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Our knowledge of how extinct species looked is changing all the time, as is the art that depicts them.
Known as palaeoart, the artform is linked closely to the latest scientific research on dinosaurs, pterosaurs and other forms of extinct life. But how has palaeoart changed over time? And how does it affect science?
Extinction is a natural part of life. But that doesn't mean we have to forget what extinct species look like.
Whether in cave paintings or films, humans have been depicting wildlife for tens of thousands of years. When the artist makes use of scientific information to draw an extinct species, it is known as palaeoart.
Alongside fossils, this art helps bring us closer to the extinct species of the past, as palaeontologist Dr Darren Naish explains.
'Fossils are, for many people, remote objects,' Darren says. 'While museums have gone to some trouble to make them as accessible as possible, there is still a stigma that they are dusty things hidden in basements.
'Emphasising that these are the remains of extinct organisms, which would have been exciting, beautiful and dynamic, is the function of palaeoart. Through it, fossils can be made to seem more real, more accessible and more attractive to everyone.'
While dinosaurs are often the most popular subjects, palaeoart can portray any animal, plant or ecosystem that no longer exists. The term was coined by palaeoartist Mark Hallett in the late 1980s to refer to his illustrations of extinct animals, and has since been widely accepted for a range of different styles of artwork.
These can range from drawings attempting to reconstruct a section or body part of the animal to depictions of how the organism could have looked while it was alive, including showing any potential behaviours.
These artworks are often commissioned by scientists to illustrate their latest fossil discoveries to help explain them to a general audience.
Dr Susannah Maidment, a palaeontologist at the Museum, says, 'Primarily, we use palaeoart as a way of communicating our work to the general public.
'However, there are elements of it that are used slightly more in science now. For instance, while biomechanical models don't need bones, a sculpted figure helps other scientists understand the concepts more quickly and illustrate the processes involved.'
Given palaeoart's close links to active research topics, depictions of how extinct species may have looked need constant updating. This means that older palaeoart often becomes outdated as new discoveries are made.
Museum palaeontologist Prof Paul Barrett says, 'Out of date palaeoart is useful as a record of what people used to think, so it's a very quick way of getting an idea of change.
'When they're made, they help convey an idea about what we thought the animals did or looked like at the time, and now they are a way of showing how these ideas have progressed.
'This means they have just as much educational value now as they did then, but not as a way of communicating the most current science.'
Before there was palaeoart, there were illustrations of fossils. Some of these depictions can date back centuries, with 600-year-old rock art believed to depict dinosaur footprints known from Flag Point in Utah, USA.
The tracks, and subsequent art, are associated with the legend of the thunderbird in several Indigenous peoples across North America. Mythology also played a role in Gottfried Leibniz's depiction of disparate fossils as the remains of a unicorn in the 1660s.
Mammoths tend to feature heavily among early illustrations, as their well-preserved remains were drawn by individuals such as Philipp Johann Tabbert von Strahlenberg and Roman Boltunov. As international travel and communication became more common in the 17th and 18th centuries, depictions began to circulate more widely.
As scientific illustration became more common, some artists took went beyond depictions of specimens and made the first attempts at soft tissue reconstructions. These artworks are considered amongst the earliest examples of palaeoart.
However, the reconstructions were generally based on scientific guesswork, and though some are still recognisable today, such as Johann Hermann's illustration of Pterodactylus antiquus, others are significantly different from our modern understanding, such as George Scharf and Gideon Mantell's Iguanodon watercolour.
A more ambitious example of palaeoart from the 1800s is the painting Duria Antiquior, by geologist Henry De la Beche. Depicting plesiosaurs, ichthyosaurs and pterosaurs, he used fossil evidence to depict the marine animals of Jurassic Dorset, based on finds from Lyme Regis, in an ecosystem for the first time.
'Duria Antiquior was produced to raise money for Mary Anning,' Darren says, 'and is one of the earliest attempts to produce accurate depictions of extinct animals.
'At the time, artists were trying hard to fit the animals into an environment, using living animals as inspiration. After thinking of memorable scenes with modern wildlife, they were trying to recapture that with prehistoric animals.'
One such artist was Benjamin Waterhouse Hawkins, who spent many years at Knowsley Park drawing the menagerie of the Earl of Derby. He used this experience, as well as consulting with the leading scientists of the day, to recreate extinct animals from the UK and USA.
Some of the most famous of his reconstructions are the Crystal Palace dinosaurs.
The increasing availability of palaeoart in museums and other public places set the stage for its prominence at the beginning of the 20th century.
In the late 1800s and early 1900s, palaeoart was increasingly found in museums and publications around the world. Some of these images helped inspire a young Charles Knight, who is now acknowledged as one of the fathers of modern palaeoart.
He is renowned for his large murals of dinosaurs in museums such as the Field Museum and the American Museum of Natural History. These were some of the first images to show dinosaurs, such as Laelaps, as fast-moving, dynamic animals rather than sluggish, slow-moving beasts.
It wouldn't be until decades later that scientists would begin to re-evaluate how active dinosaurs actually were.
But while Charles Knight's palaeoart likely helped to set the scene, his depictions weren't always accurate, with muscles and proportions unlike those that the fossils suggested. Despite this, many of these drawings have since become iconic, even if their message isn't entirely helpful.
'There are occasions, even to this day, where palaeoart can drive the scientific agenda because people become used to seeing reconstructions of an organism in a certain way, and those become proxies for what the animal looked like,' Paul says.
'If certain elements of a reconstruction keep coming up, it creates an unconscious bias, even in professional palaeontologists, as to what a species may have looked like without considering all the evidence.
