Thursday 25 July 2019

The science of the Crystal Palace Dinosaurs, part 4: The mammals of the Tertiary Island

An 1853 illustration of one of the Crystal Palace mammals, Palaeotherium magnum, imagined next to the plesiosaurians it would eventually share the Geological Court with. The Palaeotherium sculpture that this image is based on is now lost, just one of many misfortunes to befall the Crystal Palace mammals. From Die Gartenlaube, archived at Wikipedia.
It's time for our final visit to the prehistoric animal sculptures of Crystal Palace Park. Having toured through the Palaeozoic and Mesozoic exhibits in the last three posts (part 1, part 2, part 3), today we turn our attention to the Cenozoic section of the Geological Court, one containing exclusively mammalian palaeoart subjects. As with previous entries in this series, these words stem from writing palaeoart notes for the Friends of Crystal Palace Dinosaurs charity - please check out part 1 for additional context about their work and the ongoing need for care and maintenance of the Crystal Palace sculptures. You can help conservation efforts by donating money or volunteering your time to keep the Geological Court maintained.

At risk of being melodramatic, I find it difficult to escape a sense of tragedy when concerning myself with the Crystal Palace mammals. They are the least documented, least discussed and most suffering of the sculptures, with several bearing obvious hallmarks of neglect and low-quality repair work. Even in the 1850s they were being sidelined to make way for the more spectacular fossil reptiles, a fact all the more tragic because extinct mammals were the inspiration for having prehistoric animals at Crystal Palace in the first place (McCarthy and Gilbert 1994). Initially, Hawkins planned to restore a woolly mammoth and other large mammals but, when developing his display, his attention was drawn to the dinosaurs and fossil reptiles which would ultimately consume most exhibition space and public interest. Once located on their own 'Tertiary Island' (Doyle and Robinson 1993), the mammals are today situated on the 'mainland' component of Crystal Palace and it's hard not to view them as being a little tucked away. Although close to the reptile displays (within sight of the Mosasaurus), most of the mammals are located on a separate path to the reptilian sculptures and they are often obscured by foliage. It's quite easy to miss them on a casual walk around the park.

A sense that the mammals were being overshadowed by reptiles may explain why Hawkins wanted to expand this component of the park. He wrote to Owen in 1855 with a plan to augment his Cenozoic fauna considerably, including models of a mammoth, a bathing Deinotherium, glyptodonts, Sivatherium and extinct bison, as well as moa, dodo, turtles and snakes (Doyle 2008; Dawson 2016). This letter was dated a full month after Hawkins was no longer working for the Crystal Palace company however, who thought he had constructed enough models and would not even let him finish a half-completed mammoth sculpture. The official reason for terminating Hawkins' work was allegedly an artistic one of “less being more” (McCarthy and Gilbert 1994), but the abrupt termination of Hawkins’ project surely reflected the financial struggles of the Crystal Palace Company shortly after the park opened (Dawson 2016). It’s certainly difficult to believe that the same company who filled their park with reconstructed ancient buildings, expanded the Crystal Palace to incredible dimensions and built fountains rivalling the biggest in Europe would suddenly be concerned about artistic excess. Some idea of what Hawkins’ mammoth, Sivatherium and turtles may have looked like may be taken from his later drawings and paintings, including his 1860s poster series that borrowed heavily from his Crystal Palace designs (Rudwick 1992).

Hawkins' grand plans for the Tertiary Island, drawn on the back of a letter to Owen in 1855. He wanted it packed with Cenozoic mammals, reptiles and birds, but the Crystal Palace Company thought this was excessive (or, more likely, couldn't afford to pay for the work). From Doyle (2008).
To my knowledge, very little information survives regarding Hawkins’ construction of the four Crystal Palace mammal species. They were built and installed at the same time as the other models but were not mentioned in Owen’s (1854) Geological Court guidebook despite his interest in fossil mammals (e.g. Owen 1846). Perhaps this is further evidence of Owen’s general disinterest in the Crystal Palace project? Victorian visitors had to make do with a very brief and incomplete overview of the mammal fauna provided in the general Crystal Palace Park guide (Phillips 1854). Later versions of this book would tweak their text on the mammals to provide short, but often historically important, insights into their composition and display. So lacking is the documentation of the mammals that we're sometimes reliant on the throwaway text in these guides to tell us how many sculptures were originally installed!

