Tribute to Richard Fortey and his trilobites

Trilobite expert Richard Fortey (15 February 1946 – 7 March 2025) brought the prehistoric world to life through an unprecedented literary oeuvre. His eloquently written books – on palaeontology and far beyond – are interwoven with wonder and a deep passion for nature. Trilobites, a fossil class of arthropods, lay at the heart of his focus. Even in their gaze, now hardened into stone, he could breathe sparks of life. Those remarkable kaleidoscopic eyes were among the first to see the ocean, to witness long-lost underwater worlds, which Richard Fortey described with both zest and warmth. Through his literary legacy, he opened many eyes in many ways. Mine too.

By Kathelijne Bonne.

When I turned the last page in his book Trilobite! in March this year, I was shocked to learn that Richard Fortey had just died. He had personally recommended this book to me. It is a deep dive into wonderful lost worlds, brimming with unusual views of life, and interspersed with subtle British humor – so elusive to non-Brits (and a true test for translators). I find some solace in knowing AI will never be able to produce anything like this book.

I do not know how unexpected his passing was, but the news caught me by surprise as his last e-mail showed that he was bubbling with activity. In denial of the frailty of life, I thought he still had decades ahead, and would at least live to become an unstoppable nonagenarian like my other heroes Jane Goodall, Sylvia Earle and David Attenborough, globetrotting and churning out book after book. 'Contingency' however, had other plans. 

But Richard Fortey did not leave without a legacy, one in which he infused science with magic and mystery, things many had forgotten were ever part of it. Science and discovery even grant the practitioner a touch of immortality, he wrote, the very thing humanity has sought since time immemorial:

It may be the access to a kind of immortality, of however unusual a variety, which make science such an attractive option for intelligent people seeking meaning in their lives. ... And he continues, showing his usual wit: Death cannot be cheated, but the discoveries made in one's prime may well outlive bodily decay.

And every contribution counts, no matter how small:

The beauty of a life devoted to science is that every honest practitioner contributes an indelible stone to the great edifice of knowledge. Perhaps among their intellectual successors there will be a few who remember them, but what really counts is their contribution, even if it is anonymous.

Charisma, chimera, and a trident for jousting

Trilobites are a fossil class of arthropods that branched into a vast diversity of species during the Paleozoic era. They spread throughout the farthest corners of ancient oceans. They occupied all ecological niches. The Paleozoic can rightly be called the era of trilobites, even though Richard acknowledged the importance of other life forms too. 

Thanks to Richard Fortey, certain trilobite behaviours, evolutions and lifestyles came into sharper focus. Since he first looked a trilobite in the eye as a 14-year-old after a tap of the hammer, he discovered many. After his appointment at the Natural History Museum in London, he also became authorised to give them names. Through his literature, they gained not only fame and a wider appreciation, but also charisma, personality and vivacity.

The earliest recognised case of chivalry came to my attention directly through Richard, who forwarded me an article (see bibliography) on the incredible Walliserops trifuncatus, a trilobite with a trident. He wrote in Trilobite! about this fairytale-like creature: The trident bearer was a dream, a chimera, that should not exist. Yet it did. And ignoring the fact that this was just a remarkable find, he hoped and knew that the trident bearing trilobite may one day stimulate a moment of wonder in a child.

The trident enabled the trilobite to ward off other males via jousting to court the females (a behaviour known as intraspecific combat or sexual combat). Heroic (or macho?) behaviour is at least 400 million years old and probably older.

Enrolled into a ball, legs of gold, and the Pompeii of trilobites

Among more endearing trilobite antics is the tendency to enroll into a ball when in danger, reminiscent of hedgehogs, armadillos and pangolins and some insects such as woodlice. Many trilobites died enrolled. They were startled by an approaching threat and did not survive the attack; giving us a glimpse into a snapshot in the life of an animal that died hundreds of millions of years ago.

As their name suggests, trilobites consist of three lobes (tagmas), a head shield (cephalon), body (thorax) and tail shield (pygidium). And also in transverse section, they show a tripartite structure with a central axis and lobes on both flanks. Their body is divided into segments and to each segment an articulated leg was attached – typical of all arthropods, including insects, spiders and crustaceans – the most diverse group of animals on Earth. The hard armour of their calcified exoskeleton favoured fossilisation. In some cases, they are extremely well preserved. Every decorative tip, protrusion or notch has been immortalised in microscopic detail.

In contrast, soft tissue such as antennae, gills, muscles, organs and legs fell prey to decomposition. But in exceptional circumstances, even these were preserved (for example in lagerstätten such as the Burgess Shale), or in low-oxygen environments. In the latter, the delicate leg tissue was replaced during fossilisation by pyrite, fool's gold, giving the legs a golden tinge.

And another level of anatomical details were revealed more recently, in the Pompeii of trilobites, an excavation in Morocco in which trilobites were killed in a pyroclastic flow.

Eye-to-eye with trilobites

Trilobites lived exclusively in the sea, where they played roles of predators and prey, from tiny pelagic species that flitted through sea currents in schools like tiny fish, to larger fearsome hunters, filtering sea-bottom dwellers and scavengers, and herds of grazing species, almost cow-like.

Evolution moulded their compound eyes into the strangest shapes, depending on what needed to be seen. Some trilobites had eyes on stalks, like snails; others had no eyes at all because they lived in the midnight zone. The eyes had been filtered out by evolution. Still other pairs of eyes looked only in a two-dimensional plane, apparently enough to catch sight of opponents.

