatomies, but the executive summary would be: out of Africa, into Europe, and then back into Africa.

“[…] Begun takes the long view, surveying the deep evolutionary past of the ancestors of great apes and humans”

A bird’s-eye-view version of his argument starts with the great diversity of catarrhine* primates flourishing in Africa from the Oligocene into the early Miocene (~33 to 17 mya), including Aegyptopithecus (a “not-quite-ape”), Kamoyapithecus (“possibly the first ape?”), and the intermediate monkey-ape genera Proconsul and Ekembo. Then, by 17 mya, Afropithecus and the closely related Heliopithecus show up in East Africa and expand their range into Eurasia. While Asia sees its own flourishing of species in what is by now the middle Miocene, Europe is the epicentre of the overall story and sees the evolution of Griphopithecus and Dryopithecus around 15–13 mya. Four descendants of Dryopithecus (Hispanopithecus, Rudapithecus, Ouranopithecus, and Oreopithecus) are discussed here in particular.

Meanwhile, the fossil record in Africa seems to dry up. Are we yet to find crucial fossils from this period? Possibly, but Begun points out we have plenty of other fossil mammals from these areas and the environment back then would have been suitable for apes. There are some questionable African fossils from this period discussed here, but nothing convincing. By ~10 mya, climatic changes forced the Eurasian great apes to reinvade Africa where the first hominins appear by ~7 mya; Sahelantropus tchadensis is mentioned in particular.

Now, in case you think that this makes for a neat linear story, let me stop you there. For the sake of brevity, I have skipped over many side branches and dead ends that Begun discusses here. Evolution resembles a tangle more than a tree. Although his ideas are not accepted by everyone (show me a group of scientists not endlessly arguing finer points), are his ideas really this much at odds with the established Out of Africa story? Let me offer three observations.

“[…] your narrative depends on what period you take as your starting point and what groups you consider, which can make the whole Out of Africa narrative sound a bit arbitrary”

First, definitions matter. Casual use of the phrase “Out of Africa” in the popular press often fails to mention the when. Some researchers use it to talk about Pleistocene migrations into Europe by hominins such as Homo erectus, though it more commonly refers to the spread of anatomically modern humans from Africa between 300,000 to 200,000 years ago. Begun ends his story some 7 mya, which is when many popular accounts start theirs. Although I do not know if Mark Maslin agrees with Begun’s thesis, the scenario laid out in his book The Cradle of Humanity is perfectly congruent with it. Maslin’s starts around 5 mya and Begun agrees that the fossil record situates early hominins in Africa by this time. My thoughts after reading this book are that your narrative depends on what period you take as your starting point and what groups you consider, which can make the whole Out of Africa narrative sound a bit arbitrary. Even Begun’s ideas could be cast as an Out of Africa scenario.

Second, the recent ancient DNA revolution has splendidly revealed how populations have constantly moved and mixed. Though this technology only allows us to look back hundreds of thousands of years, there is no reason to think the pattern differed drastically before that. Especially as pretty much all palaeoanthropologists agree that climatic changes are an important driver of migration: Begun does so here, Maslin does, even the Leakeys—focused as they are on human evolution in Africa—do.

Third, Begun’s claims might be provocative, but his delivery is not. Instead, his account is tempered and reasonable, clearly indicating where and why his ideas diverge. He might not always agree with others but nevertheless goes on record here to praise their track record. The most severe criticism he expresses (and that is putting it strongly) is that he follows a philosophical approach of putting evidence before process (e.g. the Out of Africa hypothesis). In his opinion, others lean towards putting process before evidence, ignoring or downplaying evidence that does not fit. He thinks that most of the research community has mischaracterised the European branch of the ape family during the middle to late Miocene as little more than an exotic sideshow. But he also repeats throughout how he is open to changing his mind with future fossil finds. Basically, I see someone of intellectual integrity here.

“[…] Though [ancient DNA] only allows us to look back hundreds of thousands of years, there is no reason to think [migration patterns] differed drastically before that. Especially as pretty much all palaeoanthropologists agree that climatic changes are an important driver”

And there is much else to like here. Begun clearly, and if necessary repeatedly, clarifies terminology, whether it is cladistics or skeletal morphology. He also offers excellent explanations of how we reconstruct past climates. How the rules around taxonomical nomenclature work. How plate tectonics influenced biogeography, specifically how, as the Tethys Sea was closing up, rising and falling sea levels revealed and sundered land bridges, opening or closing migratory routes in the Mediterranean and Arabian Peninsula. And how, with modern technology (and often a lot of hard work) we can extract unbelievable detail from fossil teeth, which are frequently the only remains we find. Additionally, he spices up his writing with introductions to the typical fauna of the different periods, short excursions into the history of palaeoanthropology as a discipline, and personal anecdotes of old-guard researchers he met or worked with.

