The perception of palaeontology in commercial off-the-shelf video games and an assessment of their potential as educational tools
Video games now comprise the largest sector of the media entertainment industry. Hundreds of video games, spanning a huge variety of genres and platforms, use extinct animals and/or palaeontological themes as a basis for their gameplay. Because of this, many players, especially children, spend long periods of time engaging with and being immersed in palaeontological concepts and themes. Video games may be the first medium of implicit or tangential science communication they engage with, especially with regards to palaeontology. However, commercial off-the-shelf video games are not primarily designed to be educational tools, and the proliferation of some common tropes can disseminate harmful and/or unethical (mis)information regarding palaeontology. This paper introduces the major types of palaeontological video games and discusses their educational potential. We highlight the most common palaeontological tropes, both positive and negative, observed in video games to better equip science communicators regarding the perception of palaeontology (and ancient animals) in this massively influential medium that they may encounter when undertaking scientific engagement. Furthermore, by highlighting common misconceptions and harmful tropes, we aim to bring awareness to game developers, who may be unaware that they could be propagating negative tropes about palaeontological science.
Video games are the largest and fastest growing sector of the media entertainment industry. In 2020, approximately 2.7 billion users generated an estimated USD 159.3 billion in revenue; more revenue than the movie and music industries combined (Newzoo Global Games Market Report, 2020). The rise of games playable on mobile devices has massively increased the accessibility of video games to a broad range of demographics, mainly because they do not require specialist hardware, are often free-to-play, and have huge diversity in regard to subject matter and necessary skill level. Furthermore, gaming-related entertainment, such as live-streaming services on websites like Mixer, Twitch, YouTube, etc., is hugely popular, with over 12.4 billion hours of gaming live-streams having been consumed in the first two quarters of 2020 (Stream Hatchet Video Game Streaming Trends Report, 2020). Combined, this indicates that not only are video games increasing in popularity but that, when not playing, many users will still engage with video games by watching others play online via streaming services. Palaeontological science communicators should recognise the size of the captive audience and how influential the medium of video games is as a resource for engagement practices.
Broadly speaking, video games can be separated into two categories: educational and commercial games. Educational games actively promote learning by introducing information and engaging the user with specific content, known as “digital game-based learning” (DGBL; see Habgood and Ainsworth, 2011). Video games that focus on specific STEM subjects have been shown to be beneficial to both non-students and students by positively aiding learning outcomes (e.g. Shute et al., 2013; Mani et al., 2016; Pringle et al., 2017), especially for students who struggle with traditional teaching practices (Mayo, 2009). However, studies also indicate that some users can find educational games patronising (Klopfer and Osterweil, 2013) or lose interest rapidly and disengage, decreasing the likelihood of positive learning outcomes (Kerawalla and Crook, 2005; Habgood and Ainsworth 2011). Furthermore, educational video games are typically niche in their subject matter, difficult to programme (e.g. Pringle et al., 2017), difficult to implement (e.g. Mani et al., 2016; Mozelius et al., 2017), and require extensive funding to develop (Annetta, 2008; Mayo, 2009). These games are often made in association with specific outreach projects with limited funding (e.g. Mani et al., 2016; Pringle et al., 2017), making protracted development an impossibility (Mayo, 2009). It should also be noted that it can be difficult to monitor the learning outcomes of these educational video games (e.g. Mani et al., 2016), especially if they are released to the public. Ultimately, educational video games are viable learning aids, but they do not appeal to the wider public.