From a palaeoartistic perspective, a clear distinction between Hawkins’ task with the mammal reconstructions and his more famous reptilian efforts was the availability of anatomical information. Mammalian palaeontology was considerably more advanced than studies of fossil reptiles in the early 1800s. Complete skeletons had been known for several of the Crystal Palace species for several decades, allowing scholars to describe, illustrate and restore the osteology of these animals in detail. Hawkins surely benefitted from Owen being an authority on the anatomy of mammals (e.g. Owen 1846), including the Crystal Palace species, and probably also made use of several pioneering skeletal reconstructions, muscle studies and life restorations published by Georges Cuvier. Neglected and somewhat forgotten as they are, the Crystal Palace mammals are actually pretty good takes on the form of their subject species, and clearly demonstrate Hawkins as the equal of later palaeoart masters.

Palaeotherium

The surviving Palaeotherium sculptures in their original site at Crystal Palace, photographed in 2013. These models were temporarily moved for a period in the mid-20th century, which may explain the damage to the sitting model and loss of a third, larger sculpture.
The Eocene equoid Palaeotherium was one of the first discovered fossil mammals and was studied in detail by Georges Cuvier during the early 1800s. Its entire osteology was understood from more or less the moment it was found thanks to near-complete skeletons being recovered from French gypsum deposits at the turn of the 19th century. These brought several Palaeotherium species to the attention of early palaeontologists and led to it being the subject species for some of the oldest palaeoartworks. Both its skeleton and body outline were restored by Cuvier and artists in his employ in the early 1800s (Rudwick 1992, 1997) and these images - after some initial hesitation from Cuvier - were eventually widely published in European literature. With so much data available, Hawkins probably had little difficulty restoring Palaeotherium in three dimensions for the Geological Court.

The Crystal Palace Palaeotherium have an unfortunate history. Two models survive today but photographs from 1958 (see McCarthy and Gilbert 1994), 19th century illustrations, and later editions of Crystal Palace Park Guide (Anon. 1871) indicate that a third model once existed. It was clearly larger and anatomically distinct from the surviving models, but at present no-one seems to know what happened to it - a most regrettable circumstance. It may have been relocated or destroyed when the Tertiary Island site was taken over with a small zoo in the 1950s (we know that parts of the zoo directly encroached into the space for the models - the base of the Megatherium was part of a goat enclosure, for example (see McCarthy and Gilbert 1994)) or else when the smaller mammal models were temporarily moved in the post-war period (Doyle and Robinson 1993). I hope it hasn't been destroyed and may still turn up in some neglected part of the park or emerge from someone's garage.

Two of the three Palaeotherium models photographed in 1958, printed by McCarthy and Gilbert (1994). The standing model shown here is remarkably different in form and size from the surviving Palaeotherium sculptures and almost certainly represents a different species (P. magnum?). Its whereabouts is unknown today.
The surviving Palaeotherium have not escaped misfortune either. The sitting sculpture lost its head at some point in the late 20th century and has been fitted with a replacement, but photographs show that the original head was quite different to the one it has now (compare image above with that below). The replacement is probably a replica or cast from the other surviving Palaeotherium. Both heads are very similar in the snout, ear and eye region, and the differences - the abbreviated cheek and braincase in the sitting statue - are likely results of marrying the head of the standing animal to a sitting one. The neck has also evidently been lengthened since the 1950s and the head/neck join lacks the well-executed muscle contours characteristic of Hawkins' work. It is not the only example of strange, somewhat crude, restoration work on the Crystal Palace mammals (see below).

