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Friday, 31 July 2020

Benjamin Waterhouse Hawkins: the first grand master of palaeoart

Few of us need an introduction to the Crystal Palace dinosaur sculptures (here shown during construction in 1854), or their artist, Benjamin Waterhouse Hawkins, but not many of us know much about Hawkins or his work beyond this one installation, thus overlooking a major contributor and pioneer in the early history of palaeoart. Image from the Friends of Crystal Palace Dinosaurs blog.

Most of us would agree that palaeoart history is marked by a few key figures that, for one reason or another, define their generation. Knight, Burian, and Paul are perhaps the most considered of these, each producing an iconic portfolio of work that set the bar for other artists and recast how we might view and consume palaeoartworks. But the professional grandfather to all these famous figures is Benjamin Waterhouse Hawkins, a familiar name to anyone interested in palaeoart or vertebrate palaeontology, and yet one that doesn't always receive the accolade and praise it perhaps deserves. Hawkins - whose life spanned 1807-1894 - became the first great palaeoartist at a time when both palaeoart and palaeontology was still finding their feet. We all know of Hawkins' work at Crystal Palace, where he built 33 life-sized restorations of prehistoric mammals, reptiles and amphibians as part of the grand educational and commercial extravaganza, but many of us do not know much about him, his other works, or even how groundbreaking and unique his Crystal Palace sculptures were, demonstrating the sort of good practise and insight that we regard as essential in modern palaeoart.

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Hawkins was a skilled anatomist who wrote several books on animal anatomy and form. These images, from his 1860 book A comparative view of the human and animal frame, show his mastery of osteology and anticipate the now-common convention of a dark soft-tissue silhouette behind our skeletal reconstructions.

An appreciation of Hawkins is helped by learning something of his life. Although featuring frequently in accounts of Victorian-age natural history, the details of Hawkins' life had not been pieced together in real detail until his great great, great-granddaughter Valerie Bramwell and historian Robert M. Peck synthesised his personal and professional records into a concise biography (Bramwell and Peck 2008). Hawkins' existence was an eventful one, including both great professional success and tragedy, as well as a complex and somewhat mysterious personal life. The latter is perhaps the most cryptic and unexpected part of the Hawkins story, as he somehow sustained nearly four decades of bigamy (juggling two wives and 10 children, seven of whom survived infancy) without his families discovering the deception. His strange domestic life was found out in the mid-1870s, not long before a string of personal tragedies and a stroke brought Hawkins' life to a sad, inauspicious end. But despite his bizarre private dealings, Hawkins seems to have been a well-liked, generous man known as hardworking, personable and charming, equally confident of his abilities as an artist and anatomist but also reverential and respectful to his peers, especially the academics and intellectuals he frequently worked with.

Hawkins never secured permanent or long-lasting employment but instead moved from commission to commission and project to project. Though he dabbled in a few other careers, his chief profession was creating art of animals - initially living, and eventually extinct. Known best today for his sculpture, Hawkins only began working in 3D during the 1840s, well after he had established himself as an expert painter and illustrator. He was regarded as a high-quality, experienced natural history artist early in his career such that, even before he reached his career peak - the Crystal Palace geological restorations - he was sought-after as an illustrator of zoological specimens. He produced art for many of the biggest names in contemporary palaeontology and biology, including William Buckland, Gideon Mantell, Richard Owen, Thomas Henry Huxley, Charles Darwin and Joseph Leidy (interestingly, Hawkins could have worked with another big name, Edward Cope, but he disliked and refused to work with him - see Desmond 1976). Hawkins' association with Darwin and Huxley is of interest because he became an outspoken anti-evolutionist in later life, a position he expressed directly and clearly in many of the popular public lectures he delivered in his post-Crystal Palace career.

Before Hawkins recreated extinct dinosaurs, he was a prolific and sought-after illustrator of the modern kind, along with many other types of animals. This illustration of a White-crested Kalij was produced for John Edward Gray's Illustrations of Indian Zoology (1830-1835). (From Bonhams; incidentally, original Hawkins' lithographs like this sell for hundreds or even thousands of pounds to private collectors, a sad contrast with the under-financed conservation of his Crystal Palace works.)

Hawkins was well-known for his expertise in animal anatomy and form, which he learned primarily from zoological museum specimens as well as drawing captive animals (Bramwell and Peck (2008) recount an urgent trip to Paris in 1849 to sketch a newly born giraffe calf). As would happen later with Charles Knight, this expertise with living animals was instrumental in his later work at reconstructing extinct ones. Hawkins authored several books on animal anatomy and from the 1850s onward spent a large amount of time - and commanded a high fee - for grand public lectures. At his lecturing acme, Hawkins produced large drawings of prehistoric animals on stage (of such size that a ladder was required to reach the top of the board canvas) to bring his restorations direct to his audience. A gleaming reputation among British academics meant he enjoyed a warm reception in the US where he was offered many auspicious platforms and opportunities by well-regarded officials and academic institutions. These included the production of (sadly never realised) Crystal Palace-like prehistoric animals installations in Central Park and the Smithsonian, and production of the first-ever mounted dinosaur skeleton (Hadrosaurus), of which several casts were made (of which only one skull survives).

Hawkins' Central Park Workshop, c. 1869, with an assembled Hadrosaurus mount and the beginnings of a Laelaps skeletal reconstruction alongside - note the mount outline to the right of the Hadrosaurus ;skeleton. This terrific image (reproduced from Bramwell and Peck 2008) is a treasure trove of detail: look out for various mouldings of reptile skin, including a carnivorous species on the left, real animal skeletons in the background and unmounted bone replicas on the floor (are these bits of Laelaps?). There's also a lack of vermin - Hawkins' Crystal Palace workshop was shown with rats and birds scampering over the floor, and described as lying among a muddy swamp. The content of this workshop was destroyed by order of corrupt New York politicians, which devasted Hawkins.

There's a lot more we could say about Hawkins career, but you get the point: Hawkins wasn't a flash-in-the-pan artist who happened to land the Crystal Palace gig, nor was he someone who just knew how to throw clay and concrete around to someone else's design. He was a distinguished and respected intellect with expertise in natural history and anatomy, and brought considerable experience to his palaeoart commissions. Indeed, historians are uncovering increasing evidence that Hawkins was the principle intellect behind many of the Crystal Palace sculptures. We touched on this in my previous discussions of the Crystal Palace palaeoartworks but it's worth repeating: Richard Owen, lauded by many (e.g. Phillips 1854; Owen 1894; Desmond 1979; Beaver 1986; McCarthy and Gilbert 1994) as the brains and overseer of the Crystal Palace models, was actually barely involved and may have even held disdain for this grand project (Secord 2004; Dawson 2016). Numerous pieces of evidence, the best of which are the private correspondence from Hawkins, Crystal Palace officials and others, show that Owen only visited the models once during their construction for the famous 1853 New Years's Eve banquet held in the clay Iguanodon. Most tellingly, Hawkins is on record as stating that Owen "afforded no assistance" while the models were being built (Dawson 2016). Owen later attempted - as was his custom - to take greater credit for his role in the project, but his actual contributions seem restricted to input on some early clay maquettes constructed by Hawkins; a short, incomplete guide book about the sculptures (Owen 1854); some promotional duties; and allowing his name to be used to give the restorations scientific authenticity. We can thus draw two conclusions: 1) having been largely abandoned by his consultant, Hawkins' expertise and anatomical confidence were probably instrumental in seeing the models realised and completed on time; and 2) the sculptures are not - as so often stated - Owenian theory brought to life, but Hawkins' personal take on the works of Owen, Mantell and Cuvier. This probably explains why, as we discussed previously, many details of the models are at odds with Owen's ideas, as well as the nitpicking tone his 1854 sculpture guidebook.

File:Benjamin Waterhouse Hawkins Moas of Prehistoric New Zealand ...
Hawkins' 1870s take on the moa, produced for College of New Jersey (now Princeton University). The moa is not a creature we intimately associate with Hawkins, but its research history - where its form was predicted from a small amount of fossil material - underpinned Hawkins career as a palaeoartist. From Wikipedia.

It's in this largely unguided context that we have to view many of the details in Hawkins' Crystal Palace work as the product of a genuine palaeoart master, and especially so given their incorporation of an important but ultimately flawed 19th century way of interpreting extinct animals. Hawkins was a student of the then-in vogue Cuvierian philosophy of anatomical correlation: that is, the idea that whole animals could be reconstructed with some degree of precision from very few anatomical remains. This concept, explored in depth by Gowan Dawson in his 2016 book Show Me the Bone, gained traction after successful demonstrations in the early 1800s by Cuvier and then, most famously, by Owen's seemingly miraculous reconstruction of the moa from a femoral fragment. Anatomical correlation became the principle philosophy guiding the earliest visualisations of all poorly known extinct creatures and became celebrated as a scientific marvel in popular and academic spheres alike. Although some scholars - including Mantell - developed misgivings about anatomical correlation by the time the Crystal Palace project began in August 1852, Hawkins relied on this technique to restore his dinosaurs as well as Dicynodon, Mosasaurus and Labyrinthodon, none of which were known from more than scraps of bone at the time. In lieu of a reliable consultant, such predictive restorations might have been disastrous, had they ever been achieved at all, but Hawkins' anatomical expertise instead saw the creation of genuinely lifelike, plausible-looking restorations. Moreover, as discussed in my previous posts, his restorations of Crystal Palace are often more precedent than they first appear, anticipating not only many genuine aspects of prehistoric animal anatomy but also demonstrating sound, logical palaeoart approaches that we can approve of today.

