The Alien Prehistoric World Trope: Part 4 - Familiar Beasts?

This is part 4 of a series. If you have not read the previous part, please do here.

Fig. 1: A young Rudolph Zallinger (1946) standing in front of his nearly completed mural, just to give you an idea of its size.

Last time we stayed in the dinosaur-cinema of the late 30s and early 40s, but as these will grow into increasingly stale stop-motion affairs in the following decades we will instead return to the world of artwork. In 1947 a major piece was finally completed after having been worked on for almost 5 years. The Age of Reptiles is a gigantic mural painted on the eastern wall of the Peabody Museum of Natural History’s Great Hall. The museum was first founded in 1866 by George Peabody as a sort of gift for his nephew Othniel Charles Marsh, whom we have met in part 2. In 1925 a new building, the Great Hall, was opened to house the gigantic mounted skeletons which Marsh had uncovered. Many of these still stand there today, including the Apatosaurus/Brontosaurus. In 1942 oceanographer Albert Parr, then director of the museum, noticed a big problem with the hall. The large, grey bones in combination with the large, grey concrete walls made for a rather drab and unappealing look. He wanted to decorate the walls with some paintings that showed the dinosaurs in life restoration. At the time a young man of the name Rudolph Franz Zallinger was illustrating fossil algae for Parr. Zallinger was born in 1919 in Irkutsk, Russia, as the son of an Austrian war-captive and a Polish railway-engineer. When he was still a baby his parents fled with him to the US, where Rudolph would go on to study Fine Arts at Yale University, leading to him getting this job at the Peabody Museum. Lewis York, professor at Zallinger’s art-school, suggested to Parr that Rudy could do the job. Zallinger accepted and suggested that instead of creating many smaller paintings, he should instead make one giant mural across the entire eastern wall that shows the progression of time. His technique of choice was that of a fresco secco (Italian for dry plaster), a type of painting invented by Cennino Cennini in the fifteenth century. It is a form of mural rarely practiced today, as it requires enormous skill and many preparatory steps. Zallinger pulled it off however.

Fig. 2: The Triassic section of The Age of Reptiles. Note the trees in the foreground that separate the scenes. Because of the sheer size of the mural I had to scan a fold-out miniature from a guide-book I own, so I apologize for the low quality.

The end-result was a 4.9 x 34 meter mural showing the evolution of life from the Devonian all the way to the end of the Cretaceous. We begin on the right side with a bunch of lobe-finned fish, Eusthenopteron, crawling their way out of the swamp underneath alien-looking plants. As we progress to the left the world becomes populated by amphibians. Over a fallen tree we pass a across a river into the Permian where we see familiar fin-backed stem-mammals such as Dimetrodon. A large tree in the foreground cuts this scene off from the next one, which is set in the Triassic and shows us the first dinosaurs, such as Plateosaurus, in the familiar tripodal, upright posture. The Triassic scene is again bordered by another tree, which sections off the Jurassic. In the foreground a gigantic Brontosaurus wallows in its river, surrounded by the pterosaur Rhamphorynchus and the proto-bird Archaeopteryx. Behind it a very ungainly-looking Stegosaurus crawls around, while a very pot-bellied Allosaurus feeds off an indeterminate carcass. Another tree caps this period off as we enter the Cretaceous. We see familiar dinosaurs such as Ankylosaurus and Triceratops. Most prominent is of course T. rex, who looks even more pot-bellied and brawny than the allosaur and stands in the classic Godzilla-pose. Under the dinosaurs’ feet the small mammal Cimolestes scuttles around. At the very far left corner we see several volcanoes erupt. This signals off the end of The Age of Reptiles, as at this time volcanism was seen as the most likely explanation for the extinction of the dinosaurs, with the ash in the atmosphere creating an ice age that the giant reptiles could not survive.

Fig. 3: Zallinger’s depiction of Jurassic dinosaurs is definitely not revolutionary for his time in any way, however the quality of the art with all its details is impressive (though my bad scan probably does not accurately convey this) and on a higher level than previous work. It definitely does not have to shy away from Renaissance frescoes.

Like Fantasia, The Age of Reptiles likely only added to the Alien Prehistoric World Trope indirectly through misunderstanding and its wide circulation. A casual viewer not familiar with geologic history or the mural’s background might not understand that the piece depicts multiple time periods and that the large trees in the foreground are used to demarcate leaps in time. They might therefore come to think that the mural depicts just a single age, with all the lieforms living together at the same time. The volcanoes at the left corner, used to symbolize a singular event, might therefore also be understood as a casual occurrence of a prehistoric landscape from this time. This likely added to the already existing preconception that volcanism was high throughout prehistory, which we looked at in part 3. The creatures themselves also look quite bizarre. While preparing for the mural, Zallinger did get a crash course in vertebrate paleontology and paleobotany, but it is clear that he did not have the finest grasp of the dinosaurs’ anatomy. In many ways they resemble their lumpy and bloated counterparts from Fantasia, except their implied movement is much slower, in part because fresco secco does not lend itself well for dynamic scenes. We are reminded of the “huge, bizarre and ungainly shapes rising and subsiding in the landscape” that Charles Knight talked about in Part 2.

Fig. 4: Depicting Cretaceous dinosaurs alongside modern-looking flowering plants became a common trend from the 1940s onward. In hindsight Zallinger ironically depicted the flowers too modern, as the purported crown-group angiosperms depicted in the mural later turned out to be a lot more primitive than thought.

There are however some things that Zallinger improves over Disney and Knight when it comes to the alienation of the prehistoric world. Because of the fresco secco method, the texture and coloration of the dinosaurs looks a lot more lifelike and similar to a living animal’s appearance. While the animals are a bit ungainly in shape, they do not look as cartoonish as they did in Knight’s paintings. The coloration of the environment itself is more reminiscent of a modern landscape than anything in Fantasia. The sky and water are blue and the plants are in a pleasant green. The Age of Reptiles also began the trend of showing Cretaceous dinosaurs alongside something very familiar for us: Flowering plants! Most prominently we see a beautiful magnolia tree alongside Triceratops and T. rex. Behind them grows a willow. In previous paleoart, apart from anachronistic grass, it was convention to show dinosaurs alongside tropical  swamp- or rainforest plants, such as ferns, palms, cycads or ginkgos, all of which are vegetation we Western audiences usually associate with foreign, exotic places. What was often forgotten is that for most of the Mesozoic the dominant plant groups were the conifers, which even back then looked familiar to our modern pine trees and sequoias. Many Jurassic dinosaurs likely lived in environments that would not have been too dissimilar form North American redwood forests. What was also known for the better part of the twentieth century is that angiosperms, the flowering plants, existed at least since the Cretaceous (in recent years some evidence has pushed their possible earliest origin as far back as the Triassic or even the Permian). Zallinger was perhaps among the first to accurately reflect this in a major paleoart piece. The composition, coloration and vegetation all create something which was largely absent from previous depictions of the prehistoric world: Beauty. The thing that Knight thought prehistory entirely lacked. The Age of Reptiles is not a deriding depiction of a ruthless antediluvian world anymore, but rather a glorifying story of the conquest of land by the vertebrates, in the same style and with the same skill as any Renaissance painting. When Peter Dodson first saw the mural in person he allegedly almost wept.

Fig. 5: A lonely, sad 1950s Iguanodon standing in a dinosaur graveyard. It should be mentioned  that Augusta and Burian had to live through both World Wars and saw the beginning of the Cold War with its global threat of nuclear annihilation. Someone standing in a field of bones in this context likely adds at least some personal, perhaps even autobiographical connotations.

14 years before Zallinger another artist was born in the Eastern Bloc, more precisely in what is today the Czech Republic. Unlike Rudy he would however remain in this part of the world, as it would soon thereafter become a member of the Soviet Bloc. His name was Zdeněk Michael František Burian. As a teenager he joined the Academy of Fine Arts in Prague, however he would not complete his studies and instead improve his artwork autodidactically while working rather laborious jobs. After doing his military service (partially working as the division painter) in the 1920s, he became an illustrator for various adventure books, including those of Jules Vernes, Edgar Rice Burroughs and Rudyard Kipling. Some German readers might recognize his art from some of Karl May’s Winnetou novels (to American readers unaware, Winnetou was what we in Central Europe used to think the Wild West was like). In 1935 Burian was discovered by paleontologist Josef Augusta, who asked the painter if he could illustrate extinct lifeforms for him. Paleoart-history was made and the two would work closely together on many popular books about prehistory until Augusta’s death in 1968. Burian would nonetheless continue working with other palaeontologists to produce more art until his death in 1981.