The second type of video games are commercial off-the-shelf (COTS) games. These games make up the overwhelming majority of the gaming ecosystem and are incredibly diverse in their genres and themes. Typically, COTS games are developed to a much higher standard of graphics and gameplay mechanics than educational video games (Mozelius et al., 2017) and are designed specifically to capture the attention of the user by motivating and challenging them in an interactive environment, often reinforcing this through repetition and reward (Brown et al., 2014). Despite the dated perception that video games are nothing more than a trivial time sink, evidence suggests that video games can have positive benefits for the cognition, motivation, and social skills of players (e.g. Granic et al., 2014). Furthermore, COTS games can effortlessly provide early exposure to educational themes and concepts, even if the specific COTS game is not primarily designed for this purpose. Because of the huge variety in genres, the engaging gameplay, the ability to play online with friends, and the high production quality, it has been proposed that COTS games are a resource that could be incorporated into teaching environments (Van Eck, 2006), with multiple studies undertaking the utilisation of COTS in teaching environments (e.g. Charsky and Mims, 2008; Mohanty and Cantu, 2011; Sun et al., 2015; Caldwell et al., 2017). Recently, some COTS games have included specific sections of the game that are focused on education: for example, Assassin's Creed: Origins (2017, Ubisoft Montreal) and later instalments in the Assassin's Creed franchise contain a “Discovery Tour” mode that removes combat and quests in the game to allow players to explore the games' historical settings, accompanied by factual information and quizzes. Generally speaking, however, whilst educational video games often sacrifice entertainment for accuracy, many COTS video games do the opposite (Van Eck, 2006; Mozelius et al., 2017; McGowan and Scarlett, 2021), and must therefore be carefully vetted prior to integration into curricula, often leading to increased workload for educators (e.g. Sun et al. 2015; Caldwell et al., 2017).
One important and often overlooked aspect of computer gaming is the potential for implicit and tangential learning (Mozelius et al., 2017; Crowley et al., 2021; McGowan and Scarlett, 2021). Implicit (or unconscious) learning occurs when learning takes place in an incidental manner without the awareness of the person engaging in a given activity (e.g. Frensch and Rünger, 2003). Conversely, tangential learning is the semi-conscious process of self-education via exposure to a topic in an already enjoyable, non-educational format (Brown et al., 2014). Many COTS games impart complex scientific and historical content to their audiences by presenting the topic within fun and engaging game mechanics in a non-scholastic format (see Herrero et al., 2014; Crowley et al., 2021) and can be used as a tool within a wider educational framework (Herrero et al., 2014). However, it is also possible that any implicit and tangential learning from a COTS game may be a completely unintended or entirely accidental element of game design. Consequently, because COTS games are not primarily designed with educational purposes in mind, inaccuracies – either intentional (disinformation) or unintentional (misinformation) – could have a powerful effect on less-informed players, misleading audiences about key concepts and even reinforcing harmful stereotypes.
This is particularly pertinent to the science communication community, especially for palaeontology. For over 150 years, ancient organisms have inspired a plethora of popular books, films, cartoons, and toys. It should therefore be of no surprise that ancient life is a popular topic for the video game industry. In fact, one of the first 3D games for the home computer was 3D Monster Maze (1982, J. K. Greye Software), a maze game where the player was chased and, unless they could successfully escape, eaten by a Tyrannosaurus rex (Fig. 1). Since then, the diversity of palaeontological video game genres has greatly increased and is currently a popular theme for COTS games (see Fig. 2). This means that a considerable number of people, particularly children, will be introduced to the field of palaeontology through the medium of video games. Because players repeatedly engage with a single video game and because new games containing palaeontological themes are released frequently, COTS games have a much greater reach than traditional outreach events.
The aim of this paper is twofold: (1) to familiarise science communicators with the types of palaeontological COTS video games; and (2) to identify and discuss common tropes and misconceptions that often arise where scientific accuracy is compromised for entertainment value. This paper seeks to raise the awareness of science communicators regarding the perception of palaeontology (and ancient animals) that the video-game-playing public are exposed to, especially those that have been introduced to the subject via these games. Here, we highlight harmful tropes (i.e. unethical behaviour, misogyny, racism, etc.) to inform science communicators of the perception of palaeontology that is disseminated by COTS video games. Furthermore, this review may be of interest to COTS game writers, developers, and video game artists who may be unaware that they are propagating damaging tropes pertaining to palaeontological science.
In this review, we focus on COTS video games that incorporate fossils and/or ancient animals as one of the playable or interactive aspects of the game; we term these as “palaeo-video games”. This review does not include games based on human remains or artefacts (archaeology).
Currently, there is a huge diversity of video games that contain palaeontological content. As of 2021, there are over 270 palaeo-themed COTS games available on Steam, the largest digital video game distribution service (PC games only) (Fig. 2). It is important to note that this number does not include games that have been released for games consoles (such as the SNES, PlayStation, Xbox, etc.), and so the number of palaeo-themed COTS games is much greater. Many of the games available on Steam are made by small development teams and will not sell in large quantities; however, mainstream titles can often sell large numbers of games. For example, by 2020, Jurassic World Evolution (2018, Frontier Developments) sold over 3 million copies (Kerr, 2020). Due to the vast quantity of palaeo-themed COTS games, there are titles not explicitly discussed herein.