The buffalo-like shoulder hump of Hawkins' Megalosaurus has a slightly complex backstory. Sometimes logically assumed to reference the tall-spined vertebrae of Altispinax dunkeri (a species once part of the Megalosaurus taxonomic complex), it was actually a Hawkinsian speculation that large-skulled dinosaurs needed vast neck and shoulders musculature to support their heads. Initial interpretations of Altispinax seemingly proved Hawkins right, but, of course, we've since realised that the Altispinax vertebrae belong much further down the body, possibly representing a Concavenator-like sail.

Using the dinosaur sculptures as an example of this, it's evident that Hawkins was anticipating the sort of anatomy that giant, somewhat mammal-like giant reptiles might have had to support their vast frames, and was moving well beyond the brief lifestyle and functional commentary provided by Owen and Mantell. He predicted, for instance, that the large heads of dinosaurs would need additional support from their shoulder regions, leading to the portrayal of both Megalosaurus and Iguanodon with shoulder humps of varying size (this is especially obvious in the Megalosaurus, but a shoulder prominence is also discernable in the reclined Iguanodon, while the shoulders of the standing animal bulge into a voluminous, rhino-like neck). This was an entirely sensible prediction that mirrors how we approach reconstructing fossil animals today: when we see robust and large osteological features, we assume they have correspondingly developed soft tissues for support and motion. In this respect Hawkins was ahead of some later artists who essentially ignored aspects of functional morphology and biomechanics in their work, leading to emaciated, peculiarly proportioned animals, such as the emaciated Stout and Kish dinosaur reconstructions from the 1970s and 1980s. Moreover, Hawkins was proved correct in assuming dinosaurs with large heads needed additional neck support, once (as it would turn out, erroneously) during his lifetime when Owen referred the tall-spined dorsal vertebrae of Altispinax dunkeri to the shoulder of Megalosaurus, specifically commenting on their use to support the head (Owen 1856) and, later, by the genuine shoulder and skull anatomy of ceratopsids and certain theropods, which bear augmented anatomy related to skull support.

Hawkins also predicted that dinosaur torsos may differ markedly depending on diet and habits, moreso than they do in conventional reptiles. The herbivorous Iguanodon has a vast belly that spreads in the reclined sculpture, simulating the weight of a large animal resting on a voluminous gut. In contrast, the Megalosaurus has a taut, narrow torso without a significant gut region, more in line with that of a carnivore. At this point in history virtually nothing concrete was known about dinosaur toro dimensions, so these reconstruction choices - also borne out by later discoveries - were sensible predictions of the functional properties of dinosaur guts. A third example concerns his dinosaur musculature, which was modelled after a mammalian, rather than lizard-like fashion. Though mammals are not the best reference for dinosaur myology, Hawkins was right in assuming that large, powerful animals needed large, powerful limbs, and realised that mammals were a more appropriate model than modern reptiles. In this regard, Hawkins was more insightful than some of his successors, who would restore dinosaurs with skinny, lizard-like limbs ill-suited to their (by then well-known and obvious) limb girdles (see artwork by Cope, Knight, Smit etc.). He also went so far as to add features in his art that we associate with especially thorough extinct animal reconstructions today, such as skin webs linking limbs and body, tissue deformation, pose-based muscle bulges and so on.

It's been said before, but it's worth saying again: the reclining Crystal Palace Iguanodon is an amazing work of animal reconstruction. This 30-tonne concrete and brick model captures minutiae of superficial musculature and other nuanced features of anatomy. The world had not seen anything like this in 1854, and even today it knocks the stuffing out of the detailing seen on many dinosaur sculptures.

These points demonstrate how excellent Hawkins was at solving anatomical 'problems' thrown up by fossil remains with an appropriate and sensible corresponding morphology. They belie an experience and instinct with anatomy and biological functionality that remains essential to high-quality palaeoart today, and this was undoubtedly a major factor in how these largely speculative creatures look so realistic. Hawkins' predictions were not always correct of course, but his decisions were logical given the material available to him - the best we can aim for in any palaeoartwork, regardless of its vintage. Needless to say, plenty of people were amazed by his work, and both its popularity and seeming scientific authenticity saw it referenced and replicated by artists for decades to come.

Because Hawkins is mainly known for the Crystal Palace models few people realise that he enjoyed a significant post-Crystal Palace career revolving around reconstructing extinct life. His skills were sought out by various bodies for use in museums, posters and other artworks. During lengthy visits to the United States, he almost re-realised the grandeur of the Crystal Palace models (first in Central Park, later the Smithsonian), but circumstances were never on his side and these projects never materialised or - in the case of the Central Park Paleozoic museum - were sabotaged. Hawkins continued to produce palaeoart until at least the 1870s where he completed what could be considered as his second most substantial set of palaeoartworks: 17 paintings of different geological periods for the College of New Jersey (now Princeton University). These artworks capture the evolution of Hawkins' personal ideas on prehistoric life and are also worth tracking down as exemplary takes on prehistory from the late 19th century. Among the most interesting is the 1877 painting Jurassic Life of Europe where Hawkins revisited his Crystal Palace subjects for (so far as I know) the final time. Hawkins got a lot of professional mileage from the Crystal Palace project and many of his subsequent artworks referenced his sculptures in detail, down to their posing, but by 1877 he must have realised that many of his reconstruction choices were no longer tenable. Both the now-hornless Iguanodon and Megalosaurus bear visibly short forelimbs, long, bird-like hindlimbs, and relatively slender necks with somewhat smaller, less bulky heads. Although still quadrupedal, these restorations indicate the influence of new dinosaur discoveries on Hawkins' classic dinosaur interpretations. This 1877 work might be criticised for his animals not having attained the status of true bipeds, as Hawkins realised was appropriate for both Hadrosaurus and "Laelaps", but consider that Megalosaurus was still considered to have powerful, bulky shoulders at this time (a holdover from Owen's interpretations) and that, while Iguanodon was considered by Mantell (1848) to have relatively gracile limbs capable of non-supportive functions (e.g. grasping vegetation), an explicit case for Iguanodon bipedality had not yet been made - the famous Bernissart Iguanodon skeletons would not be found until a year after Hawkins completed his painting, in 1878. Jurassic Life of Europe is surely among the last palaeoartworks where overtly Mantellian and Owenian interpretations of dinosaurs could be said to have some validity.

Hawkins' 1877 painting Jurassic Life of Europe, produced for the College of New Jersey (from Princeton University Art Museum).

Jurassic Life of Europe and its sister paintings demonstrate Hawkins' adaptability and humility as a palaeoartist. Much of Hawkins fame came from the Crystal Palace Company promoting their dinosaurs as authentic, wholly accurate creations, so their adaption to fit more modern ideals was an admission that his most revered work was not without some significant errors, and that the predictive principle at the heart of the geological court was not infallible. Concerns that the Crystal Palace sculptures were flawed were not new at this point (Dawson 2016) but, by the late 19th century, they were subject to increasing vicious comments from detractors. Othniel Marsh, speaking in 1895, remarked
The dinosaurs seem... to have suffered much from both their enemies and their friends. Many of them were destroyed and dismembered long ago by their natural enemies, but, more recently, their friends have done them a further injustice by putting together their scattered remains, and restoring them to supposed lifelike forms... So far as I can judge, there is nothing like unto them in the heavens, or on the earth, or in the waters under the earth. We now know from good evidence that both Megalosaurus and Iguanodon were bipedal, and to represent them as creeping, expect in their extreme youth, would be almost as incongruous as to do this by the genus Homo.
Othniel C. Marsh, 1895 (quoted in Desmond 1976)

Marsh and other late 19th century critics were, of course, not being entirely fair to Hawkins by comparing his work to their contemporary knowledge of prehistory, and not judging it by the standards of decades prior. Perhaps, for those living through the rapid paleontological advancements of the late 19th century, Hawkins' attempts to reconstruct fossil animals from fragmentary remains, and the genuine belief that they were credible takes on their true form, seemed premature, arrogant and foolish. There may be some truth to this: the 19th century acceptance of anatomical correlation is surely a major case of Dunning-Kruger effect, where scholars had yet to be humbled by the bewildering anatomical diversity present in Deep Time and believed their mastery of modern natural history equipped them to make bold, confident predictions about the past. But it's surely also the case that, while the science of his time was flawed, Hawkins' work was as true to that science as could realistically be expected. Like the kids say, don't hate the player.

Mark Witton on Twitter: "Yes, they seem to linger on in mainland ...
The final extinction of Hawkinsian dinosaurs: the frontispiece to Camille Flammarion's Le monde avant la création de l'homme& (1886). In this (possibly) last use of Hawkins' restorations in a non-historic, non-ironic context, a Crystal Palace Iguanodon (right) meets an early interpretation of Stegosaurus (left) - a representation of New World dinosaurs coming into focus. By this time, new European fossils had already made Hawkins' reconstructions long in the tooth, and American specimens were showing how wide of the mark his predictions were.

Hawkins' death in 1894 occurred without note, despite his significant contributions to science, public outreach, and education. Thankfully, his legacy has not gone neglected and increasing research into his life and work has seen his stock as an artist and intellect grow considerably, at least among academics. But it remains the case that showcasing Hawkins' work to even palaeoart fans sees it quickly dismissed, as by Marsh, as inaccurate, and thus now worthless, takes on prehistory. It's surely the case, however, that drawing attention to the inaccuracies of Hawkins' palaeoart is the most superficial and least interesting observation one can make of it, and defies the obvious fact that palaeoart of any kind must be evaluated in an appropriate historic context. Of course Hawkins' work is inaccurate, and pointing it out does not make one look smart or insightful. Rather, look closely and we see the work of someone who, with no prior experience in the prehistoric animal restoration, took the early palaeaort into a different league of quality and popularity; pioneered principles of soft-tissue restoration and attention to detail that we consider essential today; and demonstrated that even the most unusual extinct animals, with sufficient knowledge of living species, could be restored to plausible, life-like forms. The outdated science underlying Hawkins' work may not it the most accessible of vintage palaeoart, but it's quality, the knowledge that informed it, and historic significance leaves little doubt that Hawkins was a true master of the genre, and worthy of deeper study and appreciation by palaeoart enthusiasts.