The sitting Palaeotherium as it appears today - note the different head to the photo from 1958 above, and the slightly awkward manner in which the head replicated from the standing animal has been grafted to the neck.
Questions about lost models and restoration work are not the only uncertainties about these models. I’m not aware of any literature that identifies the Palaeotherium sculptures beyond generic level, but my assumption is that at least one of the surviving models represents P. minus. This sheep-sized species was well described and illustrated by Cuvier and others in the early 1800s, providing Hawkins with ample reference material. I'm uncertain whether both existing sculptures represent P. minus given their historic differences in head shape and other anatomies, but each was clearly distinct from the missing third model, which was significantly larger and of contrasting form. From photographs and illustrations I estimate that the missing model was about the size of a small horse, and this almost certainly labels it as P. magnum, another species that was well illustrated in literature of the early 1800s.

I see you, cryptic P. magnum, hiding in plain sight within P. H. Delamotte's 1853 illustration of Hawkins' workshop. I've long wondered what this sculpture was given that it didn't quite fit anything on display at Crystal Palace, but it's a perfect match for the missing P. magnum model - note the concave back, upright head, straight forelimbs and Gonzo-esque nose. Image modified from Wikipedia.
It’s difficult to evaluate the Palaeotherium models against the science of their day because of modifications made since their installation. Scholars of the early 19th century regarded Palaeotherium as a tapir-like animal with a short proboscis. Cuvier went as far as to suggest that some Palaeotherium species would, should we see them alive, be virtually indistinguishable from modern tapirs (Rudwick 1997). Hawkins evidently followed this suggestion with his smaller standing model, giving it a long, tapir-like face, an arched back, a podgy, creased torso recalling the Malayan tapir, and short, round ears. He opted to give the feet a more horse-like appearance however, which is appropriate to Palaeotherium limb anatomy. The sitting Palaeotherium model also has a tapir-like body, but it lacks the obvious creases of the other surviving model. Perhaps they are, indeed, meant to be different species. The previous head of this sculpture was certainly very different in being much shorter and smaller, and doesn't bear a strong resemblance to any living animal.

Stranger still is the lost model, which was far removed from a tapir-like form except for its short trunk. This large sculpture rather recalls African bush elephants, including a concave back, wrinkled skin, stocky limbs, a deep, short face, prominent brow and conspicuous orbital margins. The skull of P. magnum was not entirely known in the early 1800s and Hawkins may have taken this as an opportunity to be creative with the facial form of the larger model. His apparent referencing of elephants may seem unusual but, even though Palaeotherium was regarded as being related to horses even in the early 1800s, it was also considered a member of Pachydermata. Today, the term ‘pachyderm’ is best known as being an obsolete taxon for elephants, rhinoceros and hippos, but in the early 1800s it included many hoofed mammals too. Under this classification, it might not have seemed much of a stretch to include some elephantine anatomy in a P. magnum restoration.

Palaeotherium magnum as we might reconstruct it today: essentially a robust, compact horse.
Hawkins’ surviving take on Palaeotherium - muddied as they’ve been by time - are not too far off how we regard this creature today: a browsing hoofed herbivore that must have looked something like a tapir or small horse. The now-lost short faces of the sitting and large sculpture are admittedly peculiar as all Palaeotherium have long, somewhat horse-like skulls (Rémy 1992), a fact well established by the 1850s. The introduction of elephant features into the P. magnum reconstruction is, of course, questionable. Elephants are now considered very distant relatives of hoofed mammals and there is no reason to think that they are a good soft-tissue analogy for Palaeotherium. The depiction of trunks is also probably erroneous. Short trunks have evolved repeatedly in perissodactyls and can be predicted for fossil species through a range of bony correlates (Wall 1980). Palaeotherium bears features indicating a particularly fleshy set of lips, but it lacks the full suite of features we associate with having a proboscis.

Anoplotherium

A parade of Anoplotherium commue hanging out at the water's edge in 2013. These remain, a few details aside, pretty darned good takes on Anoplotherium anatomy. Note the impression of musculature in the tails: even though they're hanging low, they look powerful and mobile.
Like Palaeotherium, Anoplotherium was an early subject of palaeoartistic reconstruction at the hands of Georges Cuvier. Two incomplete skeletons of this peculiar hoofed mammal were recovered from Eocene gypsum deposits adjacent to Paris at the turn of the 19th century and, with these, Cuvier was able to reconstruct most of its osteology in a series of papers published from 1804 to 1825 (see Rudwick 1997 and Hooker 2007 for discussion and references). Cuvier's skeletal reconstructions and basic life restorations of Anoplotherium were widely reproduced and would have been well-known among Victorian scholars. Cuvier privately developed muscle studies based on the same illustrations but did not publish them out of concern that they were too speculative for scientists of the early 19th century (Rudwick 1992, 1997). Cuvier was clearly ahead of his time in this regard, foreshadowing a practice that would become important to studies of functional morphology as well as an essential part of the palaeoartistic reconstruction processes.