In this review we do not discuss COTS games only incorporating ancient animals as “set dressing”. Examples include games such as The Last Of Us Part II (2020, Naughty Dog), Batman: Arkham City (2011, Rocksteady Studios), and BioShock (2007, 2K Boston), which feature levels in museums containing skeletons or animatronic replicas of extinct animals, or Battlefield 4 (2013, DICE), which features a hidden “Easter egg” that allows the players to summon a giant Otodus megalodon during live multiplayer matches. While some of these games, such as The Last Of Us Part II, do have playable museum levels that allow players to interact with fossils, typically the palaeo-content of many of these games is purely aesthetic and, therefore, would be of limited use to science communicators. Another common use of ancient animals is purely cosmetic; many multiplayer games feature “skins” that the players can unlock to customise their characters, and these often resemble ancient animals, e.g. dinosaur outfits in Fortnite (2017, Epic Games) and dinosaur-themed armoured suits in Destiny 2 (2017, Bungie Inc.) (Fig. 2). These “skins” are purely aesthetic changes that do not alter the core gameplay and will not be discussed further.
In order to understand the tropes commonly seen in palaeo-video games, it is important to identify the major categories of palaeo-video games available on the market. Despite the diversity of game types and styles (Fig. 2), most games align with one of the following categories (though some span multiple): (1) ancient animals as adversaries (Fig. 3a); (2) ancient animals as tools (Fig. 3b); (3) fossils as collectibles (Fig. 3c); (4) ancient animal management simulators (Fig. 3d); and (5) ancient animal simulators (Fig. 3e).
2.1 Ancient animals as adversaries
This category is, by far, the most common type of palaeontological video game (65 % of palaeo-game titles on Steam; Fig. 2d). Many video games use ancient animals (typically dinosaurs and pterosaurs) as adversaries that must be defeated (i.e. killed) for the player to proceed and continue the game. In this type of game, the ancient animals are programmed to be hostile if approached and/or will actively hunt the player. Examples include The Legend of Zelda (1986, Nintendo EAD), Turok: Dinosaur Hunter (1997, Iguana Entertainment; Fig. 3a), Trespasser (1998, DreamWorks Interactive), Dino Crisis (1999, CAPCOM), Tomb Raider (1996, Core Design), and Peter Jackson's King Kong: The Official Game of the Movie (2005, Ubisoft Montpellier).
Another type of palaeo-video game within this category is the “wave shooter”, in which the player fights off waves of hostile enemies with a suite of exotic weapons in order to progress. The enemies in these games are presented as cannon fodder that exhibit little or no natural behaviour, swarming en masse to overwhelm the player. Examples include Dino Stalker (2002, CAPCOM), Dino D-Day (2011, Digital Ranch), Dinosaur Hunt (2015, Racing Bros), Carnivores: Dinosaur Hunter (Reborn) (2015, Digital Dreams Entertainment LLC), Wrath of the Goliaths: Dinosaurs (2018, Ascendence Studios), Hunt: Primal Reptiles (2020, HugeLittleStudio), Prehistoric Hunt (2020, Antiproto Studios), and Second Extinction (2020, Systemic Reaction). Some games subvert this category and allow the option to play as the ancient animals in order to hunt and kill humans, such as Primal Carnage (2012, Lukewarm Media) and Orion Prelude (2012, DANKIE).
Generally, category 1 games offer very limited educational content. At best, the player can access a glossary of the hostile units, which may contain some scientific information; however, many of these games rely heavily on tropes such as monsterification (see below) to make the enemies appear and behave more frightening or lethal. In fact, in most category 1 games, the ancient animals could be replaced by any other generic adversaries (i.e. aliens or zombies) with little effect on core gameplay.