Fig. 6: This 1970 Burian-piece is often called the “Heroic Tarbosaurus”. You can clearly see why. With muscly, eagle-like legs, lifted tail and sharp sight Tarbosaurus bataar proudly dominates the landscape, looking for a new foe to face. Napoleon Crossing the Alps ain't got shit on this. The dinosaur no longer serves as a monstrous villain to the protagonist, rather it is the protagonist. I also just want to say that Burian was great at painting skies. As Clemens J. Setz notes, the literal air in many of his pieces is drawn as if the paintings were nostalgic photographs made with a 1970s-camera (Schalansky 2013, p. 6).

Burian’s work is iconic. If you don’t know the man, you nonetheless probably know his art, as it is widely circulated and its influences can be felt even today. Even Jurassic Park has payed hommage to him in one scene. His naturalistic artstyle has reached a level of feeling real which I think has not been achieved ever since, even with modern digital art. There are some elements that do create an alien atmosphere in his works. A good example is the Iguanodon standing in a field of bones, while the moon shines in the background. It is a very eerie, otherworldly scene. But even this piece is an example of how Burian subverts the Alien Prehistoric World Trope. In the original 1950s book where it is from it is used to illustrate a short story told by Augusta, named The Call of Silence. It is about an old Iguanodon whose herd gets attacked by a Megalosaurus. The attack separates the old animal from its family and it retreats to a dinosaur-graveyard to die alone. In that light the scene becomes very melancholic and humanizing. Burian and Augusta's view of evolution and Earth’s history was not one of a savage survival of the fittest nor a march of progress, as it may have been for people such as Charles Knight, but a melancholic series of individual stories about life and death. In Augusta’s stories and Burian’s accompanying artwork the dinosaurs are no longer villains or obsolete monsters, but the protagonists of their own stories, often being the victims of fate. The dinosaurs become tangible and even relatable creatures, animals that lived and died like you and me, not alien monsters.  There are two more notable things: Burian rarely, if ever, depicts scenes of outright violence. At best he shows predators running towards their prey or standing above an already killed animal, but with no blood or gore visible. He therefore largely defies the trope of the prehistoric world being more violent than the modern one. The second thing is that he applied his extremely good intuition of living animal biology onto his paleoart, which is why, unlike Knight, he managed to give his dinosaurs accurately proportioned muscle mass. Just compare his Tarbosaurus to Knight's Allosaurus from part 2. Burian's creations come off as way more believable, actual animals than in many works before him. This is especially impressive, because since he was a citizen of Soviet Czechoslovakia he never had direct access to the fossils concerned and instead had to heavily rely on photographs of mounted museum-skeletons, other artist's skeletal drawings and advice from the paleontologists he worked with. 

Fig. 7: Among Burian’s last paintings was this Barosaurus lentus from 1976. As Darren Naish suspects, it is most likely based on a similar piece by Robert Bakker. Burian was likely trying to adapt to the commencing Dinosaur Renaissance by portraying sauropods as erect-legged land-dwellers with their tails off the ground. What is often forgotten is that as far back as 1938 Burian depicted at least some dinosaurs as agile and active animals, most notably the bird-like ornithomimids, but also hadrosaurs and T. rex.

With the 1960s coming to a close it seems like dinosaurs and their world were on the way to lose their alienness. They were rather archaic and somewhat strange animals, but animals with their own beauty nonetheless. They were becoming familiar beasts, just like our lizards and tortoises, and the Alien Prehistoric World Trope may have started to die. However, the children who were inspired by Zallinger’s and Burian’s art were now growing up and became palaeontologists themselves. Looking back at the old art, the old books and the old mounted skeletons, one of them realized something that would forever change paleontology: “Something is very wrong with our dinosaurs.”

With that I thank you again very much for reading this part and hope to see you next time when we come to the Dinosaur Renaissance.

If you liked this and other articles, please consider supporting me on Patreon. I am thankful for any amount, even if it is just 1$, as it will help me at dedicating more time to this blog and related projects. Patrons also gain early access to the draft-versions of these posts.

Literary Sources:

• Brusatte, Steve: The Rise and Fall of the Dinosaurs. A New History of a Lost World, New York 2018.
• Burian, Zdeněk/Spinar, Zdeněk: Life Before Man, Prague 1972 (Revised Edition from 1995).
• Davidson, Jane: A History of Paleontology Illustration, Bloomington 2008.
• Schalansky, Judith: Die Verlorenen Welten des Zdeněk Burian, Berlin 2013 (Naturkunden 8).
• Volpe, Rosemary: The Age of Reptiles. The Art and Science of Rudolph Zallinger’s Great Dinosaur Mural at Yale, New Haven 2007.

Online Sources:

• Palaeoart Memes and the Unspoken Status Quo in Palaeontological Popularization by Tetrapod Zoology (the SciAm Years)

Image Sources:

• Fig. 1: Volpe 2007, p. 2.
• Fig. 2-4: Volpe 2007, foldout.
• Fig. 5: Schalansky 2013, p. 105.
• Fig. 6: Schalansky 2013, p. 113.
• Fig. 7: Schalansky 2013, p. 178.

The Evolution of Fur - A Historic Perspective

Fig. 1: A 1988 illustration by Gregory S. Paul, showing two South African therocephalians, Trochosaurus, feeding on their distant tapinocephalian relative Styracocephalus. The small animal in the foreground is the therocephalian Blattoidealestes. All animals are notably drawn with a thin coat of hair.

When did hair first evolve? I do not know. Neither does anyone really at this point. Compared to the evolution of feathers, which is fairly well understood thanks to the discovery of amazingly preserved dinosaur-fossils, the first appearance of mammalian fur is quite a bit more mysterious. Is it something that is unique to mammals and their immediate ancestors or did it have its origins farther back in time? Were famous stem-mammals, such as Dimetrodon, Inostrancevia or Lystrosaurus scaly, leathery or hairy in appearance? It is a contentious question and one that continues to be relevant when we try to reconstruct these animals. While this article will not offer a definitive answer, it is nonetheless interesting to see how the different takes on these questions have evolved over time and how they might change in the future.

The mammal-like “reptiles” (Late 19th century – 1968)

Fig. 2: Dimetrodon as portrayed by Charles R. Knight in 1897. The skin and posture is strongly inspired by modern living reptiles. Guessing by the spiny sail, the pelycosaur in the background may be an early rendition of Edaphosaurus.

Amniotes, the animals which develop their young inside an amniotic sac (which includes us), were classically split up into three major groups: Anapsida, Diapsida and Synapsida, named after the number or form of their post-orbital skull openings or fenestra. Anapsida, sometimes called Parareptilia, have none and it was once thought that this group gave rise to turtles. Diapsida have two openings and the group includes lizards, snakes, crocodilians and dinosaurs (birds included), as well as a whole host of extinct forms. Synapsida, the group that would eventually give rise to mammals, have only one such opening (in humans this was eventually closed off, but you can still feel where it used to be if you touch your sphenoid bone or temple). The name means “fused arches”, as it was originally thought that synapsids descended from the diapsids and evolved their opening by combining the two fenestra. All synapsids that were not mammals were therefore classified as a sub-class of reptiles in classic taxonomy. It was only later realized that they did not descend from diapsids, that they instead just shared a common ancestor among the stem-amniotes and that they evolved their skull openings independently. The name, however, stuck. To quote zoologist John C. McLoughlin: “Like the Indians of North America, our remote ancestors were named by bumbling neophytes” (McLoughlin 1980, p. 29). What also stuck was the perception of early synapsids as reptiles or reptile-mammal missing links. They were most commonly called “mammal-like reptiles”, a term that one can still find in modern books, documentaries and museums. This then had a great effect to how they were represented in art. Even though no skin-impressions were known at the time, famous artist Charles R. Knight firmly believed that these animals had scales, with hair being a development of only mammals proper, and illustrated them as such (Knight 1946, p. 10). Most famous is his illustration of Dimetrodon from 1897, showing the animal with a very crocodile-like skin and a sprawling gait. His 1946 drawing also has some elements of a tuatara. His paintings of other synapsids, such as dicynodonts and gorgonopsians, were in similar vein. Works like these of course had a great influence on public perception, which came to view these animals as examples of early reptiles to the point where Dimetrodon is even today often mistaken as a dinosaur. That many of these species were given names ending in -saurus (Ancient Greek for lizard) such as Varanosaurus, Edaphosaurus or Lystrosaurus of course did not help (in my opinion the suffix saurus is way too overused in nomenclature and often inadequate at describing a species’ appearance).