Cuvier is thus very much the architect of the Crystal Palace Anoplotherium, and Hawkins followed his vision fairly faithfully. He deviated by giving the Anoplotherium camel-like facial details, including large lips, small, rounded ears and a sloping skull roof. Referencing camel anatomy was not Hawkins’ whimsy but informed by early ideas of where Anoplotherium sat in mammalian systematics. This depiction was a departure from scientific credibility however, and Cuvier’s take, with its lower snout and modest lip tissues, was more in keeping with the underlying skull and inferred soft-tissue anatomy of Anoplotherium. This seems to be another example of Hawkins transferring anatomy from living species rather than, as he often did, reconstructing it objectively from fossil bones. Another error is the reconstruction of four toes on each foot. Anoplotherium feet actually had three toes each: two hoofed main digits, and single, somewhat opposable ‘thumbs’ on the inside of each limb (Hooker 2007). Cuvier was aware of there being three digits on the forelimbs at least, and it’s possible that Hawkins added more toes because he thought the fossils were incomplete or otherwise somehow anomalous. After all, Anoplotherium is meant to be an even-toed hoofed mammal, and even today it's an oddball for its unusual toe counts. But other than these relatively minor errors, the Anoplotherium sculptures are compelling reconstructions that are still used to illustrate the form of this taxon today (e.g. Prothero 2017). I particularly like the strong, flexible-looking tails and the form of their muscular torsos.

Anoplotherium commune, the middle portion of a Venn diagram containing Bambi, Lassie and Rory Calhoun, in its characteristic feeding posture. Our anatomical interpretation of this animal is very similar to how Cuvier and Hawkins reconstructed it, but we have different ideas about its lifestyle.
Hawkins created three Anoplotherium sculptures: one standing, one resting, and one in a curious half-crouched pose with an outstretched neck and head*. I’m not entirely sure what behaviour the latter is meant to depict. In the early 1800s Anoplotherium was regarded as a swimming animal that used its powerful tail to propel itself through water, perhaps like an otter or coypu (e.g. Owen 1846). Maybe the third animal is meant to be shaking itself, dog-style, to dry off as it emerges from the water surrounding the Tertiary Island? The soft-tissues of the neck are inconsistent with the other models, and this is not the result of damage or poor conservation. Might it represent deformation of the skin as the neck is shaken about? Today, Anoplotherium is interpreted as a fully terrestrial animal adapted for high browsing (Hooker 2007). Peculiarities of its pelvis are shared with mammals that regularly stand upright on two legs, and it’s probable that Anoplotherium adopted this pose to browse above the feeding envelope of contemporary mammals (Hooker 2007). The strong tail, in this hypothesis, becomes a stabilising organ rather than a swimming aid.

*The 1854 Routledge's Guide to the Crystal Palace and Park at Sydenham suggests there are meant to be four Anoplotherium, but I'm not aware of any other documents indicating this. Is there another missing model, or was this a typo?