In June 2020 the face of the Crystal Palace Megalosaurus, surely one of the most iconic works of Victorian palaeoart on the planet, was significantly damaged. The survival of these works, and Hawkins' legacy, is a continued fight against the elements and the thoughtless people who routinely clamber over them. Image from the Friends of Crystal Palace Dinosaurs website.

Having just written about Hawkins at length, it would be remiss not to mention the terrible damage that recently occurred to the iconic Crystal Palace Megalosaurus, where the front of the face was broken off through suspected vandalism. The Friends of Crystal Palace Dinosaurs charity is now working to repair the damage with Historic England, Bromley Council and specialist conservators, as well as to address issues with the security of the island that houses most of Hawkins' sculptures. Note that this incident has nothing to do with the new bridge to the island, which has not yet been installed and, in any case, will be a rotating structure inaccessible to the public most of the time. Instead, the chief security issue is the low water level around the island, making it easy to access. As you can imagine, fixing this damage and ensuring the security of the Crystal Palace models is an expensive undertaking, so please support the charity if you can. If you really think Hawkins' work is worth preserving long-term, you can set up a recurring monthly donation - a small amount each month can quickly build into a substantial contribution, and gives the FCPD more reliable income over time.


  • Beaver, P. (1986). The crystal palace: A portrait of Victorian enterprise. Phillimore & Company.
  • Bramwell, V., & Peck, R. M. (2008). All in the bones: a biography of Benjamin Waterhouse Hawkins. Academy of Natural Sciences.
  • Dawson, G. (2016). Show me the bone: Reconstructing prehistoric monsters in nineteenth-century britain and America. University of Chicago Press.
  • Desmond, A. J. (1976). The hot-blooded dinosaurs: a revolution in palaeontology. Dial Press.
  • Mantell, G. A. (1848). XIII. On the structure of the jaws and teeth of the Iguanodon. Philosophical Transactions of the Royal Society of London, (138), 183-202.
  • McCarthy, S., & Gilbert, M. (1994). The Crystal Palace dinosaurs: The story of the world's first prehistoric sculptures. Crystal Palace Foundation.
  • Owen, R. (1854). Geology and inhabitants of the ancient world (Vol. 8). Crystal palace library.
  • Owen, R. (1856). The fossil Reptilia of the Wealden Formations. Part III, Megalosaurus bucklandi. Palaeontographical Society. Monographs, 9, 1-26.
  • Owen, R. (1894). The Life of Richard Owen by His Grandson the Rev. Richard Owen,... with the Scientific Portions: An Essay on Owen's Position in Anatomical Science by the Right. John Murray.
  • Phillips, S. (1854). Guide to the Crystal Palace and park. Crystal Palace Library.
  • Secord, J. A. (2004). Monsters at the crystal palace. In: de Chadarevian, S, & Hopwood, N. (eds). Models: the third dimension of science, Stanford University Press. 138-69.

Monday, 8 June 2020

Racism in palaeoart and #BlackLivesMatter

The recent Black Lives Matter protests resulting from the cruel murder of George Floyd at the hands of US police officers have once again drawn attention to matters of racial inequality around the world. These events have been, for me, a belated eye-opener to the depth of institutionalised racism experienced by non-whites internationally, especially by black people. Like many white folks, I have traditionally assumed that simply not being racist was doing my part, and that the actions of others would eventually convert society at large to seeing race as the non-issue it should be. I have also felt that, as a white, straight male from a middle-class background, my voice would add nothing to this conversation or - worse - be seen as patronising or virtue signalling.

I now realise that this view was incorrect. The fact that people of colour are still fighting against global systemic marginalisation and persecution shows that being non-racist isn't enough, and that we must be outspokenly anti-racist, even if we have never experienced racial discrimination ourselves. Some may accuse me of jumping on a bandwagon with this. That's accurate, but I don't care. This is a wagon we should all be on, and I'm ashamed for not being on-board earlier.

In reflecting on racial issues for much of this week, it's been difficult to escape how prevalent racism is in Western culture once you open your eyes to it. Even palaeoart, which outwardly seems like an entirely innocent endeavour largely uncomplicated by social conventions, has been tainted. Palaeoart has actually been associated with suppression of non-white people both indirectly and rather pointedly, and not through obscure works or people, either. Famous historic figures, who are justifiably held in high regard for their scientific and palaeoartistic work, are part of this story. We're talking about people like pioneering European scientists - George Cuvier and Henry de la Beche - as well as pivotal American figures - Edward Drinker Cope, Charles Knight and Henry Fairfield Osborn - actively benefitting from black oppression or contributing to the thoroughly debunked pseudoscientific field of biological racism (the attempted justification of racist views with scientific study). The racist history of palaeoart thus hides in plain sight, and the fact that this seemingly innocuous branch of natural history illustration has this darker side is a great example of how rooted prejudice is within Western culture.

The first true palaeoart scene, Duria Antiquior, by geologist, palaeoartist and slave owner Henry de la Beche.

Three of the above-mentioned figures did not outwardly produce racist palaeoart, but strongly promoted or benefitted from racism against black people in other facets of their lives. Cuvier, who produced some of the first musculoskeletal reconstructions of fossil animals and was among the first to publish animal life reconstructions in academic works, viewed whites as the pinnacle of creation, but blacks as ugly, barbaric persons of monkey-like appearance. His work on dividing humans into 'scientifically validated' races was instrumental in later attempts at biological justifications of racism. De la Beche, famous for producing the first palaeoart scene in 1830, Duria Antiquior, owned a Jamaican slave plantation and was against the abolition of slavery. Cope, who drafted some of the first American palaeoart and mentored Charles Knight in the anatomy of prehistoric animals for a short period, wanted to see black people returned to Africa, and viewed them as degenerate forms of humans with animalistic levels of self-control.

The majority of the Henry Fairfield Osborn-directed Charles Knight mural Neanderthal Flint Workers (1924) - the right side of the image shows a valley and Coelodonta which is missing from this cropped version. This mural was hung in the AMNH's Hall of Man, an exhibition which included content based around Osborn's racist and eugenicist ideas. The 'primitive' look of these Neanderthals was not deduced from their bones, but based on features of non-white peoples that Osborn regarded as inferior to white Europeans.

Knight and Osborn, however, produced palaeoart that was genuinely racially offensive. It's not clear to me what Knight's views on race were*, but Osborn's racism, eugenicism, his support of Hitler, and his exploitation of his presidency at the American Museum of Natural History to research and promote his racist agenda, are well-documented historical facts. It was under Osborn's direction that Knight produced many now-iconic artworks of prehistoric animals in the late 19th and early 20th century but, while dinosaurs and fossil mammals can be restored without overt social agenda, Osborn's racism was captured in oils when it came to producing art for the AMNH's Hall of Man in the 1920s. Osborn viewed white Europeans as occupying a particularly high branch at the top of the evolutionary tree, with other races and hominid species well inferior. His Hall of Man exhibition sought to present the same view to the public, despite its lack of traction among his scientific peers. Neanderthals were thus shown as much more primitive, both culturally and anatomically, than modern humans, and Osborn ordered them to be restored as brutish and ape-like in Knight's famous Hall of Man mural, Neanderthal Flint Workers. But rather than pointing to apes or monkeys for 'primitive' anatomical references, Osborn pointed to non-white races that he regarded as inferior for details of skin colour and physique. The symbolism here is as gross as it is obvious, and makes the Osborn/Knight Neanderthals the palaeoartistic equivalent of golliwogs, minstrels and mammy figurines. Although this work was controversial among museum staff even when it was produced, and was removed in the 1960s out of recognition of its racist connotations, it remains an iconic, oft-used and oft-referenced illustration even today. Indeed, most popular depictions of Neanderthals are variants on this image: a dark-skinned, black-haired brutish caveman. How many artists, cartoonists and film directors know that their cavemen or Neanderthal depictions are a visual representation of eugenics and racism?

*Passages in Knight's book Prehistoric Man: The Great Adventurer (1949) suggest he shared Osborn's ideas that modern humans evolved in central Asia and not in Africa, and show that he viewed some non-Western cultures as 'primitive', but I'm not aware of any more pointed statements aimed at people of colour.

These are just a handful of easily researched examples of racism touching historic palaeoart, but there are more - particularly in depictions of human evolution. I will be accused of wanting to censor or vilify the figures mentioned above, so want to stress that this is not the point of this piece. The historic significance of these individuals and their contributions to science remain unchanged regardless of their personal views, and I don't regard it as fair to judge people living centuries ago, under very different social norms, by modern standards. Recognising and respecting the objective historic importance of their work is not the same as condoning their racist views. This said, I do condone the retiring of offensive works, such as Neanderthal Flint Workers, and believe that we should not shy away from discussing the racist views of key figures in the history of palaeontology. This is especially so for Osborn-like individuals, who had professional lives strongly influenced and guided by racist agendas.

I'm not sure what good discussing the above will do, but feel that saying something, and acknowledging the sometimes sketchy history of the discipline I work in, is important. It's an opportunity for folks like myself, who have never known the difficulties or prejudice, to say that we're listening, that we understand where we've gone wrong in the past, and that we stand with other races in their quest for fairer treatment and representation. The fact that racism has even touched and shaped palaeoart, a field which I think should be free of agenda and discrimination, is unnerving and sobering to me, especially since it involved artists and scientists who laid down its foundations. Some of the people we've discussed here are personal heroes of mine. And lest we forget, palaeoart is primarily an educational tool, and yet we've seen - via Neanderthal Flint Workers - how it can be corrupted not to show ancient realities, but to promote racist agendas. This is why Black Lives Matter is such an important movement, even if we are not black ourselves: we all have to reflect on the historic and modern ubiquity of anti-black racism if we want to stamp it out.