Hippo-skin, whiskers and cladistics (1968 - 2012)

Fig. 3: “He has no style, he has no grace, this therapsid has a funny face”, to paraphrase the DK Rap. Estemmenosuchus as illustrated by Julius Csotonyi in 2011, shown with a smooth, glandular skin. My scanner unfortunately could not catch the entire picture, showing the rest of the herd. Splashing around in the background is distant relative Eotitanosuchus. 

After decades of illustrating early synapsids as scaly reptiles, an interesting discovery was made in 1968. Skin-impressions of Estemmenosuchus, a dinocephalian from the Middle Permian first described in 1960, were found. These showed that the animal was in fact not scaled, at least in the parts found, but instead possessed a smooth, glandular skin, very similar to what you would expect from a hairless mammal. This made sense from an evolutionary viewpoint, as this means it likely possessed a form of sweat-gland, out of which milk-glands would one day evolve (or perhaps already did?). The problem was the question of how representative Estemmenosuchus was for the rest of the early synapsids. As a dinocephalian therapsid it was fairly midway on the road from the first synapsid to the first mammal. It was more closely related to us than to Dimetrodon, however too basal to give much clues about later therapsids like dicynodonts or gorgonopsians. Further compounding was the fact that Estemmenosuchus is theorized to have had a specialized semi-aquatic lifestyle similar to a hippopotamus (to the point where I fear that Estemmenosuchus-Hippo comparisons will one day become one of those cliché paleoart-memes). Its skin-type might have been similarly derived and not representative of the ancestral state. That said, if it descended from a scaled ancestor it would make little sense for it to become scaleless, as many aquatic and marine reptiles keep their scales. However, it is possible that, just like a hippo, it may have descended from a hairy ancestor and secondarily became smooth-skinned. Or all its ancestors simply were already smooth-skinned and there was nothing special about its skin-type. Artists largely went with the latter hypothesis and in many great artworks since the 60s and up until very recently non-mammalian synapsids were often reconstructed with smooth, hair- and scaleless skin like that of hippos, elephants, rhinos or naked mole rats. The art of Zdeněk Burian is a good example of this.

Fig. 4: The Triassic dicynodont Lystrosaurus, painted by Zdeněk Burian and apparently imagined to have had a semi-aquatic lifestyle and smooth, hippo-like skin. Fun Fact: At one point in time this genus was actually the most abundant tetrapod on Earth, due to the ecologic imbalance left by the mass-extinction at the end of the Permian.

Some of course went a bit further than that and it was not uncommon to see illustrations of cynodonts with hair. From my own childhood books I particularly remember Cynognathus often being shown as this quasi-mammal with fur and ears, but also sprawling legs and lizard-tail. Also common was Thrinaxodon with whiskers (often only whiskers), the most prominent example being in the first episode of Walking with Dinosaurs. The speculation about whiskers in cynodonts goes all the way back to the 1950s when it was speculated that foramina found in their snouts were attachment-sites for vibrissae. However very similar structures were also found in the skulls of modern reptiles, such as the tuatara, which obviously do not have whiskers. We can therefore not definitely say if they had whiskers or not based on these foramina.

Fig. 5: The Cynognathus of my childhood (ca. 2005). I remember no book back then about prehistoric animals that did not have its own version of the supposedly furry cynodont. Today however it feels like it is only rarely featured in books. I wonder why that is. By the way, the German text states that the animal had fur as if it were a fact, even though this would be viewed as speculative even today.

Of course, speculating about hair in non-mammalian cynodonts was not particularly spectacular, as they were the immediate relatives of mammals and showed many signs of already being warm-blooded. During the 70s and 80s some went a bit further however. In his 1979 book Archosauria, John C. McLoughlin for example illustrated an anteosaur-like dinocephalian (the exact species is not given) not only with fur but also with little ear-pinnae, something that would be seen as highly speculative even today, given how “primitive” dinocephalians still were. In the semi-sequel Synapsida from 1980 he however seems to have doubled down on this, illustrating most synapsids as smooth-skinned and speculating that fur was unique to cynodonts (McLoughlin 1980, p. 84). While 1986’s The Dinosaur Heresies was, as you guessed it, primarily about dinosaurs, author Robert T. Bakker dedicates a chapter to the evolution of warm-bloodedness and speculates about endothermy in the Middle Permian synapsids of the Karoo:

“In Kazanian times [today this would be called the Guadalupian epoch], the Karoo experienced cool winters because it was closer to the South Pole than it is now. To remain active all year round, the small animals might well have needed some kind of insulation. Orthodox paleontology insists that hair is a uniquely mammalian invention, the adaptive badge of our own Class Mammalia. Several dissenters have however suggested that perhaps hair evolved long before the first true mammal appeared in the Late Triassic. Perhaps hair did evolve at the very beginning of warm-bloodedness, back in the Kazanian, fully forty million years before the earliest true mammals. Maybe our picture of the Kazanians should include shaggy protomammals stalking their prey through a winter snowstorm, hot breath steaming from their nostrils.” (Bakker 1986, p. 414)

The illustrations in the book however still show them as smooth-skinned, as was the norm of the time. His vision however became prophetic for later paleoart.

Fig. 6: A fuzzy dinocephalian, perhaps meant to be Anteosaurus, as illustrated by John C. McLoughlin in 1979’s Archosauria. His later depictions of synapsids in Synapsida were surprisingly more conservative, perhaps because he was criticized for a lot of his speculative takes in the previous book (although many of his predictions, like feathered theropods and rearing diplodocoids, later turned out to be true).

The late twentieth century also saw the advent of cladistic classification, which made the classification of early synapsids as reptiles obsolete. They could now be viewed as their own lineage in the amniote family-tree, distinct from reptiles (which themselves became regrouped into Sauropsida, consisting of diapsids and parareptiles). Despite its still widespread usage in popular media, the term “mammal-like reptiles” became outdated and it has become more common for non-mammalian synapsids, such as Dimetrodon, to be simply called stem-mammals by experts. Colloquially the term protomammal is also used. The notion of glandular-skinned stem-mammals was further complimented by the discovery of skin-impressions by sphenacodontids such as Haptodus in 2009, which showed they possessed dermal scutes, likely similar to those found in crocodilians or turtles. These also found their way into illustrations of pelycosaur-grade synapsids (synapsids that are not therapsids).

Nobody expects the Soft-Tissue Inquisition (2012 – 2018)

Fig. 7: What? Never seen a furry Moschops? You prude. This image was posted on Tumblr by someone with the username Alphynix in 2016, right during the Soft Tissue Revolution which we are still experiencing. It follows the common trend of taking a fairly well-known creature and illustrating it in novel and speculative ways to break up tropes and traditions.