"Anoplotherium" gracilis - or, more appropriately, Xiphodon gracilis. Some authors suggest that some of the Crystal Palace Anoplotherium sculptures represent this species, but I strongly doubt this. gracilis has a completely different shape to commune and this was understood early in the 1800s. This image is by Georges Cuvier, and it pre-dates the Crystal Palace project by several decades. From Rudwick (1997).
According to McCarthy and Gilbert (1994) and Doyle and Robinson (1993), two species of Anoplotherium are represented at Crystal Palace: the two standing individuals are A. commune, and the reclining sculpture is “A”. gracilis. It’s not clear to me that this is accurate, however. Firstly, by the time Crystal Palace Park opened A. gracilis was well-differentiated taxonomically from A. commune. Cuvier placed gracilis in a subgenus, Xiphodon, in 1822, and this was erected to a full genus by M. Paul Gervais in 1845. Owen agreed with this change and stopped referring to “Anoplotherium gracilis” at some point between the mid-1840s and mid-1850s (see Owen 1846, 1856, 1857). By the time the Crystal Palace sculptures were being commissioned the disassociation between Anoplotherium and X. gracilis was thus well established, and we have to assume that Hawkins and Owen were aware of it. A complication to this is that Hawkins still referred to A. gracilis in the early 1860s (judging by the labelling on one of his 1862 posters), but this brings us to our second point: that Hawkins evidently knew how different gracilis and commune were anatomically. His 1862 posters show commune as reconstructed at Crystal Palace while his gracilis are the long-legged, long-necked, and short-tailed creatures of Cuvier and other artists in the early- and mid-1800s (see Rudwick 1997 for Cuvier’s own detailed accounts on the anatomy of this species). This isn’t surprising: in the early 19th century commune and gracilis were regular fixtures in palaeontological texts, and skilled, intelligent artists like Hawkins would not readily confuse them. This is not to say that claims of gracilis being featured at Crystal Palace are baseless, but they neither marry up with the Anoplotherium statues we have today nor the history of Anoplotherium research. As with the uncertainty about the taxonomic representation of the Palaeotherium sculptures, the poor records and deficit of historic interest in the Geological Court mammals do little to help resolve this confusion.

Megatherium

Towering above surrounding vegetation is Hawkins' Megatherium americanum, shown here as it was in 2013. There's always some vegetation obscuring this sculpture and, last time I visited the park, it was near impossible to see Megatherium at all. This obscures, among other things, the expertly sculpted feet, legs and tail. Controversy still reigned about the foot posture of ground sloths in the 1850s, but Hawkins was spot on in his depiction.
The research history of Megatherium americanum began nearly six decades before Hawkins commenced work on his model. Despite this long research lead, Hawkins’ rearing, tree-grasping Megatherium was ultra-progressive for the time. It was one of the first depictions of Megatherium in a pose that chimes with our modern understanding of giant sloth habits and actually pre-dated publication of ideas that it was capable of such feats. Megatherium was, for much of the early 1800s, regarded as Georges Cuvier imagined it in the latest 1700s: a flat-footed, trunked quadruped with particularly dextrous forelimbs (Rudwick 2005, Argot 2008). Even into the 1850s scholars were still confused over aspects of how this animal lived, with authors like François Jules Pictet-De la Rive writing long discussions about its capacity for burrowing, climbing, and harvesting vegetation. Owen's studies on another species, Mylodon darwinii, elucidated many aspects of ground sloth lifestyle and anatomy that would inform Hawkins model. Owen showed that Mylodon walked on the sides of its feet and the shorter, clawless fingers of its hands. He viewed giant sloths as browsers and tree fellers based on numerous lines of anatomical evidence (Owen 1842) and, under his direction, a Mylodon skeleton was restored as a tree-rearing biped at the Hunterian Museum in the late 1830s/early 1840s. In contrast, contemporary European mounts of Megatherium and illustrations of its skeleton retained fully quadrupedal, flat-footed stances.

The Crystal Palace Megatherium being manufactured in an image published by Die Gartenlaube in 1853. Note the size of the hands, which are much larger than the surviving hand on the model today. From Wikipedia.
Owen eventually wrote at length about tree-rearing giant sloths when discussing the anatomy and habits of Megatherium, but only well after the Crystal Palace models were completed. An article in the German magazine Die Gartenlaube shows that Hawkins’ Megatherium model was completed in 1853, a year that could - if Hawkins had been strictly following available Megatherium reconstructions - have seen the Crystal Palace ground sloth restored as a Cuvierian quadruped. The fact that Hawkins avoided this implies that he either combined Owen’s ideas on Mylodon with the anatomy of Megatherium, or else that Owen tipped him off about the direction of his future research. Either way, the portrayal of an elephant-sized mammal rearing into a tree has to be regarded as extremely progressive for the 1850s. Other early 19th century scholars assumed such animals were confined to quadrupedal poses (Argot 2008) and, while it was not a stretch to imagine the bear-sized Mylodon routinely rearing onto its back legs, it was a bold prediction to portray an enormous Megatherium doing the same. As demonstrated in many Crystal Palace models, Victorian scholars often imagined fossil animals as variants on recognisable modern forms - paleotheres as tapirs, dicynodonts as turtles, pterosaurs as birds - but Hawkins’ Megatherium was boldly different from anything alive today, and foreshadowed the way we would start visualising prehistoric species as our science and data improved.