Seeing the book’s age, it is relevant to ask what has happened since publication to change or reinforce this idea. When I contacted Begun, he confirmed that there have been no major finds to overthrow his ideas, with new work adding further support. I will next turn to the 12-million-year-old finds recently described in Ancient Bones. Also of interest is the story of the 4.4 million-year-old Ardipithecus, one of the last species Begun mentions, which is told in more detail in the recently published Fossil Men.

I was not all that familiar with the earliest chapters of primate evolution, so, all in all, I found The Real Planet of the Apes an incredibly engaging and useful account.

* catarrhines are a taxonomical group that encompasses humans, great apes, hylobatids such as gibbons, old-world monkeys, and all their ancestors before these various groups split from one another.

The Real Planet of the Apes

Other recommended books mentioned in this review:

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“Skeleton Keys: The Secret Life of Bone“, written by Brian Switek, published in the US by Riverhead Books in March 2019 (hardback, 276 pages), and in the UK by Duckworth in August 2019 with the title “The Secret Life of Bones: Their Origins, Evolution and Fate” (paperback, 288 pages)

Switek starts off with a potted evolutionary history of the skeleton, taking the reader all the way back to the Cambrian, some 455 million years ago. The small fossils of Pikaia gracilens are some of the earliest evidence we have of the starting point of skeletal evolution. Looking for all the world like a small worm, it was one of the first creatures to possess a notochord, a cartilage-like structure that is the precursor of the backbone. (For a more technical exposé of that borderline between vertebrates and invertebrates, see my review of Across the Bridge: Understanding the Origin of the Vertebrates.)

He then hops, skips, and jumps to other significant milestones; fossils of the fish Entelognathus primordialis show the transition to jawed fish, while Tiktaalik roseae (see Your Inner Fish: The Amazing Discovery of Our 375-Million-Year-Old Ancestor) shows the transition of vertebrates moving onto land. He considers protomammals, the cynodonts, whose offspring lived through the age of the dinosaurs, and the primordial primates, noting how the changing skeleton acquired more and more traits we now think of as human.

This part is far from a complete overview of the evolution of skeletons (for that, see e.g. Skeletons: The Frame of Life), but that was never the intention. It does allow Switek to exercise his funny bone, wondering whether without the evolution of jaws the book and movie would instead be known as Pharyngeal Slit, or comparing the earliest invasion of land by plants to a prehistoric salad bar. At the same time, he is keen to correct misunderstandings about evolution: “It is easy to make categorical divisions between humans and apes when extinction has removed your ancestors”. Similarly, as Tiktaalik shows, any true invasion of land did not coincide with the origin of fingers and feet, with fish evolving into amphibians only millions of years later. This should do away with the misconception of evolution being goal-oriented (beyond, you know, making it to the next generation).

“Without the evolution of jaws, would the book and movie instead be known as Pharyngeal Slit?“

But we cannot dwell here any longer. Bone as living tissue is fascinating, and Switek introduces the physiology, with osteoblast cells continuously forming new bone while osteoclasts break it down again. Something that goes off kilter when astronauts spend months in space and lose bone mass. That makes hibernating bears all the more of a miracle, how do they not lose bone mass? Switek has the answer. Bone can be moulded in life, as seen by cultures around the world that change the shape of infants’ skulls, while in death it retains a personal history of disease and injury. Archaeologists are becoming increasingly skilful at elucidating these stories (see for example Injury and Trauma in Bioarchaeology: Interpreting Violence in Past Lives and Skeletons in Our Closet: Revealing Our Past through Bioarchaeology).

At this point Switek transitions seamlessly into the cultural significance of bone, and how especially skulls have become emblems of death. Neanderthals appear to have been much more sophisticated than we have long given them credit for, as evidenced by burials (see my review of The Smart Neanderthal: Cave Art, Bird Catching, and the Cognitive Revolution), while his recounting of the exhumation of Richard III’s skeleton in a Leicester car park in 2012 is a fascinating archaeological detective story.