I began this post with an admission that I have arrived late to this cause, so want to end on a more positive and proactive note, by stating what I intend to do moving forward. I mention this in the hope that it will inspire similar action in others. Firstly, I will be making sure that my future public lectures on palaeoart acknowledge the human cost of early palaeoartworks. I will not be derailing my talks into anti-racist events, but it's nothing to point out that 19th century science had links with slavery and attempted to scientifically prove the inferiority of black people. We shouldn't just let that slide.

Secondly, I intend to obtain a deeper understanding of the pseudoscience of biological racism as well as the recent history of racism in Western nations. I have books on these topics lined up to listen to while working on art projects over the next few weeks - audiobooks being the most effective way for me to digest long reading material, as I rarely get enough time to read traditional books cover-to-cover. I especially want to know more about racism in the UK, as it remains a huge issue, but does not always get the attention it deserves.

Finally, I am also already taking a more active role in standing up to racism in online communities in an effort to provide a better online experience for people of colour. This includes taking a firm line on excluding racists from my social media. Blocking racist accounts, reporting and deleting offensive comments and calling out racism should be the norm in any decent community, but social media groups I'm part of are often surprisingly lenient on overtly racist commenters, even when their guidelines suggest otherwise. It's important to curate communities where people of any skin colour can enjoy themselves unharassed, and asking people to respect their fellow humans is a small price for engaging with their favourite scientists, artists and science communicators. Remember, our online platforms are not democracies: we get to call the shots about what is acceptable.

These are minor steps, but they are a start. Again, I'm not entirely sure what I hope to achieve with this post - perhaps it's simply a cathartic act to express and organise my recent thoughts on this topic. But I hope in sharing this that it inspires some good elsewhere, or at least shows one more supporting voice for a worthy cause.

Tuesday, 12 May 2020

Spinosaurus 2020: thoughts for artists

A 2020 take on some dinosaur or another. I forget its name. This individual has recently gorged itself, resulting in a distended belly and sleepy demeanour.

Unless you've been living under a rock for the last fortnight you cannot have escaped news on one of the most famous and controversial of all dinosaurs: Spinosaurus aegyptiacus. The appearance of Spinosaurus has once again transformed via the discovery of new fossils unearthed from the Late Cretaceous Kem Kem beds of Morocco: chiefly, a long paddle-like tail of superficially newt or crocodylian-like flavour. Keen interest in Spinosaurus, as well as a large National Geographic-led PR campaign for the new study, has seen social media awash with discussion about the new discovery, and illustrations of the latest in spinosaurine fashion have swamped online galleries since. As Chris Dipiazza eloquently explained on Twitter, it hasn't been the best two weeks if you aren't a Spinosaurus fan.
At the risk of numbing everyone further to Spinosaurus, I want to share some thoughts and reactions to this new research here. We've covered a few (but not all) of the twists and turns of Spinosaurus research in recent years (posts 1, 2, 3, 4) but, rather than simply writing another blog summary or popular rehashing of the new findings, I thought I'd write this from an artistic perspective, based on reading I conducted to produce my own take on "Spinosaurus 2020", shown above. As keen-eyed readers will note, I've not slavishly stuck to the same interpretations currently circulating the press circuit because - as we'll see - our takes on Spinosaurus are more complex than ever. Even with the tremendous amount of new data published on Spinosaurus in the last two decades, it remains the ultimate moving target for dinosaur palaeaoartists.

Spinosaurus 2020: where are we now?

Before we dive into this post, it makes sense to go beyond the recent Spinosaurus press coverage to look at what's in the new papers, as there's a lot more to them than what is being reported in the popular press. Needless to say, Spinosaurus has been an especially hot topic in dinosaur palaeontology since 2014 when Nizar Ibrahim and colleagues placed a newly discovered partial skeleton from the Moroccan Kem Kem beds at the core of a radical reinterpretation of a genuinely enigmatic animal. This was our introduction to Spinosaurus as a potentially short-legged semi-aquatic species, as well as proposals that spinosaurine material from across Northern Africa should be collected into one species, S. aegyptiacus, sinking several named taxa in the process (Ibrahim et al. 2014). It also proposed that the new partial skeleton should be the replacement exemplar specimen - the neotype - for S. aegyptiacus, after the original was destroyed in the Second World War to leave us with only Ernst Stromer's original descriptive work and photographs as records of its existence. Though widely publicised and catalysing a wave of public interest in Spinosaurus, the proposals of Ibrahim et al. (2014) proved controversial among academics. Numerous responses - some supportive, some critical - have been published by theropod researchers since.

Semiaquatic adaptations in a giant predatory dinosaur | Science
Spinosaurus as illustrated by Ibrahim et al. 2014. Many aspects of this reconstruction have been questioned and scrutinised in the last six years, but new data suggests that this may be closer to the appearance and proportions of certain spinosaurines than many of us initially believed.

Fast forward to today, and we've got not one, but two new papers by Nizar and colleagues that push discussions of all things Spinosaurus along significantly. The first is not Spinosaurus specific, but is an extensive monograph on the geology and palaeontology of the Kem Kem beds, now formally termed the Kem Kem Group (Ibrahim et al. 2020a). This is an important paper that brings some long-needed clarity and formality to details of Kem Kem stratigraphy and geology, including a new nomenclatural scheme to divide the Kem Kem into the Gara Sbaa and Douira formations. This is an important step for understanding the provenance of Kem Kem fossils which clarifies details of palaeoenvironments, relative ages of fossils, and comparisons with other fossil faunas (Ibrahim et al. 2020a). The entire fossil record of the Kem Kem Group is also reviewed, including a large discussion about Spinosaurus and its status as a Kem Kem theropod. Anyone interested in Spinosaurus and its world will need to check this paper out.

The second paper concerns additional material of the same 'neotype'* individual published in 2014, recovered from the same locality in more recent fieldwork (Ibrahim et al. 2020b). Among other finds, this includes a nearly complete tail that reinvents the appearance of Spinosaurus for the second time in six years. In addition to being short-limbed, it now seems that Spinosaurus had a deep, fin-like tail comprising narrow vertebrae with long and reclined neural spines and chevrons. This is interpreted as evidence of a swimming predatory ecology by the authors, it being argued that Spinosaurus could have swum like a crocodylian or newt to chase prey. The associated PR provides large amounts of media - videos, artwork etc. - showing Spinosaurus as a deep-diving species adapted to chasing large fish. This is not the first time Spinosaurus has been interpreted as a strong swimmer in recent years (e.g. Gimsa et al. 2016; Arden et al. 2019) but the recovery of a fin-like caudal skeleton adds a lot more weight to this argument.

*I'm going to refer to this specimen as the 'neotype' throughout this article for readability, as it's a catchier name than FSAC-KK 11888. That the proposal that FSAC-KK 11888 should be the Spinosaurus neotype remains controversial however, and will likely remain so until it's fully described and we can properly evaluate its similarity to Spinosaurus. I don't have a horse in this race but, for what it's worth, FSAC-KK 11888 looks like a member of Spinosaurus to me, although it has several differences from S. aegyptiacus that require investigation.

Artistic speculations that Spinosaurus may have borne some sort of tail fin have been common since the 2014 reinterpretation of the genus. Here's my finned version from 2016.

Both papers also provide comments in defence of Spinosaurus palaeobiology as proposed by Ibrahim et al. (2014). Criticism of their work included doubts about the authenticity and scaling of the neotype skeleton (Evers et al. 2015; Henderson 2018); its biomechanical feasibility as a swimming animal (Henderson 2018); the appropriateness of collating widely-dispersed and anatomically-distinguished North African spinosaurine material into one species (a question with particular reference to the overall number of spinosaurines in the Kem Kem) (Evers et al. 2015; Hone and Holtz 2015; Hendrickx et al. 2016; Maganuco and Dal Sasso 2018; Lakin and Longrich 2019); the suitability of the proposed Spinosaurus neotype (Evers et al. 2015; Maganuco and Dal Sasso 2018); and the general ecology of spinosaurines (Hone and Holtz 2015, 2019; Henderson 2018). The new data presented by Ibrahim et al. (2020a, b) addresses some of these concerns to an extent that some criticism - as we'll explore below - can probably be laid to rest. However, the enhanced debate around all things Spinosaurus means that these new papers have arrived in a much busier and more heated academic realm than their 2014 counterpart, and initial impressions from key players in spinosaurine research imply conversations will remain ongoing about aspects of lifestyle and taxonomy. For artists, this complicates our view of what Spinosaurus and other spinosaurines may have looked like, as well as what we can show it doing. While contributing important primary data on Spinosaurus, we have to remember that these new papers represent one interpretation of the appearance and lifestyle of a most unusual dinosaur in an increasingly busy academic debate, and that the ball is still in play.

Body plan and proportions

With that set up, it's time to dig into some art-relevant details. Firstly, I think Ibrahim et al. (2020b) adds a lot more confidence to the proposed strange proportions of Spinosaurus. The authenticity and scaling of the neotype have been questioned on grounds that it was collected, purchased and excavated by different people at different times (Evers et al. 2015; Henderson 2018), but the recovery the new tail and other elements in the same site as the pelvic, hindlimb and torso material, as well as their concordant proportions, suggests that all these remains were genuinely associated and likely belong to one individual (Ibrahim et al. 2020b). There is no evidence of other species in the bonebed and many broken bones of the neotype have now been reunited with once-missing pieces. Their histology and inferred growth stage are also matching. Courtesy of a quarry map illustration, we have a good idea of how these elements were associated in the field and how they relate to the material published in 2014.