The Crystal Palace Megatherium is of further note for being one of the oldest life restorations of a ground sloth. Though several skeletal reconstructions were published prior to the 1850s, few, if any, restorations with a complete suite of soft-tissues were attempted. (It’s curious that none have been found among Cuvier’s archives, given his links to Megatherium and his habit of restoring the anatomy of fossil mammals). Despite its vintage, Hawkins’ Megatherium has held up well as a portrayal of ground sloth form. A commendable portrayal of proportion and musculature is buried under layers of long, shaggy hair, with the muscular, relatively slender shoulders contrasting appropriately against the wide and robust pelvic region. The feet are appropriately inturned and the arms are depicted as if grasping a tree to access vegetation or push it over, entirely in accordance with Owen’s interpretations of sloth behaviour. Hawkins’ depiction of shaggy hair anticipated the discovery of giant sloth hair by almost half a century (Woodward and Moreno 1899) and, although it remains unclear whether Megatherium itself was covered in such fur, this take is certainly consistent with the fossil skin of several giant sloth species.

Megatherium americanum as we know it today - really not so different from how Hawkins envisaged it 165 years ago.
Two major difference between Hawkins’ Megatherium and our modern reconstructions are obvious. The first is the presence of a short proboscis, a Cuvierian interpretation also endorsed by Owen (1842). Hawkins’ restoration would have pleased Victorian scientists, but trunked sloths have not withstood modern scrutiny. Today, it is instead thought that ground sloths had extensive nasal cartilage and prominent lip tissues (Bargo et al. 2006), but they lack features indicative of trunks or proboscides.

It's hard to see the Megatherium sculpture in full, especially from anterior view, so I've borrowed this photo from the Friends of Crystal Palace Dinosaurs website. Note the excellent rendering of the crouching, pedolateral hindlimbs, and also the strangely small left hand (the right is missing). You can see the colour mismatch between the left forearm and elbow - this marks a site of repair where the tree outgrew the grip of the sculpture. But what's with that tiny hand? Its detailing and grafting onto the forelimb is weird and doesn't match illustrations of the Megatherium sculpture from the 1850s (above). It's surely a crudely-sculpted replacement, not a replica of the original.
The hands (or rather, the hand - the right seems to be missing at present) are the second anomaly, and are far less explicable. As represented today, the surviving hand is curiously undersized, lacks claws and is so poorly shaped that I initially assumed it was a post-Hawkins replacement, akin to the replaced Palaeotherium head. The left hand was replaced after the growing girth of the tree broke the sculpture’s left forearm, and I presumed a diminutive hand was added due to lack of space. But no, at least according to McCarthy and Gilbert (1994), the Megatherium hand currently on the model is a replica of the real deal. This seems peculiar, and I'm not sure I buy it. Hawkins’ illustrations of Megatherium from the 1850s and 60s (including artwork associated with Crystal Palace) show appropriately large, clawed hands, and an illustration of the model being constructed in 1853 (above) shows it grasping a tree with sizeable, robust extremities. The rest of the model is so exact to Megatherium form that the embarrassingly inaccurate hands are entirely out of place - they look like they were made by someone who had no idea about Megatherium anatomy, which is patently not the case for the rest of the model. I strongly suspect that the hands of the model are crude replacements of lost originals, and that there’s a missing chapter in the history of this model.