2.2 Ancient animals as tools
Another common use of ancient animals in COTS games is as a tool to help the player progress through the game. One of the most famous and recognisable examples is Yoshi, a fictional, omnivorous theropod dinosaur who first appeared in Super Mario World (1990, Nintendo EAD; Fig. 3b). Yoshi was a mount for the Mario brothers, who rode Yoshi in their adventures to rescue Princess Peach; however, Yoshi's popularity promoted him to full sidekick status and has spawned several dedicated sequels (e.g. Yoshi's Story, 1997, Nintendo EAD). A variety of other games allow tamed ancient animals to be utilised as mounts, such as Far Cry Primal (2016, Ubisoft Montreal), Ark: Survival Evolved (2017, Studio Wildcard), Total War: Warhammer I and II (2016, 2017, Creative Assembly), and World of Warcraft: Battle for Azeroth (2018, Blizzard Entertainment), with some games even going so far as to mount weapons, such as machine guns and cannons, to ancient animals, e.g. Nanosaur (1998, Ideas From the Deep), Dino D-Day, and Total War: Warhammer II.
Ancient animals as tools are also featured in the video game sub-genre of role-playing games (RPGs). In the context of palaeontology, the most famous examples are the Pokémon games, in which a diverse group of Pokémon are based on or inspired by ancient animals. In these games the player can catch, train, and fight the creatures in sanctioned competitions in order to become the best trainer in their respective worlds. Pokémon fossils can be found, revived, and used to battle other trainers as the player progresses through the game (e.g. Pokémon Red/Blue, 1999, Game Freak). Similarly, the Fossil Fighters series (2008, Nintendo EAD) allows players to take part in detailed fossil excavations, revive the discovered dinosaurs (referred to as vivosaurs), and train them to fight in order to complete the main story.
Similar to category 1 games, category 2 games have very limited educational content but may have some information about the ancient animals in databases or glossaries.
2.3 Fossils as collectibles
Collectibles are a common aspect of COTS games: players can collect items found throughout the game setting as they progress through and explore the game. These collectibles are often an optional side quest separate to the main storyline – they provide additional world-building or upgrades, but are used in some games as resources to craft tools and materials.
Fossils are frequently a form of collectible seen in COTS games (Fig. 4). For example, Red Dead Redemption 2 (2018, Rockstar Games; Fig. 4e) features a side quest where the player is tasked to locate 30 fossil specimens across the game world in order to assist an aspiring palaeontologist. Another example is in the Pokémon franchise; as mentioned above, fossils can be found and resurrected by the player in order to complete the secondary objective of the game (Fig. 4o): filling the digital Pokémon encyclopaedia, or Pokédex, with every type of Pokémon that can be found in the game world (e.g. Pokémon Red/Blue).
Most other games use fossils, alongside crystals, ore, geodes, etc., as a natural resource that can be used to collect, sell, or build other objects. In Minecraft (2011, Mojang; Fig. 4n), The Sims 4 (2014, Maxis; Fig. 4f–h), Starbound (2016, Chucklefish; Fig. 4i–m), Stardew Valley (2016, ConcernedApe; Fig4. b–d), and Animal Crossing: New Horizons (2020, Nintendo EAD; Figs. 3c, 4p–t), players can collect fossils for a variety of reasons – from displaying them in virtual museums or in their own personal collections to using them as fertiliser. However, in the majority of category 3 games, fossils are also used as (or associated with) in-game financial resources as a tradable commodity. This has implications for the public perception of fossil collecting (see Representing ethics in palaeontology).
Virtual fossil collectibles are almost always based on real organisms and often include database entries or facts pertaining to the collectibles items. This has enormous potential for educating the player, provided the information is well researched (see Table S2 in the Supplement). However, category 3 games can easily impart misinformation or even disinformation, especially when presented in an otherwise informative context.
2.4 Ancient animal management simulators
“Construction and management simulations”, sometimes referred to as park management simulators (or park sims), are games where the goal is to construct a financially viable park or zoo by managing the construction, park layout, guest management, animal welfare, and breeding programmes with often very limited resources. Ancient animals tend to be a common focus for park sim games, and they often have a diverse variety of realistically modelled organisms tending to exhibit “naturalistic” behaviours (although some games are heavily stylised and/or simplistic for aesthetic reasons) for the player to engage with. There are several park sim games based on the Jurassic Park franchise, e.g. Jurassic Park III: Park Builder (2001, Konami), Jurassic Park: Operation Genesis (2003, Blue Tongue Entertainment), Jurassic Park Builder (2012, Ludia), and Jurassic World: Evolution (Fig. 3d), but independent franchises based on ancient animals are also popular, e.g. DinoPark Tycoon (1993, Manley & Associates), Zoo Tycoon: Dinosaur Digs (2002, Blue Fang Games), Parkasaurus (2020, Washbear Studio), and Prehistoric Kingdom (2022, Blue Meridian).