In the wake of the Dinosaur Renaissance, paleontology entered a new golden age of discoveries and artwork. However not everything was fine and dandy, especially in the reconstruction-department. Beginning in the 2010s it was noticed that paleoart ran the danger of becoming too reliant on tropes and unspoken rules that were not necessarily based on facts. Especially shrink-wrapping, the act of neglecting the influence of soft-tissues on reconstructions, became a major problem. In response to all of this All Yesterdays, written and illustrated by Darren Naish, John Conway and Memo Kosemen, was released in 2012 to point out and subvert various unfounded tropes and clichés in paleoart and to encourage more well-informed speculation on things that are not preserved in the fossil-record. While the book was mainly focused on dinosaur-reconstructions it started a broader trend in paleoart as a whole called the Soft-Tissue Revolution. Reconstructions of stem-mammals were not untouched by this and many new artworks started to appear which showed these animals in new, speculative and unique ways. Perhaps in emulation of the trend of reconstructing increasingly more dinosaurs with feathers, it became more common to show therapsids outside of Cynodontia with fur. Gorgonpsians were a favorite candidate, as they already resembled dogs and wolves in overall appearance, but also dicynodonts and even dinocephalians became furrier. The squirrel-like arboreal dromasaurs also made more sense in a coat of hair. Some went even further than that and illustrated pelycosaur-grade synapsids with distinctively mammalian traits. A good example comes from All Your Yesterdays, a fan-sourced sequel to All Yesterdays. Here an illustration by Mike Keesey shows a Dimetrodon-mother with tufts of hair and a fatty sail, sweating out proto-milk from her sides which is then licked up by her young, a behaviour very much inspired modern monotremes such as the platypus. Around the internet more radical examples can be found, such as Dimetrodon with full fur-coats, whiskers and even mammalian muzzles. How likely all these speculations are remains debatable, but recent finds partially supported some of it. In 2016 Piotr Bajdek et al. described Late Permian coprolites (fossilized dung) from Russia, produced by either therapsids or archosaurs. In one of them they found traces of what was most likely hair, probably coming from the predator’s last meal. This presents the best evidence of fur having evolved long before true mammals appeared, but to whom this hair belonged is frustratingly unknown, as neither the prey nor the predator could be identified. The likeliest candidate would be a cynodont, however dicynodont bones were also abundantly present in the coprolites. A year before that, a study done on Ophicacodon, a very basal synapsid from the Early Permian that essentially looked like a Dimetrodon without the iconic sail, also showed distinctive signs of endothermy in this species. Endothermy, the ability to control one’s body-temperature through internal metabolic processes (not to be confused with homeothermy, the ability to keep a stable body-temperature), is a major step towards true warm-bloodedness and eventually necessitated the evolution of insulation, such as fur. Years before this study, in 2009, evidence for another distinctively mammalian trait, parental care, was found in the varanopid Heleosaurus, a synapsid even more basal than either Ophiacodon or Dimetrodon.

Fig. 8: A speculative depiction of Dimetrodon, titled Texan Mama, illustrated by Mike Keesey and first appearing in 2013’s All Your Yesterdays. The animal has a fatty hump surrounding its sail, tufts of hair and feeds its children with proto-milk-like secretions from its skin. Believe it or not, but I have seen speculative drawings that were way more bizarre than this. 

Both in science and art, the stem-mammals were beginning to take on a distinctive new look. No longer did they look and act like primitive reptiles, but instead came to resemble their mammalian descendants a lot more. Many of them also stopped being ugly or weird-looking and the desire to get a furry Diictodon as a pet became irresistible. Then everything changed when the Varanopid-Nation attacked.

Revenge of the Scales (2018 - 2019)

In the year 2018 a new species, Ascendonanus nestleri, was discovered. It was a member of the Varanopidae, one of the most basal clades of the Synapsida (though read on). It was significant in that it was preserved with an outline of the body and complete impressions of the skin, the first pelycosaur-grade synapsid to do so. It showed that it was covered in scales very similar to those of a lizard. This was rather surprising, as the lack of reptilian scales was up to that point used as a defining trait of synapsids. It also meant that the evolution of fur likely did not occur at the pelycosaur-grade but a lot later and that the smooth skin of Estemmenosuchus may have just been a transitional phase, not the ancestral state. This put a lot of the smooth-skinned and furry reconstructions of pelycosaurs into question. Artists were however quick to adapt. Shortly after the discovery Mark Witton produced a new rendition of Dimetrodon, clad in scales and obviously inspired by Charles Knight’s 1946 version, which is set to appear in the as of yet unreleased book Life through the Ages II. It is supposed to come out this year, however something awkward has happened in the meantime.

Fig. 9: A scaly Dimetrodon illustrated recently by Mark Witton, very much inspired by Knight’s 1946 rendition of the animal. It is however not entirely retro, as the sail-tips are now exposed (something which is currently believed to have been the case) and the animal is surrounded by juveniles, presumably its children.

The Big Sike (2019 – point of writing)

Almost like an early Christmas present, on December 23 2019, palaeontologists David P. Ford and Roger B. J. Benson presented a new cladistic analysis of early amniotes. Not only did they conclude that parareptiles were in fact a group of derived diapsids, but that the same was true for varanopids. This is not that surprising, considering that similar results had been produced by earlier studies for individual members of both, but this was the first extensive study that used both groups in their entirety. While at this point it remains to be seen if this becomes consensus, if true, it has many implications for the early evolution of amniotes, but also for the recent reconstructions of stem-mammals. Ascendonanus and its relatives would not be synapsids anymore, its scales being nothing out of the ordinary since it would be more closely related to lizards than to mammals. It therefore could not be used anymore as a reference to reconstruct early synapsids. Where does this leave us? On one hand it seems safe to go back to pre-2018 speculations, meaning smooth, glandular skin and perhaps fur, until skin-imprints of definitive pelycosaurs are discovered. On the other we might also have to abandon some older concepts. Heleosaurus, used as evidence for parental care, was also a varanopid and could therefore not be used as reference anymore as well. 

Fig. 10: I sometimes make half-assed memes in my free time. It is a peaceful life.

This entire story just goes to show how rapid and fluctuous of a science paleontology and paleontology-illustrations have become. While it means we should never grow too attached to a single piece or style of reconstruction, it is a good thing as the growing data and discussion only helps us better understand deep time and our own origins. The fact that paleoart ages is a feature, not a flaw, to paraphrase Mark Witton. Even when an artwork ages, it is still useful to us, as it literally illustrates what science and the culture around it was like at a given point in time. On a deeper level we have also seen how interested we apparently are in the evolution of fur and our earliest relatives. It is always strange to realize that we share more in common with bizarre creatures such as Dimetrodon or Moschops than one might expect, so it is tempting to familiarize them with hair and parental care, all traits that we do not only recognize in other mammals, but also in ourselves. While we humans like to think of ourselves as naked apes, hair is still very much part of our bodies and our lives (we in fact have more hair-follicles than chimpanzees, our hairs are just a lot thinner and shorter). Our hair is not a vestige from prehistory, it still serves a purpose, the most obvious in ours being that it helps us shape our appearance and personality. And hey, perhaps, just perhaps, back in the Middle Permian the first hairy synapsid got quite a lot of looks from the ladies and that is why his lineage still lives on. 

If you liked this and other articles, please consider supporting me on Patreon. I am thankful for any amount, even if it is just 1$, as it will help me at dedicating more time to this blog and related projects. Patrons also gain early access to the draft-versions of these posts.

Literary Sources:

• Bakker, Robert Thomas: The Dinosaur Heresies. New Theories Unlocking The Mystery of the Dinosaurs and Their Extinction, New York 1986.
• Benton, Michael James: Vertebrate Paleontology, 1990 (3. Edition from 2005, German translation).
• Burian, Zdeněk/Spinar, Zdeněk: Life Before Man, Prague 1972 (Revised Edition from 1995).
• Conway, John/Kosemen, C.M./Naish, Darren: All Yesterdays. Unique and Speculative Views of Dinosaurs and Other Prehistoric Animals, London 2012.
• Conway, John/Kosemen, C.M./Naish, Darren: All Your Yesterdays. Extraordinary Visions of Dinosaurs and Prehistoric Animals from a New Generation of Paleoartists, London 2013 (Second Edition from 2017).
• Chambers, Paul/Haines, Tim: The Complete Guide to Prehistoric Life, London 2005.
• Csotonyi, Julius/White, Steve: The Paleoart of Julius Csotonyi. Dinosaurs, Sabre-Tooths and Beyond, London 2014.
• Knight, Charles Robert: Life through the Ages, New York 1946 (Commemorative Edition).
• McLoughlin, John: Archosauria. A New Look at the Old Dinosaur, New York 1979.
• McLoughlin, John: Synapsida. A New Look into the Origin of Mammals, New York 1980.
• McRae, Anne/Nardi, Marco: Tessloffs Enzyklopädie der Dinosaurier, Nürnberg 2005. 
• Paul, Gregory Scott: Predatory Dinosaurs of the World. A Complete Illustrated Guide, New York 1988.