Quarry map of the neotype locality and skeletal reconstruction of Spinosaurus, from Ibrahim et al. (2020b). Note the large area in which bones were found, the absence of non-spinosaurine bones, and the absence of bone duplicates: this is good evidence of the neotype representing a single individual, no matter how peculiar its proportions are. Known elements of the neotype are shaded in the skeletal, with different colours reflecting different field seasons and quarry locations. Scale bar represents 1 m.

With these data, and the fact that another spinosaurine specimen (Stromer's "Spinosaurus B") shows the same short-limbed morphology (Ibrahim et al. 2014), I think we can be fairly confident that at least some spinosaurines really were long-bodied, short-legged creatures with a body plan basically akin to that outlined by Ibrahim et al. (2014, 2020b). I know some folks are still holding out for data proving that the pelvis and hindlimbs belong with the vertebral column, but I think the burden of proof has shifted in light of these new data. Why aren't these legs associated with the body, given what we now know about the taphonomy of the site? A common question online is how much bearing the new tail has on other spinosaurids. We have sufficient skeletal remains of baryonychine spinosaurids (e.g. Baryonyx, Suchomimus) to suggest that they weren't fin-tailed, but the tails of spinosaurine spinosaurids aren't well known. The dorsal and caudal vertebrae of Ichthyovenator compare well with Spinosaurus, however (Allain et al. 2012), and it may have sported similar tail anatomy.

Posture and balance

Within the supplementary data of Ibrahim et al. (2020b) is a discussion of Spinosaurus mass and centre of gravity based on the (estimated) 11 m long neotype individual. Using a digital model and varying takes on tissue density, a mass of 3,219-4,173 kg was predicted and the centre of gravity was found to be just over one femur-length from the pelvic limb joint. This is fractionally more posterior than modelled in the 2014 model, if not quite as close to the pelvis as predicted by Henderson (2018). The cause of this shift is the larger tail and, although subtle, this difference has forced a reassessment of one of the most controversial aspects of the 2014 study: the presentation of Spinosaurus as a quadruped. Ibrahim et al. (2020b) now favour a facultative, rather than obligate, quadrupedal gait for terrestrial locomotion.

For artists, this means we can be a little more comfortable posing Spinosaurus as a biped, and I wonder if further work will substantiate bipedal poses further. Elsewhere in the supplementary data, Ibrahim et al. (2020b) suggest that the volume of restored tail musculature is conservative, and it stands to reason that models with more substantive tail volumes will pull the centre of gravity rearwards. Moreover, I wonder if the restored neck bulk is a little on the thick side, making the model more front-heavy. Among the neotype elements are long cervical ribs which, assuming typical tetrapod neck anatomy, could indicate displacement of some ventral neck muscles towards the torso (Taylor and Wedel 2013). Given that Spinosaurus already seems to have had a longish, low neck skeleton, displacing some of the neck muscle fraction posteriorly could have made for a relatively slender neck that would lighten the front end. If Spinosaurus also walked a little more upright than a typical theropod - using poses proposed by Andrea Cau, say - it might have avoided quadrupedality altogether.

Extended Data Fig. 8
Centre of mass estimates from Ibrahim et al. (2020b), compared to that of Henderson 2018 (C) and Ibrahim et al. 2014 (D).

I'm aware that some people feel that the legs of the neotype specimen are too slender to support the weight of Spinosaurus on land. The predicted 3 - 4-tonne masses of the neotype individual are relatively lightweight compared to theropods of similar length (>10 m theropods in the dataset of Benson et al. 2014 mass at 6-7 tonnes, for instance) and the hindlimbs would have to be held pretty straight to clear the animal from the ground (see illustrations, above). If so, the hindlimbs might have been loaded more like columns and imparted greater support than a traditionally bent theropod limb. Using hindlimb measurements from Ibrahim et al. (2014), I ran some very basic calculations on the strength of the neotype femur and found it critically weak against bending: it would fail when loaded with less than one 4-tonne body weight. When loaded as a column, however, it could take multiple 4-tonne masses. These calculations were very basic and ignore a lot of the nuance associated with theropod femoral posture but, if basically accurate, they suggest that the hindlimbs were strong enough to support Spinosaurus on land without help from weight-bearing forelimbs. I won't share the full details of these sums here as this post is already very long, but I can produce a follow-up article if it's of interest. Furthermore, while the hindlimbs themselves are small, there is evidence that aspects of their musculature - such as the caudofemoralis (a powerful hindlimb retractor) - were not reduced. In occupying much of the top half of the femur, the fourth trochanter of the neotype Spinosaurus femur is proportionate to the rest of the body (see for yourself in the 2014 image above, panel I, label 'ft') and suggests that the legs were capable of propelling their owner forward with suitable force, perhaps without propulsive assistance from the forelimb.

A topic I'm going to avoid here is the swimming posture of Spinosaurus, as this is an area that warrants further investigation before anything concrete can be said. I feel that the digital floating experiments with the 2014 Spinosaurus reconstruction by Don Henderson (2018) presented several worthy criticisms of Spinosaurus as an underwater swimmer, including its inability to sink due to pneumatised skeletal components (though some bones of Spinosaurus were pachyostosic (Ibrahim et al. 2014), its skull, neck and dorsal vertebrae were not) and the elevated centre of mass created by the tall, dense sail. A caveat about this study is that Spinosaurus had a relatively wider torso than was factored into the floating model, which would likely impact placement of the centre of mass and thus stability. We shouldn't dismiss Don's work because we assume this will correct the tipping issue, however: we need to see this investigated. We also have to consider the impact a wider torso would have on the suggested 'unsinkable' nature of Spinosaurus, as a wider torso will increase the lung volume fraction and impact buoyancy. For the time being we perhaps need to recognise that the body plan of Spinosaurus, even with its new tail, is entirely unlike any swimming animals alive today and that it's challenging to know how it functioned in water. Our science on this unusual dinosaur is in its infancy, and forming robust ideas about its swimming pose and capability is going to take time.

Floating spinosaurids in lateral and dorsal views.
Floating spinosaurids from Henderson (2018). One of the take-homes from Don's work is that Spinosaurus did not have an unusual floating posture among theropods, and that theropods were, in general, capable of floating with their heads well clear of the water to breathe. This questions whether features of the Spinosaurus skull linked to aquatic lifestyles - like the position of the eyes and nose - were specific adaptations to aquatic lifestyles.

Sail shape

One area where I'm less certain about the proportions of our new Spinosaurus reconstruction is the shape of the torso sail. Reconstructing the sail shape of Spinosaurus has always been difficult because the original S. aegyptiacus vertebrae were already not in great shape before Allied bombs blew them to pieces. As shown in Stomer's 1915 plates, the Spinosaurus neural spines were mostly disassociated from their centra; some were broken or deformed at their tips; and their arrangement within the vertebral series was not clear, even to those who saw them in person (Smith et al. 2006). Accordingly, several ideas about Spinosaurus vertebral order and sail shape have been proposed in the last century. While we seem to have a reasonable handle on the arrangement of the anterior sail vertebrae (artists, note that the neural spines project somewhat forward as well as up here: this is a common mistake in spinosaurine art), the shape of the posterior sail slope is more open to interpretation. Originally mounted in the Paläontologische Staatssammlung as a short, tightly-arced sail, Stromer rearranged the vertebrae into a longer, more gently sloping sail in 1936. Later, noting the reclined nature of the posteriormost-known sail spine, others proposed that the sail extended onto the tail (proposed independently by Andrea Cau in 2008 and Jaime Headden in 2010; Paul (2016) shows a similar arrangement while also matching Stromer's 1936 interpretation). More recently, Ibrahim et al. (2014; 2020b) have revived aspects of the pre-1944 Munich arrangement which brings shorter, sometimes anteriorly-positioned spines into a more posterior position (below).

Various restored shape shapes from a century of Spinosaurus. Images from Smith et al. 2006 and Ibrahim et al. 2014. Be sure to check out other takes on this sail by Andrea Cau, Jaime Headen and Scott Hartman.

I don't want to pretend that I know which of these arrangements is correct. Arranging these vertebrae is complicated, and there are multiple, perhaps equally viable ways we can order them at present. Based on the new tail data, I suspect the interpretation of Stomer and Ibrahim et al. are correct in restoring the sail plunging sharply into the tail base, but I also see merit to Stomer's 1936 model where vertebra 'f' - the cause of the dip in the Ibrahim et al. model - is positioned more anteriorly.

Unfortunately, the neotype material seems to complicate the resolution of the sail shape further. The sail spines of the neotype are noticeably more slender than those of the holotype despite coming from animals of generally similar size (the neotype is an estimated 11 m long, vs 12 m for the holotype; Dal Sasso et al. 2005; Ibrahim et al. 2020b) and, as preserved, they are quite a bit shorter. Does this imply a lower, less robust sail in the neotype individual, or is this something to do with growth, sexual dimorphism, or another form of variation within Spinosaurus? It's here where our taxonomic assumptions start impacting our reconstructions. Ibrahim et al. (2020a, b) regard S. aegyptiacus as an anatomically variable species, suggesting that we might be OK to blend data from the holotype and neotype sails. Conversely, other schemes regard S. aegyptiacus as potentially confined to Egypt and cast the 'neotype' as a closely related animal (e.g. Evers et al. 2016; Maganuco and Dal Sasso 2018), in which case we might focus more on the sail shape specifically indicated by the Kem Kem specimen. I don't know that there's a clear answer to this conundrum, so artists probably have several options for Spinosaurus sail shapes at present. My own reconstruction follows a somewhat more Stromer 1936-compliant model, as well as a sail height conservatively modelled on the neotype specimen.