Category 4 games often contain the most detailed compendiums on ancient organisms, allowing the player to find information regarding their required virtual living conditions and about the relevant science and/or discovery of the organism. They can also contain detailed information about fossil sites, fossilisation processes, phylogenetic relationships, and evolutionary processes. As with previous categories, this information can yield useful educational content if accurate, but can be misleading if incorrect information or disinformation is presented.
2.5 Ancient animal simulators
The last decade has seen the emergence of “survival simulators” – games where the player can control an individual animal through an entire life cycle in a natural, open-world environment. This category does not include games such as Yoshi’s Story, Project Bolan (Dinosaur Game) (2014, Google) and DinoRun DX (2015, PixelJAM Games). We do not consider these games ancient animal simulators because although they allow the player to play as an ancient animal, the player’s avatar could be replaced by any other organism without altering the gameplay experience.
In survival simulators, the player's ancient animal avatar must survive by finding water and food (by hunting A.I.-controlled animals or, potentially, other players) while avoiding natural hazards and being predated themselves. Typically, these games have focused on dinosaurs, e.g. Saurian (2017, Urvogel Games, LLC; Fig. 3e), The Archotek Project (2017, The Archotek Project Team), Beasts of Bermuda (2018, Sastrei Studios, LLC), Dinosaurs: Prehistoric Survivors (2018, Arcupion Art), The Beasts of 9500 (2020, Dragons), Path Of Titans (2020, Alderon Games); however, some games do focus on non-dinosaurian animal groups such as early hominids, e.g. Ancestors: The Humankind Odyssey (2019, Panache Digital Games). Survival simulators tend to market themselves as being as scientifically accurate as possible in both their palaeo-environmental and animal reconstructions (e.g. Saurian).
Another genre of game within this category allows the player to visit ancient ecosystems as a passive observer. Video games such as Abzû (2016, Giant Squid Studios) allow the player to swim through ocean environments inhabited by a plethora of ancient aquatic organisms from throughout geological time. Robinson: The Journey (2016, Crytek) utilises virtual reality headset technology to create an immersive experience for the player, who must journey through a world inhabited by dinosaurs. This genre of game introduces the player to a diverse range of flora and fauna from the ancient past but is relatively uncommon.
Within the context of this paper, a “trope” is a recurring theme or motif that occurs across video games. Although many tropes are considered clichés, they are still commonly used throughout popular entertainment as recognisable touchstones to provide familiarity to the audience (see García-Sánchez, 2021). Many tropes seen in COTS video games are shared across the entertainment industry (especially in television and film); however, due to the interactive medium of video games, some tropes are unique to the format. Here we identify some of the most common tropes appearing in video games to highlight how palaeontological science is represented in pop culture and the entertainment industry and to demonstrate that some tropes propagated by the video game industry may be scientifically misleading and/or perpetuate harmful stereotypes.
3.1 Ancient death machines: monsterification
The most frequent use of ancient animals in video games is as enemies for players to combat. Typically, these animals are designed to appear vicious, frightening, and brutish; this is known as “monsterification”. This is often achieved by “shrink wrapping” (reducing the organisms' soft tissue until they are just skin and bone. See Conway et al., 2012), increasing the animal's body size to exaggerated proportions, and grotesquely exaggerating features like claws, teeth, and horns. However, monsterification is not limited to an organism's appearance — it can also include behaviour such as being unrealistically aggressive, erratic, and, most often, mindlessly and noisily torpedoing towards their prey. By consciously combining these aspects, ancient animals are made to appear more terrifying while also being physically dissociated from real animals, meaning that players have a desensitised and guilt-free experience slaughtering them. Interestingly, Jański (2016) found that when extant animals were used as background assets or companions for the player in video games, they were more likely to be accurately represented, whereas when animals were depicted as enemies or tools, their appearance and behaviour were more likely to be inaccurate, exaggerated, objectified, and even monsterified. Indeed, some games actively task the player with “monsterifying” ancient animals: Jurassic World: Evolution, for example, encourages the player to breed dinosaurs to create hyper-aggressive monsters in order to have them fight each other for the entertainment of park guests.