Papers:

• Botha-Brink J, Modesto SP. A mixed-age classed 'pelycosaur' aggregation from South Africa: earliest evidence of parental care in amniotes?. Proc Biol Sci. 2007;274(1627):2829–2834. doi:10.1098/rspb.2007.0803
• Ford, D.P., Benson, R.B.J. The phylogeny of early amniotes and the affinities of Parareptilia and Varanopidae. Nat Ecol Evol 4, 57–65 (2020). https://doi.org/10.1038/s41559-019-1047-3

Online Sources:

• Ancestry of mammalian 'warm-bloodedness' revealed by Science Daily
• Getting to the Root of Fur by National Geographic
• The Stem-Mammals--a Brief Primer by Tetrapod Zoology (Ver. 3).

Image Sources

• Fig. 1: Paul 1988, p. 52.
• Fig. 2: Wikimedia
• Fig. 3: Csotonyi 2014, p. 6-7.
• Fig. 4: Burian 1972, p. 106
• Fig. 5: McRae 2005, p. 25
• Fig. 6: McLoughlin 1979, p. 102-103
• Fig. 7: Tumblr-user Alphynix
• Fig. 8: Conway 2019, p. 99.
• Fig. 9: Mark Witton's Twitter. 

Belated Game Review - Ancestors: The Humankind Odyssey

“When I joined the Corps, we didn’t have any fancy-schmanzy tanks. We had sticks! Two sticks, and a rock for the whole platoon – and we had to share the rock!”
- Sergeant Avery Johnson, Halo 2 Anniversary

In the very first post of this blog I said I would also take a look at how videogame developers have imagined the prehistoric world. Consider that fulfilled with this post and similar ones I plan for the future. Ancestors: The Humankind Odyssey is the first and hopefully not last game by indie-developer Patrice Désilets’ Panache Digital Studios and has been in development since 2014. It originally released for PC on August 27 2019, but since my computer is trash I had to wait until December 6 when the game was also released on PS4. I therefore call this post a belated review. 

Fig. 1: Ever looked at a painting like this and thought: “I want to experience this for myself”? Well, now you can, sucker!

Given how Désilets had previously worked on the Assassin’s Creed games (of which I have only played the first third of the third instalment) and the game had been in development for quite some time, some considerable hype seems to have built around it. For me personally however, the game only appeared on my radar very close to release and I only got seriously interested in playing it after watching some youtubers getting eaten by Miocene giant otters. I was therefore able to approach the game without any real expectations that could be disappointed. The original concept of the game was for it to have an episodic-content-structure in which you relive the most important moments of early human history, from the first hominin to the start of civilization. This was eventually changed and the game seems to have now been broken up into a trilogy, with this first instalment taking place during the first 8 million years of human evolution. While this post mostly tries to be a game review, I also want to be a bit experimental, just like the game, and treat it like a piece of paleoart, because that’s what it essentially is (unless you do not count videogames as art, but that is a discussion for another place). Videogames are rarely talked about on these types of paleo-blogs, perhaps because most games featuring prehistoric animals cause a paleontologist somewhere to cry (looking at you, ARK).

On a final note before we start, this article may include spoilers for the late-game, but I will try to mark these where possible so you can avoid them.

Part 1: Jungle Japes

There are not that many games set in the stone age, at least not many original ones. Most opt for a 10’000 B.C.-setting at the end of the last ice age, but these, like for example Far Cry Primal, seem to be more inspired by the Roland Emmerich movie than the actual time period, featuring some quite outrageous anachronisms. Other, less serious games, like Joe & Mac, go for a 1 Million B.C.-setting (by that I mean the Ray Harryhausen movie) with Frank Frazetta-style cavemen fighting dinosaurs. By his own account, Désilets was tired of these settings and went for something unique that was not inspired by pop-culture. As the opening cutscene tells us, we begin in the steaming jungles of Africa, a whopping 10 million years ago, which would be during the mid-to-late Miocene. Judging by the geography and time, the game world seems to be based on the Olduvai Gorge in Tanzania. We see the food-chain in full action, as we follow a fish getting caught by a Pelagornis, which lets it fall back into a river by accident. This scares a group of monkeys, which would have otherwise been attacked by a python. The fish gets caught again by a heron, which gets eaten by a crocodile, causing it to lose the fish as well. As we follow the dead fish further downstream, we see a crocodile getting attacked and killed by a black sabertoothed cat, before the fish falls down a waterfall. As we see the wide valley in front of us, the title card appears and tells us that this is volume 1 of the story, Before Us. At the foot of the waterfall, the fish gets grabbed by an ape, carrying its child on its back. With it he climbs up a tree to enjoy his food in peace, but a giant bateleur eagle appears, grabs the ape and violently kills it with a pluck to the head. The child the ape was carrying can flee however and falls down the canopy. Now it is all alone on the forest floor with all these predators around, as fear cloaks its vision.

Fig. 2: Just because a game is set in the stone age does not mean there can’t be terrifying dinosaurs. The opening cutscene seems to  be a direct reference to the real life Child of Taung, an Australopithecus which was found inside the fossilized nest of a bird of prey.

This is how you start the game. It gives you some tutorial-text, which tells you how to move and how to use your intelligence to scan the environment. With that you find a hiding spot and hide inside it. Your perspective switches to an adult ape still at the settlement who hears the child’s cry for help and your task is now to venture out into the jungle to rescue it. You learn to use your other senses, mainly hearing, and how to climb and swing from tree to tree. After you rescue the child, you bring it back to the settlement and a rather heartwarming cutscene plays. Everything that happens after this is up to you. Explore the area, eat everything in sight, bang rocks together, figure out the game’s mechanics… in the words of Scarface: The world is yours. The game will occasionally give you some short tutorial-texts (which you can turn off or read again in the Help-section of the pause-menu), but only for the most basic controls, like which button does which action and what the symbols on the UI mean. I believe these were also only added in a later patch. What you can do with this type of rock, which types of food can be eaten and which ones not, what animals are easily intimidated and which ones will eat you for dinner and so on and so forth is never told to you, instead you have to find that out yourself. Ancient humans did not have tutorials either, so why should you?

If that is too overwhelming for you, the game can give you mission objectives depending on the game-mode you’re playing on. These range from recruiting new apes into your clan, finding a mate and producing a baby to exploring certain spots on the map or intimidating a certain animal. These are optional and exist to help you understand what you can do and accomplish in the game. As this was my first time playing it, I naturally tried to follow these as close as I could. I ran into problems however when it came to the objective to explore a site where a small meteorite had impacted into the jungle. Actually finding the spot was not difficult, as it was quite obvious by the smoke column coming from it. The real problem was that there was a significant section of jungle and swamp between me and the meteorite that I had to cross. In the process of reaching it, I think I got eaten by the same crocodile about three times in a row, interspersed with sabretooth- and warthog-attacks. I was straight-up not having a good time and actually decided to restart the game about three times because I did not want to live with the consequences of my actions. On my final try I decided to tackle the problem in a smart way. With a child on my back, a spear in my hand and accompanied by an elder ape (I believe it was the child’s grandpa) that was also armed, I ventured out again to find this damned space rock. I still had bad luck. I was ambushed by a sabertooth and unable to dodge it in time. I got tackled onto the ground and could not defend myself. Adrenaline rushed through me as I would not only die again but also lose a child in the process, but the game let me immediately switch to the elder ape that came with me. With nothing but a sharpened stick in my hand, I ran towards the cat and, either by good reflexes or pure luck, finally managed to kill something. As the beast goes down with a violent stabbing to the stomach and eye and even with some punches to the face, I am overcome with great relief as I see my family-members badly bruised but still alive. Using two granite rocks I create a makeshift grinder and turn some horsetails into paste, which I give my grandson to stop the bleeding. It was this entire moment that finally made something click with me and I realized I was going to have a great experience with this game.