Tail flexion

Among the more interesting aspects of the new Spinosaurus tail is the reduction of zygapophyses in the distal region. This potentially allowed the tail to flex far more than was typical for a theropod and to be used for swimming (Ibrahim et al. 2020b). I was initially sceptical of this claim because the long neural spines of the tail extend not only upwards, but also backwards over several other vertebrae, meaning that any movement between vertebrae required the spines to bend in multiple places or else project at wide angles from the tail curve. This is not a novel observation on neural spine length in potentially aquatic animals: I'm basically rehashing arguments made by Silvio Renesto et al. (2010) about the unusual tail of the drepanosaur Hypuronector, and how its extremely long, backwards-projecting chevrons stiffened the tail against sculling-like swimming motions. What I forgot, however, was that Hypuronector also had very developed 'clamping' zygapophyses (Renesto et al. 2010), and what I didn't realise is that - according to folks who know a lot more about biomechanics than I do - the 15 mm wide neural spines of the Spinosaurus tail could probably bend quite far. The bones of healthy living animals are somewhat plastic and capable of flexion, but I was surprised to learn that muscles and ligaments binding the Spinosaurus tail together would let relatively thick bony rods bend considerably without failing. So perhaps there's less of a problem here than I anticipated, though I admit to wondering how this would work given that Ibrahim et al. (2020b) only reconstruct a very slight covering of soft-tissues on the distal neural spines (below). If muscles only extended up the basal portion of the spine, was this enough to hold the tail together as it sculled the animal through water?

Fig. 1
The new tail of Spinosaurus, as presented by Ibrahim et al. (2020b). Note the reduction of musculature in the distal tail ('e') in relation to the discussion of bone bending, above.

Based in part on these discussions, I've been wondering how much flexibility we can safely reconstruct in the tail. Some of the recent PR imagery has shown a degree of tail flexion that seems beyond that of crocodylians, which seems excessive even allowing for some plasticity in the tail bones. Crocodylian tails have relatively short neural spines and chevrons, as well as large transverse processes to anchor large, strong musculature along much of the tail length. This allows them to pull their tails into tight arcs but, as noted by Ibrahim et al. (2020b), the transverse processes in Spinosaurus are restricted to the anterior tail region in a pretty typical theropod fashion. This musculoskeletal arrangement is thus not very crocodylian-like, and I wonder if the tail was more flexible than usual for a theropod, but maybe not to the degree where it could form a tight, crocodylian-style arc. I also wonder if the energy stored in bending neural spines would spring the tail straight once muscular effort was relaxed, which might have been especially significant when the tail was unrestrained during walking or floating. Maybe, for all its potential flexibility, the tail was held largely straight unless it was actively being used in swimming, or braced against something in the environment.

Facial anatomy and lips

To close out this post, I want to briefly touch on a topic not directly covered in the recent Spinosaurus work, but that comes up whenever spinosaurid illustrations are discussed: did these animals have lipless, crocodylian-like faces? In my experience, lipless spinosaurids are justified by several lines of evidence: their superficially crocodylian-like jaws and teeth; the size and configuration of their anterior teeth (where large premaxillary teeth overbite the lower jaw and long dentary teeth - unusually for a theropod - protrude over the upper jaw during occlusion; Dal Sasso et al. 2005), and the development of liplessness in other semi-aquatic fishers, such as crocodylians and river dolphins.

But when looking at spinosaurid jaws with the same criteria generally used to predict extra-oral tissues in fossil animals (tooth size, tooth orientation, jaw bone foramina counts), spinosaurids do not seem unusual compared to other theropods. Their jaws appear peculiar in some ways - check out that foramina rich anterior rostrum, below - because of their atypical geometry, but beyond this, much of their jaw configuration is typically theropodan. Their jaw foramina counts, for example, are not significantly high. Foramina frequency in tetrapod jawbones (premaxilla, maxilla, dentary) have been provisionally hypothesised as indicating the presence of extra-oral soft-tissues in tetrapods (Morhardt 2009), so we can compare foramina counts of Spinosaurus to other tetrapods to infer their facial configuration. Ibrahim et al. (2014) give a Spinosaurus upper jaw foramina frequency of 125, which seems high, but this value represents four bones worth of foramina. Crocodylians have this many foramina, and perhaps many hundreds more, in a single jaw bone. Morhardt (2009) suggests that we need about 100 foramina per jaw bone to infer a lipless condition, which Spinosaurus is well short of. This point recalls comments that the foramina counts and inferred sensitivity of Spinosaurus jaws, which have been correlated to aquatic lifestyles by some authors (Ibrahim et al. 2014), may have been pretty standard for large theropods (Barker et al. 2017), and are possibly not related to aquatic lifestyles or unusual facial anatomy.

Spinosaurids are often suggested to be among the more likely dinosaurian candidates for liplessness and exposed teeth, but the key features we might look for regarding this condition - labial foramina counts and distribution, as well as jaw bone texture - are not atypical for theropods, nor are they especially crocodylian-like. Their large teeth, including those at the jaw anterior, are no larger (relatively speaking) than those of extant animals with immobile lips and sheathed dentitions (bottom row). Spinosaurus elements after Dal Sasso et al. (2005), Neovenator after Barker et al. (2017); American alligator cropped from original on Wikimedia by Didier Descouens, CC BY-SA 4.0.

We can also observe that the maxillary and dentary jaw foramina of Spinosaurus are arranged in a more lizard-like row along their oral margins, and not - as in crocodylians - distributed in a dense pattern across the entire jaw. In Spinosaurus at least, they seem to be placed some distance from the toothrow (Dal Sasso et al. 2005) in a lizard-like configuration. This would keep the nerves and blood vessels running into any lip tissues well clear of the overlapping dentary teeth when the mouth was closed, but - based on lizards with similarly displaced foramina - I don't think this means anything too radical for life appearance (lips, if present, would not look unusually big or weirdly anchored). The absence of unusual, epidermally-derived textures on Spinosaurus jaw bones is a further distinction from crocodylians. As we've discussed at length in other posts, the characteristic rugosity of crocodylian skulls is reflective of their facial skin and sensory tissues, so the absence of comparable characteristics in Spinosaurus is strong evidence of a different anatomical regime. I'm also not convinced that the teeth of Spinosaurus - so far as they are known (to my knowledge, Spinosaurus jaws with a complete set of teeth remain elusive) - are too large for sheathing behind lips. We have reptiles today with large teeth at their jaw tips and they do not protrude from their lips (above): to the contrary, you'd have no idea they were there from their external appearance.

Finally, what about the purported link between liplessness and fishing aquatic lifestyles? I feel that this reflects a focus on lipless semi-aquatic or aquatic tetrapods but ignorance of the great number of secondarily-aquatic fishers that have retained fully-sheathed dentitions. Yes, crocodylians and river dolphins have unsheathed teeth, but many other fishing swimmers - cetaceans, seals, otters, mink, water monitors, numerous snake species and so on - do not. In fact, many have facial tissues little different to their terrestrial relatives. This questions whether lifestyle is a useful predictor for facial anatomy in Spinosaurus. This is surely a problematic line of evidence anyway, given that it remains to be determined exactly what sort of habits were common to Spinosaurus. Was Spinosaurus an underwater pursuit predator (Ibrahim et al. 2014, 2020b; Gimsa et al. 2016), something more akin to a heron (Hone and Holtz 2015, 2019; Henderson 2016), or something in between? As with so much about Spinosaurus, we have a lot of primary questions to answer before we can start thinking about their implications for behaviour and life appearance.

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  • Taylor, M. P., & Wedel, M. J. (2013). Why sauropods had long necks; and why giraffes have short necks. PeerJ, 1, e36.

Wednesday, 22 April 2020

In pursuit of giant pliosaurids and whale-sized ichthyosaurs

Some marine reptiles - like Shonisaurus popularis - were big. So big, in fact, that you can't fit them all into a picture. But just how large did the largest marine reptiles get? Finding the answer can be more involved than you'd think. This image is a cropped version of a larger picture, which you can see at my Patreon.
The size of Mesozoic marine reptiles is undeniably a big part of their charm and popular appeal. And yet, if you want to know how big some of these animals really got - and by 'really', I mean 'size estimates preferred by contemporary scientists based on our latest data' - you might have a struggle on your hands. Marine reptile media is full of varying size estimates for different groups, some of which are accurate to modern science, some of which were accurate to old science, and many of which were never accurate to begin with. It's not always easy to tell which is which.

I've recently had cause to ascertain reliable size estimates for a number of marine reptile clades and found it far more involving than I anticipated, especially for pliosaurids and ichthyosaurs. This isn't necessarily because it's hard to find an article or paper giving figures of total length or mass, but because confused taxonomies, a deficit of complete skeletons and opaque comments in technical literature complicate efforts to recover and understand maximum size estimates from peer-reviewed, reliable sources. I thought it might be of interest to share some of my findings here, focusing specifically on those two aforementioned groups - pliosaurids and ichthyosaurs - as I feel their upper size range is perhaps most in need of clarification.

Before we get going, it's worth reminding ourselves of how estimating the size of very large extinct individuals is fraught with challenges, and how the numbers and stats we throw around are often based on relatively little data. Many well-known scaling problems apply to ichthyosaurs and pliosaurs. Our remains of especially large individuals are not only rare but are often relatively scrappy, and our capacity to scale reliably from other, smaller animals is frequently more limited than we might like. As is often necessary, there's a need to distinguish between size estimates that are generally reliable, being based on relatively complete large skeletons, and those which are only indicative, on account of being scaled from fragmentary remains. At the risk of spoiling details of the article below, there doesn't seem to be a clear answer to "what's the biggest pliosaurid" or "what's the biggest ichthyosaur" because the size estimate error bars for most very large specimens are too wide to give precise assessments.