Megaloceros, the Irish elk

The Crystal Palace Megaloceros bucks and doe on display in 2018. The attention to detail on these models is superb and, today, their situation close to pathways around the Geological Court allows visitors to get extremely close.
Probably the most spectacular mammal sculptures at Crystal Palace Park are the four Megaloceros giganteus situated in at the northeastern extent of the Geological Court. A reposed doe and fawn feature alongside two large bucks, each standing in a classic ‘regal’ pose with antlers aloft. So imposing are these sculptures that they would not look out of place situated among grand governmental buildings, or atop an enormous plinth in a city square. It seems strange that no iterations of the 19th century Crystal Palace Park Guides suggested starting tours of the Geological Court with this display. The Megaloceros, after all, slowly leads us into the strangeness of extinct animals ("they're deer, Jim, but not as we know them") as well as demonstrates Hawkins’ ability to create believable animals (a fact far easier to deduce with a deer than a dicynodont). Beginning a tour from the other end of the court, with the far less impactful and more distant Dicynodon, robs us of this effect. I also feel that the grandeur and strangeness of Victorian dinosaurs, marine reptiles and giant sloths overshadows Megaloceros somewhat. Sure, it's big and the antlers are impressive, but its "wow" factor is diminished after meeting the stranger, larger reptiles situated a few hundred metres away. I can understand why Hawkins was pushing for additional, less-familiar species - mammoths, dodos, moas - to place around this end of the Court before his funding was pulled.

Hawkins had no concern for missing anatomy or predicting proportions with Megaloceros. Decades before the Crystal Palace project was even conceived, Megaloceros osteology was extensively described and illustrated. From Cuvier (1827).
Megaloceros was a historic fossil species even to Hawkins and Owen. Remains of this animal were found in the late 1600s and, by the early 1800s, enough material was known to reconstruct the entire skeleton. Cuvier (1827) published several such illustrations, including two skeletal reconstructions that were widely reproduced in later texts on fossil mammals. This, and the glut of Megaloceros material held by British museums, would have given Hawkins an excellent insight into its anatomy. Visitors to Crystal Palace would not have known the Irish Elk as Megaloceros giganteus, however, but under Owen’s 1844 name for the species, Cervus (Megaceros) hibernicus. The nomenclatural history of M. giganteus is confused by several names being coined for this species in the 18th and 19th centuries. Owen’s subgenus (eventually promoted to a ‘true’ genus) Megaceros was the first to enter widespread use and almost became the accepted generic name for giganteus. However, Megaloceros was resurrected in 1945 (albeit somewhat corrupted from its original spelling, Megalocerus) and both names were applied to the Irish Elk until the 1980s. Adrian Lister (1987) finally brought an end to decades of confusion by establishing Megaloceros giganteus as the most appropriate name on grounds of both nomenclatural priority and usage.

Hawkins' Megaloceros doe and fawn, as seen in 2013. I really enjoy the detailing on their feet - if there was any doubt that Hawkins could make realistic-looking familiar creatures as well as weird fossil ones, these models dispel it.
Megaloceros was surely the least demanding of Hawkins' reconstruction assignments because of its close relationship to living deer. As did Cuvier, Owen realised that early interpretations of Megaloceros as a giant moose-like cervid were incorrect, and he placed it among Cervus, a genus that includes several other large, Old World deer species (e.g. Owen 1844). Hawkins appears to have referenced several Cervus anatomies in his reconstructions, especially the thick neck manes, deep fur over the withers, and a line of long, shaggy fur along their bellies. These are especially obvious on the bucks, but also present on the reclined doe. Manes are not common to many deer females, and I suspect Hawkins was referencing the winter appearance of certain elk subspecies (‘elk’ as in the wapiti Cervus canadensis, not the Eurasian elk/moose). The short, blunt tails seem to agree with this interpretation too. It would later be traditional to reconstruct Megaloceros like the red deer Cervus elaphus, but I wonder if Hawkins thought the shaggy appearance of winter elk was more apt for an Ice Age animal, or else if he thought the longer fur would look more obvious on his sculptures.