Part 2: Hominin see, hominin do

So what do you actually do in this game? From a gameplay-perspective, the closest thing I could describe it as is a mix of Spore, Saurian, Niche and perhaps the cancelled Peter Molyneux game B.C. The overarching goal is to move from 10 million to 2 million B.C. and in the process develop a tribe of hominids into something you might call humans. You can switch between all your tribe members, which you play from a third person perspective. A circle on the lower right of the UI shows you all the members of the tribe (except the one you are currently playing as), represented by points. This circle also shows you if any of your tribe-members are in danger, outside the settlement, following you, pregnant or carry a genetic mutation. At maximum you can have 18 tribe members, 6 elders, 6 adults and 6 children, each fulfilling different roles. The lower middle of the UI is a green circle, which doubles as your stamina-, health-, food- and water-meter and is limited by your life-expectancy. Lower left is a dopamine-meter, which also shows you if your ape is calm or alerted by danger. If you are alerted, you can run faster and intimidate animals, but it will drain your dopamine. If it drops to zero you will actually lose control over your ape and in a panic it will run back to the settlement. Upper middle of the UI is a clock that shows you the time of day. There is no map whatsoever and the game even gives you the option to play completely without UI to give you a better immersion.

Fig. 3: The user interface, very early into the game.

Controlling your ape is pretty straightforward for a third person game. Holding X makes you run or climb, releasing or tapping X makes you jump or grab objects. Circle and square are used for context-sensitive actions and triangle lets you use your senses to scan the area. You have three senses: Intelligence (basically seeing), hearing and smelling. Climbing, your main mode of locomotion at the beginning, is pretty fun and intuitive and the only good defence you have against predators in the early game. Just don’t climb too high or the eagle will get you. Climbing also is about the only thing you are really good at in the beginning, as the apes you begin with are a pretty pathetic excuse for humans. They look like a weird mix of chimpanzee, bonobo and gorilla, they cannot walk bipedally, they can only eat plant-matter without getting sick, their stamina, dopamine and life-expectancy are low, their senses are limited, they cannot talk and their motor skills are too bad to alter most types of rock or wood. How do you advance with these prospects? Here the game functions similarly to an RPG. You unlock neuronal points on a braincell-skill-tree by doing certain tasks often enough. This is pretty situationally based, so while the game still technically shows evolution inaccurately as a goal-oriented process, you nonetheless just adapt to your environment, which does make it more realistic. For example, if you live in a place where you get attacked by animals a lot, you will eventually unlock a lot of neuronal points that allow you to better dodge or counter-attack them. If you live close to water and have to swim a lot, you will mostly unlock neurons that will improve your swimming. Actually unlocking the neuronal points does not automatically grant you the abilities however. You first have to unlock them using your neuronal energy. The amount of neuronal energy you have is displayed as a white orb inside the settlement circle in the lower right. You gain neuronal energy by performing actions in front of children. This is how the game simulates learning in primates. Monkey see, monkey do. While this is easy when you do things inside your settlement, when you go on expeditions you have to bring the children with you by carrying them on your back or belly if you want to gain energy. The amount of energy you get from this is pretty high, but you also run the risk of you and your child getting killed. Even losing one child can put a significant dent in your already low population. Thankfully you can craft weapons to defend yourself, but these are not your caveman-style spears, clubs or bows. The strongest weapons you will ever wield are a sharpened stick and an olduvai-style basalt chopper, but they get the job done. Many Machairodus died to give you this information.

Part 3: Homo: Combat Evolved

Of course you cannot realistically advance 8 million years by simply living on a day-to-day basis. The game features two ways with which you can go forward in time and advance your species: generational leaps and evolutionary leaps. To enter the menu for these you actually have to build a nest/bed first in which you have to lie down. Only here can you also access the previously mentioned skill-tree and use your neuronal energy to unlock new abilities. When you leap a generation, 15 in-game years pass. Your children grow up to be adults, your adults age into elders and your elders turn into a pile of bones. The downside of this is that you cannot bring all the unlocked neurons with you to the next generation. In general, the game only allows you to permanently lock as many neurons as the number of children you have in your tribe, which at maximum is six. The neurons you could not carry over you have to unlock again by gaining new neuronal energy, which many have understandably described as grindy at times. Personally I was never really that frustrated by it for some reason and towards the late-game, when you have already unlocked most neurons, it becomes a negligible issue. There are also ways by which you can fix more than just six neurons. Across the map there are a number of meteorites and gem stones. Finding them will grant you one additional point with which to fix neurons. Meteorites can even cause genetic mutations in case you are carrying a child with you. Meteorites are relatively easy to find, as they announce their presence through large smoke-columns, although these vanish after some time. Gem stones are usually hidden inside caves or on tall rock-towers and make their presence known by a faint, creepy whisper. 

Fig. 4: The neuronal skill-tree, very late into the game.

The second way of moving forward in time is the evolutionary leap. Throughout the game you fulfil evolutionary achievements, either by finding certain places and objects, killing animals or making them fight with each other. These achievements you can cash in during the evolutionary leap, as they will be converted into additional time you can jump forward to. The amount of children you have born also adds to the time-counter, while the amount of deaths you have caused will subtract from it. If you are doing well you can jump hundreds of thousands of years this way and once you hit certain time-markers you will evolve into new species known from the fossil record, which will grant you better stats and new abilities. The game also gives you an extra-challenge here, as you race in time against a red bar which represents the speed at which actual science says our lineage evolved. Can you evolve faster than our real ancestors? In theory it is possible to reach 2 million B.C. by just living in the same place all day and hitting the generational and evolutionary leap buttons repeatedly, and I have actually read about people who have played the game this way, but this is not only boring but will also make you lose this challenge. The game therefore greatly incentivizes you to go out there, explore the world and accomplish things. Evolutionary leaps will also unlock genetic mutations. Genetic mutations are neurons you cannot unlock the usual way. They can appear at random in a child when it is born and you are informed what the mutation does when it grows up into an adult, but you can only unlock the mutation’s abilities when they are carried by an adult or elder and you make an evolutionary leap. I did not realize this during the first couple of hours I have played, which caused me to severely lag behind in things like bipedality and omnivory.

Part 4: I bless the rhinos down in Africa

Let us talk about the world and the life in it a bit. The map is pretty huge and features a wide variety of biomes. I have seen many people who actually did not realize this when starting the game, even though the trailers already showed off some places. I guess this is because (when you play the beginner-mode) you start in the jungle, which is not only cluttered but also hard to get through, so most do not move far. Once you go past it, travelling becomes much easier, however. You find large lakes, caverns and woodlands and once you go past them you come to a wide and open savannah. Connected through a canyon, you can go from the savannah to a barren desert. At the easternmost end of the map you finally hit the African coast of the Indian Ocean. While the game’s graphics are not on the highest end of the current spectrum, they still manage to create a quite lively and beautiful world. There is a lot of detail, from tiny frogs that hop around in swamps to a river that starts out in the savannah and slowly dries out as it passes through the desert. The lack of any in-game map may make it difficult in the beginning to navigate yourself. However, despite lacking any man-made structures, the world has enough charismatic natural landmarks which can help you orient yourself. With your intelligence-sense you can also memorize certain locations. Another thing that helps is that the entire map is essentially placed inside a giant gorge/valley, so you can also use the gigantic stone walls for orientation. Each new biome comes with new plants, animals and hazards. When it rains in the jungle, you can run the risk of freezing, while an average day in the savannah can cause you to overheat. There are plants and other food which can help you prevent these effects, but it is up to you to find which ones have which effect.

Fig. 5: Me and my tribe on our way to Pride Rock to kill Mufasa. 