A far darker chapter that Switek tackles with panache is the heritage of anthropology. The pseudoscience of phrenology (where measurements of the bumps on a skull supposedly predicted someone’s mental capacities and traits) was long used to justify white man’s superiority. As he mentions, anthropology may not have invented racism, but it certainly fueled it through the 19th and 20th century. Switek is deeply troubled by the resurgence of the idea that race is biologically meaningful. As has been documented at length, there is more variation within populations than between populations, and the overlap between what we thought of as races is enormous (see strident takedowns in The Myth of Race: The Troubling Persistence of an Unscientific Idea and Superior: The Return of Race Science).

“[…] anthropology may not have invented racism, but it certainly fueled it through the 19th and 20th century.”

This naturally leads on to the literal skeletons in the closets of many research collections. Lance Grande, a curator at Chicago’s Field Museum, dedicated a chapter to this in his book Curators: Behind the Scenes of Natural History Museums. The repatriation of old skeletons to for example Native American tribes, for whom the bones of the deceased hold particular spiritual significance, is becoming more commonplace. But not all museums are going along with it, as evidenced by Switek’s story of the skeleton of the Irish Giant Charles Byrne, who is still on display. Another one of those eye-opening tidbits in the book is his peek at the online trade in human bones on platforms such as eBay and Etsy (who have since cracked down on it), and now Instagram. You are far better off building your own.

Each chapter opens with a drawing from the 1733 book Osteographia or the Anatomy of Bones. These are lovely, and my only bone of contention (sorry) is that there aren’t more illustrations in the book. I would have loved to see some photos included, as Switek describes many wonderful things. For that, readers will have to turn to, for example, the work of palaeoartist John Gurche (see my review of Lost Anatomies: The Evolution of the Human Form) or, two of my personal favourites, Evolution in Action: Natural History Through Spectacular Skeletons and Skulls: An Exploration of Alan Dudley’s Curious Collection. Other slightly less spectacular but still noteworthy books are The Skeleton Revealed: An Illustrated Tour of the Vertebrates and Skeletons: The Extraordinary Form & Function of Bones.

Skeleton Keys is a multifaceted exploration of bones and their biological and cultural importance that is very absorbing. Far from a macabre gawk-fest (Skeletons! Eek!), Switek capably handles a range of serious topics, smoothly transitioning between them. The narrative sizzles, whether it is with witty jokes or genuine ire at the disrespect to bones and the questionable ideas they are used to prop up. An incredibly enjoyable book that comes highly recommended.

*Note that the book is published in Europe by Duckworth in paperback with a completely different title: The Secret Life of Bones: Their Origins, Evolution and Fate.

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

The Secret Life of Bones paperback / Skeleton Keys hardback

Other recommended books mentioned in this review:

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“The Art of Animal Anatomy: All Life is Here, Dissected and Depicted“, written by David Bainbridge, published in Europe by Herbert Press (a Bloomsbury Publishing imprint) in October 2018 (paperback, 256 pages)

Bainbridge has divided The Art of Animal Anatomy into five chronological chapters and limited himself to depictions of vertebrates. The early records of anatomical artwork are scant, though works have come to us from the Middle East and Asia. In Europe, we rather saw monstrous and fantastical depictions in bestiaries. Bestiary: Animals in Art from the Ice Age to Our Age shines more of a light on this phase of art history. Bainbridge picks Leonardo da Vinci and Albrecht Dürer as his starting point. These polymaths, who, unbeknownst to me, were contemporaries, drew and painted many different subjects , so only a few representative anatomical images are here included (see their complete oeuvres in Leonardo Da Vinci: Complete Paintings and Drawings and Albrecht Dürer, but see also Nature’s Artist: Plants and Animals by Albrecht Dürer).

A specific chapter is reserved for the horse, which was a hugely popular subject for artists, and a wonderful mixture of black-and-white engravings and colour paintings is included. Bainbridge especially highlights the work of the 16th century Italian Carlo Ruini and 18th century Englishman George Stubbs. Skeletons, skulls, legs, viscera – you will see the horse as never before. The next phase Bainbridge highlights is the fascination with the variety of life in works from the seventeenth to the nineteenth century and with it the rise of comparative anatomy. This includes works by the Dutch Volcher Coiter, but also well-known scientists such as George Cuvier, Richard Owen, and many others, showing us (parts of) birds, snakes, turtles, primates, and many other animals. At this point in time, scientists were particularly struck by the many similarities on the inside of animals, and their drawings compare animal structures to show how skeletons and organs of vertebrates hint at a shared origin.