'I think this is one of the slight negative connotations of palaeoart, because it's pervasive and engaging but doesn’t always have the same outlook as the scientific consensus.'
The work of Knight was also influential in the early film industry, with the dinosaurs of 1919's The Ghost of Slumber Mountain based on Knight's reconstructions. This helped pave the way for dinosaurs to become big business in blockbusters such as 1925's The Lost World and 1933's King Kong.
These films, alongside the work of emerging artists such as Zdeněk Burian and others, helped drive public interest in palaeontology even as scientific interest was waning.
It also helped to drive changes in society. For instance, Alice B. Woodward was a pioneering illustrator whose scientific drawings and life reconstructions were highly sought after at a time when women were only just starting to break through into the male-dominated scientific community.
While artists had previously been generalists, with specific palaeoart commissions from patrons such as museums, over the coming decades the rising popularity of extinct organisms, especially dinosaurs, would see the first full-time palaeoartists emerge.
Professional palaeoartists began to become more common during the dinosaur renaissance of the 1960s and 1970s. During this time, the ancient reptiles were reassessed as being much more active than thought, while the idea they evolved from birds began gaining acceptance.
Changes in scientific thought led to changes in palaeoart. Dr Robert Bakker, who helped promote the idea of active dinosaurs, produced a variety of artworks showing dinosaurs running, jumping and hunting to make his point, which inspired other artists like Gregory Paul and Doug Henderson.
This more science-focused artwork meant that more research than ever before was going into palaeoart, as palaeoartist Bob Nicholls explains.
'Science is everything,' Bob says. 'Palaeoart is an odd artform because when you start a new artwork the aim is to learn as much as you can about your subject to minimise any opportunity for self-expression, which is the opposite of other forms of art.
'I want to minimise the number of knowledge gaps in my work, and when I do come across them, I use informed speculation through comparisons with relatives and other scientific techniques to come to the most plausible conclusion.
'The place where an artist has the opportunity to express themselves is in the composition, such as the viewpoint, the texture and the style, and different palaeoartists would all conclude the same project with different interpretations.'
This increasing role of research in palaeoart saw it move away from portraits of particular organisms on a white background, and from impressionistic scenes featuring volcanoes and other 'prehistoric' backgrounds.
'In the old days, for dinosaurs in particular, the standard palaeoart reconstruction is an animal standing in a bare foreground with a few trees dotted around in the distance, and maybe a smoking volcano,' Paul says.
'In these images, the emphasis is very much on the animal. During the dinosaur renaissance, there’s been a move away from this paradigm, to thinking about the other animals and plants that would have been a part of the species' ecosystem.'
This has helped draw attention to topics such as fossil plants, which are harder to reconstruct from fossils and can often be overlooked.
'It's really difficult to get information on how the world looked and hone it,' Darren says. 'There's always been demand for fossil animals, but getting the specialist information to depict environments is very difficult. The information to reconstruct plants or more obscure animals in these scenes may simply not exist.
'The available information is improving over time, but it's common for leading palaeoartists to devote weeks, months or years of research to find out about plants, landscapes, and potentially even weather systems in their work.'
Part of these changes are the result of new technology and digital connectivity, which is providing another revolution in the artform.
With the age of the internet, palaeoartists now have access to more information than ever before, allowing them to achieve greater levels of accuracy.
'When I started out, you had to find the information you wanted in books, and for topics such as fossil plants, there weren't a lot out there,' Bob says. 'It's a lot easier now that you can use the internet to find sources for a particular species, and it also allows you to compare these sources to decide upon the most convincing interpretation.'
This has opened palaeoart up to a new generation, who have made it their mission to recreate as many extinct species as possible.
'We're probably not too far off the stage now that if you think of a genus level fossil animal, there is probably a reconstruction somewhere online,' Darren says. 'I think it's helpful not just for palaeontology, but for humanity as a whole, because it's increasing the available imagery for extinct species which we didn't have that beforehand.'
Professional palaeoartists have also changed how they work as new technology becomes available.
'I've been working as a palaeoartist for 20 years, and for the first 10 years I worked almost entirely traditionally using paints, pencils and other materials,' Bob says. 'But for the past decade my work has been almost entirely digital.
'I can change compositions really quickly by moving components around, so making a living from palaeoart is much easier. I'm also able to share my work on social media and connect much more easily with clients, rather than having to physically post samples.
'It's completely changed the game and it is a huge advantage I didn't have when I started out.'
Palaeoart itself is also entering a new era, as scientists and palaeoartists reconsider and challenge preconceptions of different prehistoric animals.
In All Yesterdays, which Darren co-authored in 2012, a variety of extinct animals are depicted in often overlooked contexts, while showing how modern animals could be reconstructed inaccurately by using the same ideas applied to extinct species.
Debate in palaeoart reflects how it can challenge scientists to think more widely about their ideas when they assess their own work.
'I think palaeoart, in some ways, challenges researchers to think about what we know and what we don't,' Susannah says. 'For instance, there are many dinosaurs with long neural spines, and when I'm describing a species, I don't always consider what the function of this might be.
'It might mean they had a hump or a sail, but palaeoart's emphasis on soft tissue characteristics might help palaeontologists to go back to the bones and look for evidence of this.'
While it is difficult to predict where palaeoart will go next, TV series such as Prehistoric Planet and film franchises such as Jurassic World are investing more heavily in palaeoart than in recent memory. This could lay the foundations for further innovations in the field and bring new ways to bring extinct species to life.