Now for the more interesting part: The animals. The game’s fauna is largely based on species from Miocene Africa, which so far, I think, is a first for any popular piece of media, let alone videogame. The Miocene is an epoch that is usually not well represented, even in paleoart, as the more bizarre-looking mammals of the Cenozoic lived in the Paleogene, while more familiar ice age creatures come from the Pleistocene. If you are a particularly big fan of the Miocene, this is the game for you. I actually learned about a lot of animals for the first time thanks to it. In general, the game represents animal-types one would also encounter in modern day Africa, but usually represented by extinct relatives. Instead of African elephants, you encounter the four-tusked Stegotetrabelodon, the rhino and hippo are Ceratotherium neumayri and Hippopotamus gorgops respectively, for hyenas we have Adcrocuta, for crocodiles we got Crocodylus thorbjanarsoni and instead of zebras you will encounter the three-toed Hipparion (whose coloration seems to be based on the recently extinct quagga, which is a nice detail). However there curiously are also some modern species, such as the African Rock Python, mambas and Dorcas gazelle, which I somehow doubt existed since 10 million years ago. Your main enemy in the game is the sabertooth cat Machairodus, which comes in differently colored variations to represent different species/sub-species. After every generational leap there will actually spawn a specific machairodont in the world, which has been nicknamed Stalker Cat by the community, as it can randomly ambush you at any moment unless you manage to kill it (which the game does not tell you). Killing or attacking enemies is a bit peculiar, as there is no actual way in the game to directly attack animals, perhaps as a reference to the fact that australopithecines are not thought to have been big game hunters yet. Instead you have to wait for them to attack you first, which makes you enter a quick-time event. To dodge, hold X and move in a certain direction away from the enemy. When you hear an audio-cue (a short chime), release X and you will dodge the attack successfully. To attack, you have to hold a weapon in your hands and do the same, but move towards the enemy. Most interactions in the game, like grooming or making tools, are based on a similar QTE mechanic where you have to hold a button and release it once you hear the audio-cue. It takes some time to get used to, but eventually muscle-memory will set in. The downside to it is that people with hearing-problems will probably have a harder time playing, so maybe there should be an option for a visual cue instead. If you are successful in your counter-attack, an entertainingly gruesome cutscene will play in which your hominid will violently stab and punch the animal to death. Maybe it is because I am a madman, but it gives you a satisfying feeling of catharsis, as most creatures in the game honestly deserve it. The great majority is openly hostile to you and either hunts you on sight for an easy meal or attacks you if you just violate their personal space too much. The catharsis-factor of the cutscenes is of course reversed when the counter-attack or dodge fails and the animal gets to kill you instead in graphic detail. For me the most disturbing death scene has to be when the eagle gets you, as it strangely reminds me of a similar one from Dead Space 2 of all things (the one where Isaac gets hammered to death by a Stalker). You can also make the animals fight each other, often for your own benefit. For example, one day as I was strolling through the savannah, I accidentally walked next to a hyena-burrow. As they noticed and chased me, I ran away, but was not fast enough to outrun them. So instead I ran towards a nearby herd of water buffaloes, which dealt with the hyenas while I could escape. The cutscenes that play when two animals fight are also a bit disturbing, especially when you see the Stegotetrabelodon kill things. Despite being a herbivore I found it to be the most terrifying animal in the game.

Fig. 6: There is of course nothing more terrifying than man itself however…

There are a few things that can be criticized about the in-game fauna. For one, even though you yourself evolve into different species and technically move in time from the Late Miocene to the Early Pleistocene, the animals stay the same. Secondly, there are also (of course) some scientific inaccuracies, which ironically befall the modern species more than the fossil ones. Most, like the bateleur eagle and rock python, are simply oversized, likely to present a more serious threat to you. The most outrageous example is the Scolopendra-centipede, which you encounter in and around cave-systems. In real life it is about 30 centimeters long at most, but in the game it is roughly the same size as the pythons. It is in fact large enough to kill a sabertooth cat. Speaking of which, the game identifies its sabertooth as Machairodus giganteus, which is nowadays regarded as a dubious species and has been reassigned to the genus Amphimachairodus.The third and last point is that, despite the developers including species as obscure as the giant otter Enhydriodon dikikae, they mostly went for creatures that resemble modern species instead of also including more weird and interesting animals. This is Miocene Africa after all. Realistically, you should still be encountering chalicotheres, hyaenodonts, amphicyonids, deinotheres, primitive giraffids and weird gomphotheres. It is a really missed opportunity to not include at least some of these as animals you could encounter in the game. All that said, in the opening crawl that plays before the intro cutscene, the developers point out that they did not try to make an accurate depiction of a specific time and place, but rather create a generalized prehistoric setting as a playground for you to evolve in. It is also still cool to simply have a game where you can encounter and interact with obscure Miocene mammals. I still would have liked to mess with some chalicotheres though. 

Fig. 7: Bizarre Miocene mammals like Platybelodon, here restored by Zdenek Burian, unfortunately do not appear in the game, which is a bit of a shame.

On a last note regarding presentation, the music is quite good. It is largely a mix of African folk music with some synth(?) elements and seems to differ from biome to biome and time of day. It can get quite atmospheric at times. My only complaint is that it is not memorable enough, at least for me. Perhaps when it comes to ape-games I am simply spoiled by David Wise’s Donkey Kong Country soundtracks.

Part 5: Hominin Hijinxs  (Warning: Contains spoilers about the species you evolve into and the ending. May be skipped.)

When you start the game you begin 10 million years ago as a species that is simply titled “?” or The Missing Link. Based on the name and appearance, one presumes this to be the last common ancestor of humans and chimpanzees (but read on). When you reach 9 million years ago, you become Sahelanthropus tchadensis. At around 7.2 you reach Orrorin tugenensis. At 5.3 you become Ardipithecus ramidus, at 3 million Australopithecus afarensis and at 2.5 Australopithecus africanus. When you reach 2 million years B.C. the game ends as you see your species evolve into Homo ergaster. Every new species brings with it a higher dopamine-, and neuronal energy-capacity than the previous one, as well as higher life expectancy. Most also bring new gameplay elements with them. Orrorin for example can control its emotional status, allowing you to run faster even when you are not in danger. Ardipithecus can slow down time using its concentration, while Australopithecus afarensis has higher heat-resistance. A cute detail is that, much like the other animals in the game, each species is given a nickname based on their respective type-specimens, such as Lucy, Toumaï or Ardi. The difference in physical appearance of each species is very subtle when compared with the one that comes before and after, but over time you will notice drastic changes. The Missing Link and Sahelanthropus look like how you would generally imagine an ape: knuckle-walkers, covered in fur and long faces. The sounds they make are a mix of stock chimpanzee-sounds and what sounds like humans making monkey-noises into a microphone (although considering what you are playing as this maybe is appropriate). Orrorin is not much different, but the white of its eyes is now visible, which gives the facial expressions a sudden human touch in a very subtle way. By the time you reach Ardipithecus you probably evolved at least some degree of bipedality. Ardi has a bit less fur and some of the sounds it makes now start to resemble primitive words. Australopithecus afarensis follows this trend until you hit full uncanny valley with A. africanus. It is now hairless on most parts of its body, its face has almost-but-not-quite human proportions and its voice-commands sound like the beginnings of language. By this time you should also be fully bipedal and able to digest meat from animals. The changes will become most apparent to you in the ending cutscene, where we see all the stages you played as in a classic march of progress. It is ended by a pair of now definitely human-looking Homo ergaster, a male and a female holding a child, looking back on the hominids you played as, perhaps sentimentally, before walking off (unfortunately you do not get to play as H. ergaster, most likely as a sequel-hook). The game has many moments like these throughout it which are oddly touching and perhaps will make you think about what it actually means to be human. Every time you make an evolutionary leap, a short cutscene plays where we can see the new hominid being in a state of deep thought or confusion, as if he is pondering about what just happened. When you go to sleep you can see your hominid’s dreams in form of pictures floating above its head, which in their composition foreshadow the cave-paintings that will be created millions of years in the future. Even with the earliest species you play as, you can see heartwarming moments of hominids caring for their children, smiling to show happiness (in non-human apes smiling is usually seen as a sign of aggression), hugging newcomers to the tribe or comforting scared relatives. Interestingly you can also gain a lot of neuronal energy by simply examining the remains of your dead relatives, which may be a reference to the development of spirituality. It is notable that there is also no option to harm anyone of your own species, even though you can kill everything else in the game. We have come a long way from the very brutish and barbaric cavemen of past depictions.

Fig. 8: In general, the way Ancestors portrays early hominins is more similar to Zdenek Burian’s peaceful and naturalistic paleoart than other games, like, say, Far Cry Primal, which took more cues from Frank Frazetta. Pictured here is Burian's vision of Australopithecus gracilis. I once replaced a framed photo of my father with this picture and it took months until someone noticed.