By the 19th century, description and classification were starting to give way to mechanism and process, not least because of the impact of the theory of natural selection. In his book Across the Bridge: Understanding the Origin of the Vertebrates, Henry Gee pointed out how evolution and embryology became intertwined as disciplines, which is largely thanks to the prolific artist and scientist Ernst Haeckel (see his oeuvre in The Art and Science of Ernst Haeckel). He became a strident proponent of the theory of natural selection, and wedded it to his embryological work, confidently stating that ontogeny recapitulates phylogeny (i.e. development re-enacts evolutionary history). The full story of his notorious drawings comparing the development of vertebrate embryos has been told in full elsewhere (see Haeckel’s Embryos: Images, Evolution, and Fraud, reviewed here).

Another important development was the acceptance of deep time and a more accurate determination of the Earth’s age, and with it the rise of palaeontology as a discipline, resulting in artwork depicting fossils. Some early drawings of fossils and skeletal reconstructions from the hands of Mary Anning and Richard Owen are shown, and mention is made of the Crystal Palace dinosaurs, but there is no space here to explore the discipline of palaeoart (for that, see Paleoart: Visions of the Prehistoric Past, 1830-1980).

“Skeletons, skulls, legs, viscera – you will see the horse as never before”

Other important figures highlighted are D’Arcy Wentworth Thompson, who placed physical and mathematical principles above all else and drew diagrams geometrically distorting animal shapes to show relationships (see his book On Growth and Form). The work of Edweard Muybridge, who first experimented with stop-motion photography, is shortly covered, including his famous sequence of a galloping horse (for more, see Eadweard Muybridge: The Human and Animal Locomotion Photographs). But also the work of the less well-known Spaniard Santiago Rámon y Cajal is highlighted. He pretty much single-handedly transformed the discipline of neuroscience and his drawings remain relevant to this day.

A final brief chapter shows some contemporary examples of anatomical artwork, whether this is outdoor graffiti, the plastinated anatomical specimens of Gunther von Hagens, or the remarkable anatomical body-paint applied to horses by Gillian Higgins from Horses Inside Out (see, amongst others, Horse Anatomy for Performance: A Practical Guide to Training, Riding and Horse Care). Not included, but worth pointing out, is the contemporary anatomical artwork of Katrina van Grouw (see her beautiful The Unfeathered Bird and Unnatural Selection) who shows there is still mileage in artwork that harks back to centuries past.

Bainbridge has done an admirable job compressing a lot of history in a relatively small book, showing how illustrations became more and more informed by science, and served as educational tools for anatomists and veterinarians alike. He highlights particularly noteworthy artists in each chapter and provides short commentaries with all artwork. Given the price point of this book, don’t expect lavish production values such as those of German publisher Taschen, or shenanigans such as fold-out plates. Nevertheless, the artwork itself is wonderfully reproduced on matte paper in numerous half- and full-page images, as well as full-page spreads. Some of these span one-and-a-half pages, ensuring the centre of attention is preserved and visible, whereas others span a full two pages which sometimes means important details disappear in the fold of the pages. Luckily these cases are few. In Europe the book is only available as a paperback, but if you’re on the other side of the Atlantic pond, Princeton University Press bought the rights for the US market and has published it in hardback with a slightly different title, Stripped Bare: The Art of Animal Anatomy – a format which I would have preferred.

The Art of Animal Anatomy is a wonderful visual trip that, far from being gory or gruesome, contains fantastic imagery spanning a wide span of time. It contains many works well-known to most biologists, but will undoubtedly introduce you to new and unknown artworks. I was particularly taken with the paintings of superficial muscles from the hand of J.E.V. Boas, for instance. Though not intended as a handbook (for that, see books such as Animal Anatomy for Artists: The Elements of Form, Bird Anatomy for Artists, or The Palaeoartist’s Handbook: Recreating Prehistoric Animals in Art) this book will provide plenty of inspiration for artists. And it makes a sure-fire gift for that person in your life who loves the interface of biology and art.

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

The Art of Animal Anatomy paperback or hardback

Other recommended books mentioned in this review:

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