The eyes consisted of lenses composed of calcite. Each eye contained one to thousands of lenses. Most trilobite eyes were holochroal, with lenses set in a dense, hexagonal close-packed arrangement. Only one suborder of trilobites (Phacopina) had schizochroal eyes. However they perceived the world, trilobites are invaluable for deciphering the evolution of the eye, which dates back to the Precambrian. The eye is possibly a billion years old.

Along with other scientists such as the late Euan Clarkson, Richard Fortey shed light on how exactly trilobites saw, and how much more there is to learn, that higher goal of science: "I can imagine that we will see more clearly how trilobites saw. Their eyes will gaze upon vanished worlds with a pristine clarity."

Eyewitness, Tornquist Ocean and fossils vs machines

The early evolution of trilobites is still shrouded in mystery, but Richard Fortey shone a light on how they evolved and how they were eyewitness to evolution throughout the Paleozoic, from the Cambrian until they disappeared forever in the Permian mass extinction.

Research on trilobites improved understanding of certain aspects of evolution, such as punctuated equilibrium, and they even provided evidence for plate tectonics. The distribution of different trilobite faunas allowed the identification of an ancient ocean, the Tornquist Ocean of the early Paleozoic, a name chosen by Richard Fortey. 

This ocean of the Palaeozoic unleashed a battle between palaeontologists and geophysicists. The latter study palaeomagnetism to reconstruct movements of tectonic plates. They denied the existence of the Tornquist Ocean because they found no evidence in their data, and bluntly told the palaeontologists that "one palaeomagnetic data point is worth a thousand trilobites". But trilobites, including a species called Merlinia, do not lie, and this 'fossils versus the machines' clash was settled in favour of the former.

Cambrian Explosion?

Richard nuanced the concept of the Cambrian Explosion of life, which I dwelled on in the article on supercontinent Gondwana. From about 540 million years ago, life would have suddenly and massively developed, 'exploded', in many directions. But Richard had reservations about the 'explosiveness'. He believed that the transition from the Precambrian to the teeming, life-seething Cambrian was not so spectacular and explosive after all. His ideas are worth a separate article or book.

There is much more to say and discover about this crucial period, the possibilities are wide open, a barrel brimming with worlds and dreams yet to be shaped. I cannot thank Richard enough for opening the doors to those worlds for me, a quarter of a century ago.

Amphitheatre, horizons, Matterhorn

When I was 17, the thought of ​​studying geology took shape. I had finished reading the book I had been given for my birthday, Life: An Unauthorised Biography of Four Billion Years of Life on Earth. A world had opened. Curtains were drawn aside and an amphitheatre came into focus. Suddenly I saw everything in multiple dimensions, including that of time – life does not take place in the flat plane of the present but in a kind of layered, immeasurably deep multiverse in which every contemporary event is created by a network of millions of small happenstances that connect the past to the present via invisible links. Thanks to that seminal book, I began to suspect the existence of these worlds, and now I see them clearly, although I know that I only see a fraction. That is the exciting thing about science, it never stops, we never know enough and we wouldn't want to know enough. Behind every horizon lies another horizon.

I take pleasure in the fact that the ground under the place where I am writing this article (central Spain) consists of granite that solidified as a large batholith during the Carboniferous when this country was in the middle of Pangea, the last in the majestic cycle of supercontinents. High mountain peaks and probably volcanoes too, towered kilometres above my head, only to be worn away by mega-anni of erosion, ground to sediment, now part of the seabed and other rock layers, washed away and deposited elsewhere, "like discarded chiffon". Wolves and mouflons now roam the higher hillslopes of this granite landscape. Vultures glide high in the sky. Blue-tailed Iberian magpies are causing a racket outside my window. And when this familiar granite, now speckled with cork oaks, was still molten magma, the seas around Pangea teemed with trilobites.

Every book by Richard Fortey takes you on a grand sweep through multidimensional worlds. In "Earth" I was struck by the chapter on the formation of the Alps. How that mountain range consists of several 'nappes' pushed over each other like a wrinkled tablecloth. The whole mountain range is seen differently when you know that the pyramidal peak of the Matterhorn is a fragment of Gondwana and that the Dolomites were in the Tethys Sea.

And while our human world is polarized by leaders who bend the rules of the chess game, using peoples as pawns in their schemes, my gaze remains focussed on the nature that surrounds us and the beauty and pleasure that can be found there. The way Richard Fortey described nature is unsurpassed. Perhaps his greatest contribution is to express how science contributes to happiness and how it brings out the best in people:

"Yet I know that there can be nothing better than to pursue such dreams [of possibility]; that the will to know the truth is one of the better parts of human nature; and that trilobites will reward the investigator in a currency more valuable than dollars, and more tangible than fame."

Rest in peace, Richard. 

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Also inspiring: marine algae in cyanotype, Ocean Planet, total solar eclipses, amateur astronomy, and trees of heaven.

In a uniquely British way, Jane Goodall also wrote many sparkling books. A Spiritual Journey and In the Shadow of Man stayed with me most. I wrote two articles featuring Jane: one about how I met her, and another on the animals we love and eat. Life on Air by David Attenborough is written with equal fluidity. An American hero is Rachel Carson. Like Richard Fortey, she was born a writer, but it was marine biology that gave her something to write about—and she did so until the end of her life. Silent Spring is her most famous work, but The Sea Around Us may be the most beautiful.

Kathelijne: I am intrigued by how earth, life, air, ocean and societies interact on geological and human timescales.

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