Let’s get the gomphothere in the room out of the way: The line of descendancy the game shows is not exactly unproblematic. First and foremost, the timing at which you evolve into these species is a bit off. Sahelanthropus lived 7 million and Orrorin 6 million years ago, not 9 and 7 respectively (although of course the exact temporal range of species is hard to determine). Ardipithecus did exist around 5.3 million years ago, but under the species A. kadabba. A. ramidus came later. The game also follows the classic, but severely outdated “March of Progress”-style of depicting evolution, popularized by the Rudolph Zallinger painting from 1965. The game in fact uses it as its logo. The major problem with the March of Progress is that it portrays the evolution of man as a straight line of progress with us as the end-goal, while shoehorning species into our ancestry that should not belong there. Similarly, the game portrays some species as our ancestors, even though there is a significant chance they may not have been. While it is very likely that the genus Homo descends from Australopithecus (which brings a lot of nomenclature problems with it) and that the latter descends from or is closely related to Ardipithecus, Orrorin and Sahelanthropus are a lot more controversial. Sahelanthropus may be an ancestor of hominins, but has also been interpreted by some as a female proto-gorilla. Orrorin could be an ancestor of Ardipithecus, but has also, very controversially, been argued to have been the direct ancestor of Homo, which would make Australopithecus a side-branch. It is also possible that Orrorin was just an unrelated case of convergent evolution. That you go from Australopithecus africanus directly to Homo ergaster is also a bit questionable. Shouldn’t Homo habilis or H. rudolfensis come somewhere in-between? A. garhi or A. sediba have also been argued to be more likely ancestors to Homo than A. africanus. That said, the line of descendancy the game shows is not necessarily wrong and I would not blame the developers in case it becomes outdated soon. Like most branches of paleontology, paleoanthropology is currently going through a large boom and is in constant flux. The major problem seems to be that we have way too many transitional forms and possible cases of convergent evolution, many of them living at the same time, to reconstruct the evolution of modern man as a species-to-species line with much certainty, unless we have DNA-evidence available. Speaking of which, molecular evidence has shown that the last common ancestor of humans and chimpanzees lived about 7 to 5 million years ago. That you start the game 10 million years ago would therefore in fact mean that you are not just playing the ancestors of humans, but also of chimpanzees and even gorillas (our split with them was about 8 million years ago). The “Missing Link” you start as therefore cannot be the last common ancestor of humans and chimps, but would be closer in time to something like Nakalipithecus. Perhaps the game is aware of this, as the menus and UI call your fellow tribe-members “hominids”, a term that is nowadays generally used for great apes as a whole, instead of the more human-specific “hominins”. However, given how the two terms are often conflated in popular media, I am not sure if this was intentional. If it is intentional, the March of Progress-style the game goes for makes it miss an interesting opportunity for branching evolutionary paths by only portraying the human side of the story. What if I want to evolve into gorillas or chimpanzees instead? Similarly, wouldn’t it have been interesting to meet and interact with parallel human lines as you progress through the game, like Kenyanthropus or Paranthropus?

Fig. 9: Timeline of appearance of major hominin genera. Note that this does not directly imply a line of descent. Also note how many of them overlap in time.

All that said, the simple fact that the developers went for this part of human evolution and chose such obscure species is commendable, as is the very subtle attention to detail in how they look, act and evolve. The fact that there exists a videogame where you can play as an Ardipithecus and get eaten by a Pelagornis is simply amazing to me.

Part 6: In conclusion

In the end, what can be said about Ancestors? Is it a good game? If not, is it at least a good piece of paleoart? Personally, I had a lot of fun, but only after I fought myself through the very steep learning-curve in the beginning. Ancestors is extremely open-ended about how you can approach the game and learn its mechanics, but in order to best put you into the perspective of a prehistoric ape with no idea about the future, it does not easily give away its secrets. I found out that the best way to enjoy it is as an exploration game. It heavily rewards you for discovering and interacting with new things and even on my second playthrough I am still finding new stuff. The game is also not simply about reaching a goal, but about writing your own stories. I experienced intense life-and-death situations, like the Machairodus-attack I mentioned in the beginning, and moments where smart problem-solving helped me out of tricky situations, like the hyena-buffalo gang-war. I went on a long, exodus-like migration through the desert with my tribe to reach the peaceful coast. While walking through the woodlands I accidentally fell down a hole into a cave-system where I was confronted with a giant centipede. Out of pure fun I made a hyena follow me to a watering hole, just so I could see it get attacked by a crocodile. I raised children and rescued them from an angry hippo, armed with only a bone-club. I experimented with all kinds of tools to figure out the best way to crack and digest snake eggs. I singlehandedly killed an elephant with only sharpened sticks and a rock. Ancestors truly is an odyssey and much like Homer’s epic, it is the journey itself that counts, not the goal. Some of the most memorable gaming-moments I had in the last decade were made in this game and none of it was scripted. There is also good potential for replayability, as the game features two other modes next to the “story” one. In the survivor mode you start out alone 10 million years ago in a random location and first have to recruit other hominids before you can even form your tribe. Then there are user-defined scenarios where you can decide in which location you want to start and which species you want to play as.

And how well does Ancestors do as a piece of paleoart? Of course there are some inaccuracies due to artistic licence, but most, like the oversizing of the animals and the anachronisms of the hominid-species, as well as the goal-oriented evolution, were taken to benefit the gameplay. It is great that the fauna and even the proto-humans you play as largely consist of species that are rarely depicted in popular media, although there are some missed opportunities to show off some even more obscure or bizarre creatures. A lot of research did go into the game and I think it shows. For non-paleontology enthusiasts it will be a neat introduction to the Miocene epoch and perhaps to paleoanthropology. While the graphics may not be the highest, the presentation is great, as the world and animals feel alive and are full of subtle attentions to detail, as are the hominids you play as. While not 100% accurate, this is probably the most authentic video game depiction of a specific moment in deep time until Saurian will be released (and that will probably still take a while).

Fig. 10: Homo heidelbergensis, as illustrated by Zdenek Burian. If Ancestors ever gets a sequel, which I really hope it does, we may get to play as this species. While still controversial, it is generally thought that Homo sapiens descends from an African variant of H. heidelbergensis.

In conclusion, I really like this game, but I would perhaps not recommend it to everyone. The learning curve, especially in the beginning, is very high and you can get easily overwhelmed with all the new information. There will be times where the game may feel a bit grindy or slow and bad luck can lead to great frustration or even to the extinction of your lineage. It can be irritating that there is no way to directly attack animals or to manually save and the game arguably ends just when evolution starts to get really interesting. However, if you master the difficulties of the start, let the flow of the game be guided by your own curiosity, play at your own pace, explore everything there is to discover, are interested in human evolution or prehistory in general and hate big cats with a passion, then I believe you will have a great time going full homo in Ancestors: The Humankind Odyssey.

Did I write this entire review just so I could make that joke? Yes. Do I regret it? Absolutely not. 

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Literary sources:

•  Agusti, Jordi/Anton, Mauricio: Mammoths, Sabertoothsand Hominids. 65 million years of Mammalian Evolution in Europe, New York 2002.
• Anton, Mauricio/Turner, Alan: Evolving Eden. An Illustrated Guide to the Evolution of the African Large-Mammal Fauna, New York 2004.
• Anton, Mauricio: Sabertooth, Bloomington 2013.
• Burian, Zdenek/Spinar, Zdenek: Life Before Man, Prague 1972 (Revised Edition from 1995).
• Gould, Stephen Jay: Wonderful Life. The Burgess Shale and the Nature of History, New York 1989.
• Müller-Beck, Hansjürgen: Die Steinzeit. Der Weg des Menschen in die Geschichte, München 1998 (4. Edition).
• Naish, Darren: Tetrapod Zoology 1, London 2010 (2. Edition).
• Schalansky, Judith: Die Verlorenen Welten des Zdeněk Burian, Berlin 2013 (Naturkunden 8).

Papers:

• Guy et al.: Morphological affinities of the Sahelanthropus tchadensis (Late Miocene hominid from Chad) cranium
• Hawks et al.: An Ape or the Ape: Is the Toumaï Cranium TM 266 a Hominid?
• Wood et al.: The evolutionary context of the first hominins

Online sources:

• The Cautious Climber Hypothesis by Tetrapod Zoology
• Systematic Classification of Life by Aron Ra

Image Sources:

• Fig. 1: Anton 2013, p. 221.
• Fig. 7: Burian 1972, p. 177.
• Fig. 8: Burian 1972, p. 205.
• Fig. 9: Wikimedia
• Fig. 10: Burian 1972, p. 217.
• All images of gameplay were taken during my own playthrough, using the PS4's capture-mode.