Giant pliosaurids

The great size of certain Late Jurassic and Lower Cretaceous pliosaurids has made them famous, perhaps especially so in the UK and Australia where giant animals are particularly well represented. Since their discovery in the mid-19th century, there has been no doubt that certain pliosaurids attained enormous sizes and dwarfed virtually all other organisms in their respective palaeoenvironments (e.g. Owen 1841; Longman 1924; Tarlo 1959).

How the world met pliosaurids: plate 68 of Owen's Odontography (1841). That's a life-sized Pliosaurus tooth at the bottom of the page. Note the spelling - "Pleiosaurus". It's not clear why Owen changed the name later, but "Pleiosaurus" has not been widely used since the 1840s and Pliosaurus has, by rules of zoological nomenclature, won out.
But the question of how large pliosaurids were has, for a long time, not had a straightforward answer. An ongoing problem with estimating their size is that complete pliosaurids, in stark contrast to plesiosauroids, which are known from abundant complete remains, are actually pretty rare. Any attempt to estimate their size thus has to tackle uncertainty about their body proportions. It took over a century for palaeontologists to attempt the first calculations of pliosaurid body length, and these were still produced while aspects of vertebral number, torso length and so on remained unknown (Romer and Lewis 1959). This estimate was associated with the famous 12 m long Harvard Kronosaurus mounted skeleton, a reconstruction that has unfortunately covered the original fossil in so much plaster that it's now hard to examine in detail. It's now generally considered that the Harvard mount has too many vertebrae and is thus too long (McHenry 2009). The subsequent recovery of relatively complete pliosaurid specimens has supplied useful data for estimating their size (e.g. Noè et al. 2001; McHenry 2009) but we are still working with a very provisional dataset and assuming that the proportions of a handful of species apply to pliosaurids in general.

Newman and Tarlo's (1967) unnamed pliosaurid reconstruction, interpreted by McHenry (2009) as the 10 m long Pliosaurus macromerus (a species of uncertain validity today - see Knutsen 2012 vs. Benson et al. 2013) or Liopleurodon ferox, based on a near-complete, 4.88 m long specimen held in Tübingen, Germany (Noè 2001). I like the generous soft-tissue outline of this illustration - it looks appropriately bulky and well-muscled.
Famously, 1999 saw claims of whale-sized pliosaurs take root in public conscience courtesy of the BBC documentary Walking with Dinosaurs. The antagonist of the Jurassic marine-focused third episode was a 25 m, > 100-tonne Liopleurodon ferox which was, even given the general uncertainty about pliosaur size, a very controversial reconstruction for a species only known from much smaller remains. Our largest Liopleurodon skull (discovered in the early 20th century, so known to the producers of the programme) is 1.54 m long (Benson et al. 2013) so, assuming a skull:body length ratio of about 1:5 (Noè et al. 2001; McHenry 2009), we can predict that large Liopleurodon individuals were about 8 m long. How did this value become tripled for the documentary? Writing in 2000, two of my colleagues - David Martill (my PhD supervisor, also a consultant for WWD) and a pre-TetZoo Darren Naish explored the rationale of this decision:
This size created much debate in palaeontolgoical circles following the first airing of the programme, as no palaeontologist thinks Liopleurodon really got this big.
Although several complete [Liopleurodon] skeletons have been discovered, these are of individuals of between five and ten metres in length. It is less complete remains discovered in the Oxford Clay that indicate lengths greater than this, though here we move into an area of rough estimates and guesswork. A vertebra at the Peterborough Museum, brought to light in 1996, would seem to indicate a pliosaur of between seventeen and 20 metres in length, and various fragments of snout and lower jaw in other museum collections suggest specimens of similar size. Whether these fragments are actually from Liopleurodon is uncertain, and the animal to which they belonged has been nicknamed 'Megapleurodon'. Given that it is unlikely that these bones really represent the very biggest pliosaur specimens that ever lived, some experts cautiously suggest that Liopleurodon and related forms may have achieved total lengths of around 25 metres.
Martill and Naish 2000, p 80.

The WWD Liopleurodon was a showstopper back in 1999, and one of the stars of the series. Its arrival brings the end to this sequence showing Ophthalmosaurus giving birth. Clip from Walking with Dinosaurs, uploaded to Youtube by BBC Earth.

As is clear from this text and subsequent Tetrapod Zoology articles, both Dave and Darren knew that this size estimate was very speculative and an indulgent move by the programme-makers. But while a 25 m long pliosaur isn't defensible, supersized pliosaurs were not entirely out of the question 20 years ago. In addition to the specimens mentioned by Martill and Naish, a fragmentary Mexican pliosaurid (the so-called "Monster of Aramberri") was inferred as reaching 15 m just a few years after WWD aired (Buchy et al. 2003). While this is still some distance from 25 m, these data were pointing to pliosaurs of much larger sizes than generally anticipated during the late 1990s and early 2000s.

Unfortunately for those excited by the idea of a whale-sized pliosaurid, the specimens touted as rationalising the WWD monster have not delivered on their promise. In what is perhaps the most detailed assessment of maximum pliosaurid size conducted to date, Colin McHenry (2009) discussed all the fragmentary material from Britain and Mexico linked with supersized pliosaurids and found that they represented very large animals, but not whale-sized giants. Colin confirmed that the vertebra housed at Peterborough Museum was indeed very large - 252.5 mm wide by 219 mm tall - but its total body length estimate was just 11.6–14.2 m when scaled to well-known pliosaur remains [Note added 24/04/20: also see the comment from Colin below, for an update on this specimen). A large Jurassic mandibular symphysis archived in the Natural History Museum, London might represent an animal anywhere between 9.1 and 15.1 m long, while OUM J.10454, a near-complete, 2.8 m long lower jaw dubiously referred to Pliosaurus macromerus and known informally as the 'Cumnor mandible', scales to a surprisingly low 12.7 m. In subsequent years this estimate has also become questionable as we've realised how reconstructed the Cumnor mandible is: it remains to be determined what size range the original specimen represents (Benson et al. 2013).

The Tübingen University specimen of Liopleurodon ferox, a near-complete 4.8 m long juvenile. This is a key specimen for our understanding of Jurassic pliosaur proportions, and thus the size of the biggest individuals. From Wikimedia user Ghedoghedo, CC BY-SA 3.0.
The same story plays out across all other giant pliosaurid fragments, including the Mexican Monster of Aramberri. It seems that when scaled using our best data on pliosaurid proportions and growth allometry, most 'giant' pliosaurid fossils return total length estimates in the 10 m range, with only the upper bounds of our least reliable length estimates falling outside this (McHenry 2009). There's perhaps a moral here about reading too much into the size of giant but fragmentary bones. As stated by McHenry (2009):
Estimates made on incomplete series of vertebrae, or even a single vertebrae, are subject to the natural variation of vertebral dimensions and should be used with caution. The dimensions of individual vertebrae can be affected by taphonomic processes, in particular sedimentary compaction, and when size estimates are extrapolated from single elements small errors can be greatly magnified. The same applies to any allometric variation than is not accounted for in scaling models... any estimates based upon more complete pliosaurid material requires extrapolation over at least an order of magnitude of body mass, a leap that means even small errors in the estimate of allometric or intraspecific variation will produce a large range of results.
McHenry 2009, p. 422

My take on one of the largest Jurassic pliosaurids, Pliosaurus kevani. It was probably about 10 m long, assuming similar proportions to other pliosaurids. The smaller animal above is a calf.
So, putting the ideas of gigantic pliosaurs to the side, what are our most reliable estimates for the maximum sizes of these animals? Detailed assessments of the very large and relatively well-known pliosaur Kronosaurus queenslandicus suggest that a total length of 10.5 m and a mass of 11 tonnes is likely (McHenry 2009). Kronosaurus is one of the few large pliosaurids known from decent and articulated postcranial remains so this figure involves minimal extrapolation, and should be regarded as one of our more reliable insights into the maximum size of these animals. It seems that c. 10 m is about right for other large pliosaurids, too. The skulls of large Kronosaurus are 2.3 m long, a value comparable to measured and estimated skulls lengths of the largest Jurassic pliosaurid, Pliosaurus (2-2.5 m; Knutsen et al. 2011; Benson et al. 2013). If pliosaurid skulls account for roughly 20% of their body length, these are on target for a Kronosaurus-like size of 10 - 12.5 m. While that upper range is obviously higher than Kronosaurus, we shouldn't put much stock in it yet. Our 12.5 m Pliosaurus is a body length extrapolation based on a rough skull size calculation, and both these values hinge on the specimen in question having the same skull and body proportions as other, mostly non-Pliosaurus pliosaurids. Our 12.5 m Pliosaurus is thus perched on a stack of assumptions, each one compounding any errors of the one beneath it. Such methods are fine for getting ballpark size figures but they are far from precise and we should treat them with appropriate caution. Greater understanding of the size of the biggest pliosaurs will require discovery of more substantial skeletons of large pliosaur individuals.