Hawkins' reconstructions of Megaloceros are, of course, some of the most scientifically credible of all his Crystal Palace artworks. But I have to admit that, on grounds that their restoration was nowhere near as complex as the other sculptures, I don’t think they’re the best examples of his palaeoartistic abilities. They certainly leave little doubt that Hawkins could produce convincing portrayals of semi-recognisable animals, but Megaloceros probably wasn’t much of a stretch for him. His artistic expertise included illustrating living mammals and he eventually wrote a series of books on this very topic (including one featuring deer, in 1876). For me, it’s his deductions about lesser-known and wholly unfamiliar fossil species that place him among the old masters of palaeoart, even though these insights are associated with sculptures that are scientifically more dated.

Today, we imagine Megaloceros almost as Hawkins did, excepting some different ideas about their colouration and soft-tissue anatomy. These have been provided by cave art and revelations about their relationships with modern deer.
While Hawkins’ Megaloceros are impressive reconstructions, they differ from our considerations of this animal today. It seems that Megaloceros was more closely related to fallow deer Dama than Cervus, and this implies some differences in facial anatomy and colouration, as well as some particulars of fur and soft-tissue distribution (e.g. a bulging laryngeal region and brush-like genital sheath). Some of these anatomies are confirmed in Megaloceros cave art (Geist 1999; Guthrie 2005), which also records other details unknown from fossils. These include a shoulder hump (presumably long hairs, fat or both) and a series of dark stripes: one at the base of the neck, one running from the shoulder towards the knee, and another surrounding a pale rump. Cave art also suggests, though not conclusively, that the head and neck were pale or white, while the body was darker, perhaps light brown. (There's a terrific summary of Megaloceros life appearance over at Tetrapod Zoology - check that out for additional details and discussion). This information was entirely unknowable to Hawkins, however, as the discovery of ancient European cave art post-dated the Crystal Palace project by over a decade, and its acceptance as the work of authentic Palaeolithic humans, and not modern vandals, was even longer coming. Moreover, even once ancient cave art was accepted as a genuine part of European history in the early 20th century, it would take decades to discover enough Megaloceros cave paintings to deduce meaningful details of its anatomy and colouration.

So, about those Crystal Palace Dinosaurs...

That brings our palaeoartistic review of the Crystal Palace palaeoartworks to a close, kudos to anyone who's read the entire series. This was meant to be little more than a series of brief notes and it's ended up being a number of long articles. Having already been interested in the Crystal Palace sculptures before writing this, I must admit to having a true fascination with them now. Writing these pieces has revealed so many gaps in our understanding of their history and development, allowed me to appreciate just what a good artist Hawkins was, and driven home the fact that the Geological Court sculptures were really not an Owen-Hawkins collaboration, but almost a solo Hawkins project. The latter point is already well made by records of correspondence between Owen and those involved in the Crystal Palace project, but trying to 'reverse engineer' Hawkins' artwork further demonstrates the token involvement Owen must have had. In this light, Hawkins really needs to be discussed more widely as one of the all-time greats of palaeoart. Despite relatively little scientific assistance, he produced spectacular, realistic and charismatic takes on fossil animals at a time when our understanding of animal anatomy - both fossil and modern - was a fraction of what it is today. Yes, he got many things 'wrong' with respect to our modern understanding and he perhaps leaned on living animals more than we would nowadays, but to focus on this, and not his achievements in anatomical prediction, his knowledge of contemporary science and attention to anatomical detail, does him a disservice. We have to evaluate the Geological Court models in light of what was known at the time, and in this respect they are truly first rate. Far superior in science and art, in fact, to the vast majority of palaeontological sculptures exhibited today.

Furthermore, I feel more committed than ever to the fact that these models should, no, must be conserved for future generations. As globally unique monuments to Victorian science and culture, we should regard them with pride, reverence and admiration, and not allow them to deteriorate through neglect, underfunding and (sad to say) deliberate vandalism. The ongoing work by the Friends of Crystal Palace Dinosaurs is essential to this mission and I salute them for pushing the value of these sculptures against the odds, and for their successes so far. As I've said repeatedly throughout this series, if you share my interest and concern for Hawkins' Crystal Palace palaeoartworks then check out the FOCPD website you see how you can help: chip in some money to help conserve the models or provide some elbow grease to help maintain the Geological Court. If we let these models slide too far into disrepair there's no coming back for them: all their artistry, history and scientific significance will be gone for good. Please take interest and help out if you can.

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References

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