Giant ichthyosaurs

It's remarkable to me that the giant ichthyosaurs, which are without doubt the largest marine reptiles of all time, appeared in the Late Triassic - just a few tens of millions of years after marine reptiles entered the seas. Ascertaining details of how large these animals got is complicated but not, as with pliosaurs, because of scant remains. To the contrary, we actually have several excellent fossils of very large ichthyosaurs (Camp 1980; Kosch 1990; McGowan and Motani 1999; Nicholls and Manabe 2004), and our challenge is instead related to working out what taxa they represent. The history of two ichthyosaur genera associated with giant size - Shastasaurus and Shonisaurus - is complex and intertwined. While there's no doubt that Shonisaurus contains at least one species of very large ichthyosaur (Sho. popularis), it's not always been clear whether Shastasaurus also contained giant animals, nor to which of these genera the largest ichthyosaur remains should be referred to. The confused treatment of these animals in technical literature has spread into the popular realm and it's very easy to find articles and art treating Shastasaurus and Shonisaurus interchangeably. This is unfortunate because, as we'll see, they're actually very different animals.

Quarry map of the holotype specimen of Shonisaurus sikanniensis, the biggest marine reptile yet known from a decent portion of skeleton, from Nicholls and Manabe (2004). The last decade has seen differing teams debating whether this animal is actually Shastasaurus or Shonisaurus. Note the 1 m scale bar at the bottom of the illustration - this animal was very big.
I won't recount the full taxonomic history of these genera here, it will suffice to say that long-running uncertainty about recognising and diagnosing Shastasaurus and Shonisaurus is the crux of this issue. Both genera have housed an ever-changing number of specimens and species in their long histories (Shastasaurus was named in 1895, Shonisaurus in 1976) and most of their new species have proved problematic one way or another: generally being undiagnostic, based on controversial specimen allocations, or being better referred to other named ichthyosaurs. The conflicted history of Shastasaurus and Shonisaurus continues today as authors continue to disagree over which specimens and species should be allocated to each genus (e.g. McGowan and Motani 1999, 2003; Nicholls and Manabe 2004; Sander et al. 2011; Ji et al. 2013).

This affects our assessment of ichthyosaur size because the taxonomic fate of several giant specimens are at stake in these disagreements, including the holotypes of "Shastasaurus careyi" and Sho. sikanniensis. This might not seem like a big deal - a giant ichthyosaur is a giant ichthyosaur, right? - but not all Triassic ichthyosaurs are alike, and the generic identification of these specimens influences our predictions of body dimensions. Although both Shonisaurus and Shastasaurus are generally considered to be shastasaurids, this group is not anatomically uniform enough that we can liberally borrow proportions from other species to calculate size. Some taxa, such as Shonisaurus, were long-snouted, deep-bodied forms, while other genera, such as Guanlingsaurus, were very long, slender animals with small skulls and short faces. For Shastasaurus in particular, trying to work out the anatomical characteristics of this genus is very challenging from all the material referred to it. Even the configuration of the skull seems open to question: was it a long-snouted, toothed form (e.g. Callaway and Massare 1989) or a short-snouted, edentulous form (Sander et al. 2011)?

Not all shastasaurids were deep-bodied, Shonisaurus-like animals. Several genera, like Guanlingsaurus and probably Shastasaurus, were slender, long-bodied animals with shortened faces, so it's important we know what sort of body plan our giants had to calculate their proportions. From Ji et al. 2013.
Happily, it seems that some clarity is emerging from this murk. A large number of ichthyosaur workers are now taking relatively conservative taxonomic approaches to Shastasaurus and Shonisaurus, restricting their specimen inventories to incontrovertibly assigned historic material and rationalising species taxonomy down to better-known, well-diagnosed fossils. This renders Shastasaurus a monospecific genus containing just Sha. pacificus, while Shonisaurus has two well-represented species, Sho. popularis and Sho. sikanniensis. This tidying up means we have a more concrete idea about their shape and form, and has had one surprising outcome: after decades of being touted as an ichthyosaur giant, it turns out that Shastasaurus was actually of unremarkable size. I couldn't find a published body length calculation for Shastasaurus, so I roughly predicted a total length of just 6-7 m using the partial skull illustrated by Sander et al. (2011) and the proportions of a large Guanlingsaurus. I admit to being a little disappointed, as I quite liked the idea of giant ichthyosaurs including both robust, deep-bodied forms like Shonisaurus as well as long, slender animals like Shastasaurus but... oh well. Unless something changes in future, we need to stop talking about Shastasaurus as a giant ichthyosaur.

The skull of Shastasaurus pacificus, illustrated by Sander et al. (2011). Stripped to the core hypodigm and type species, Shastasaurus is neither a well-known nor particularly big animal. I roughly estimate this skull to have been c. 60 cm long, which is big, but not remarkably so for a shastasaurid. Note the large eye socket and pinched snout: Shastasaurus had quite a different skull to the more traditionally ichthyosaurian Shonisaurus.

This leaves the Shonisaurus species as the named record holders of ichthyosaur size, and by some margin. Both species are known from substantial remains that allow us to be fairly confident in our body length estimates. We can actually get a lot of data from simply measuring their articulated skeletons. Our best size predictions for these animals shake out to 13-15 m for Sho. popularis (McGowan and Motani 1999) and a whopping 21 m for Sho. sikanniensis (Nicholls and Manabe 2004). Using data from Gutarra et al. (2019), these equate to approximate body masses of 20-30 and 80 tonnes, respectively. The Shonisaurus species were huge animals, among the largest to ever swim the seas.

As with our giant pliosaurs, several fragmentary Triassic ichthyosaur fossils are touted as indicating larger animals. Himalayasaurus tibetensis, if valid and not another representation of Shonisaurus, is not well known but seems to have rivalled or exceeded Sho. popularis in size, though by how much cannot be reliably ascertained (Motani et al. 1999). Giant remains dubbed the "Mount Potts ichthyosaur" from New Zealand, first reported in 1874, included vertebrae of some 450 mm diameter and ribs exceeding a metre long, making them comparably-sized or bigger than other known ichthyosaur fossils. Alas, they are now lost and these claims cannot be investigated further, making them fairly meaningless anecdotal evidence for extreme ichthyosaurian body size (Fleming et al. 1971). A portion of posterior lower jaw known informally as the "Lilstock ichthyosaur" from Somerset, UK, has been roughly estimated as being similar in size to Sho. sikanniensis at 20 - 25 m long, and another UK fragment (from Aust Cliff, Gloucestershire) possibly hints at an even larger or more robust individual (Lomax et al. 2018). These specimens are genuinely large chunks of bone and perhaps represent our most intriguing hint of even larger marine reptiles, but they're also such small pieces of evidently gigantic animals that we can only very roughly anticipate their size, especially since our scaling calculations for giant ichthyosaurs are still unable to factor proportional changes with growth (Lomax et al. 2018). It's thus hard to know exactly what to make of these specimens, except that they show additional evidence of roughly Sho. sikanniensis-sized creatures in the Late Triassic. For now, values around 21 m remain our most substantiated length estimates for giant ichthyosaurs, and the upper size threshold for marine reptiles as a whole.

My take on a 21 m long Shonisaurus sikanniensis, shown here with generic, 6 m long, Sha. pacificus-sized shastasaurids to stress the size difference between these two 'giant' genera. Shastasaurus is a little older than Shonisaurus and the two didn't live alongside one another but, if they had, Shastasaurus would have been dwarfed by its larger cousin.

What hope is there for blue whale-sized marine reptiles?

As an epilogue to the discussion above, I want to briefly share some thoughts on a marine reptile Holy Grail: a species comparable in size to the modern blue whale Balaenoptera musculus. Despite no marine reptile approaching the size of the largest blue whales (c. 30 m total length, over 100 tonnes) it's easy to find media comparing marine reptiles against our biggest modern cetaceans. Size estimates for the largest shastasaurids match the proportions of very large baleen whales, including certain Pacific blue whale populations - which generally reach 20-something metres in length - but obviously fall short of the largest, primarily North Atlantic and Antarctic B. musculus individuals. Should ever expect to find a 30 m long, 100+ tonne marine reptile?

It's easy to be blase about whales, but the fact we share the planet with the biggest animals to have ever lived on Earth is not to be taken for granted. It seems that the extreme sizes of large rorquals, like this blue whale, may be tied to certain unique biological and environmental properties, which means 30 m body lengths may not be attainable for just any marine tetrapod. (Image from Wikimedia, in public domain).
Studies on the factors influencing body size in marine animals indicate that food availability and foraging efficiency might be strong limiting factors on their upper size limits, and that increasing feeding efficiency via bulk filter-feeding gives rorquals a substantial edge over other species (Goldbogen et al. 2011; Gearty et al. 2018). Enhanced capacity to obtain energy from foraging means more resources to build body tissues, which feeds directly into physiological advantages of larger body size. Together, these strongly incentivise the development of gigantism where rorquals can find enough food, and it seems large blue whales have run with this about as far as they can, reaching predicted biomechanical limits for operating their jaws in water (Potvin et al. 2012). We can thus infer that the extreme size of living baleen whales is dependent on both their unique foraging mechanism and periods of high oceanic productivity (Gearty et al. 2018). This latter factor might explain why whales only developed extreme gigantism in the last few million years, despite their likely development of baleen 23 - 35 million years ago (Gearty et al. 2018; Peredo et al. 2018).

We should, not, therefore, look at the maximum size of rorquals and assume that they represent a universally obtainable figure for all marine vertebrates. As with all organisms, upper size limits are dependent on a complex interplay of anatomy, physiology and environment that are unique to every species, and it's not a given that marine reptiles had the same adaptive potential to reach the same size as the largest rorquals. While it's hard to know exactly how productive Mesozoic oceans were, we can certainly identify the lack of bulk filter-feeding mechanisms as a probable size-limiting factor for marine reptiles, and this may well explain why we've yet to find indications of ichthyosaurs much above 20 m long. Of course, we can't say that fossils of larger marine reptiles aren't out there, waiting to be found, but my guess is that, if giant blue whale-sized marine reptiles existed, they would be anatomically and ecologically very different to our currently known species.

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