Monsterification is not a novel concept, nor is it unknown to science communicators. From as early as the 19th century, palaeo-reconstructions have liberally and creatively portrayed ancient life provocatively as monsters to stir the imagination, and these images have permeated into modern popular culture. However, for science communicators, there are some important aspects of monsterification that are not normally considered: (1) monsterification is not restricted to carnivorous animals in video games. Examples such as Orion Prelude, Dinosaur Hunt, and Second Extinction utilise hyper-aggressive herbivores such as stegosaurs, ceratopsians, and ankylosaurs as opponents for the player to dispatch. (2) Monsterification is not limited to dinosaurs. For example, ARK: Survival Evolved applies some degree of monsterification to almost every animal in the game – several taxa (e.g. Diplocaulus, Paraceratherium) are much larger than their fossil remains indicate, while the giant fish taxon, Leedsichthys, is coated in jagged spikes and scutes and is depicted as highly aggressive despite fossil evidence suggesting they were filter-feeding organisms. Similarly, Far Cry: Primal depicts “ice age” (Pleistocene) mammals as colossal, aggressive monsters that are, in the case of some animals, such as mammoths, twice the size of their largest fossil remains. (3) Extreme monsterification can lead to a situation where ancient animals are indistinguishable from and/or confused with mythical creatures. In ARK: Survival Evolved, a wide variety of ancient animals live alongside fictional animals, such as wyverns, basilisks, and gryphons, that the player can also interact with and tame. In-game information and even achievements do not differentiate between the mythical and real animals, which could lead to confusion for some players.
Monsterification of ancient animals is not the same trope as video game monster designs that are heavily based on the appearance of ancient life. This is very common in video games, with examples including Monster Hunter Rise (2021, CAPCOM), where many of the monsters clearly resemble theropod dinosaurs, and Horizon Zero Dawn (2017, Guerrilla Games), where a post-apocalyptic world is populated by an ecosystem of mechanical creatures loosely based upon extinct megafauna, including sauropod and theropod dinosaurs as well as terror birds, giant crocodyliforms, and cave hyenas.
3.2 Fossil = dinosaur = Tyrannosaurus rex: lack of palaeodiversity
Since their discovery in the 1800s, dinosaurs have captured the public imagination. Today, that popularity has continued, as dinosaurs dominate popular culture – so much so that they are almost synonymous with ancient life in the entertainment industry. The video game industry is no different, with the vast majority of video games incorporating ancient animals featuring only dinosaurs (Fig. 2c). The perpetuation of this trope poses a challenge to science communicators – as it presents a false impression to the audience that ancient ecosystems lack complexity – and also diminishes ancient diversity and species distribution through geological time and space.
Yet, even within this trope, the representation of dinosaur diversity is typically limited to a handful of “iconic” dinosaur species that are perpetually recycled by the entertainment industry (Fig. 5). Many of these, such as Tyrannosaurus, Stegosaurus, Triceratops, Brachiosaurus etc., species are well documented from North American fossil sites like the Morrison or Hell Creek Formation (see Dodson et al., 1980; White et al., 1998), while other commonly used species are typically predatory (e.g. Velociraptor, Spinosaurus). Within the video game industry, this recycling of dinosaurs serves a practical purpose: for smaller game developers, building new digital assets can be highly labour intensive and costly. It makes commercial sense to use pre-made, often free resources (e.g. 3D models, animations, sound effects, and reference material) to recreate popular and recognisable dinosaurs despite the fact that this further oversaturates their prevalence in the video game market place. Another group of ancient animals within the overused trope are the “ice age” (Pleistocene) animal pair of the woolly mammoth (Mammuthus primigenius) and the sabre-toothed cat (Smilodon sp.). As charismatic mega-mammals frequently used in media, they are recognisable to the public and often used in some video games as more relatable monsters than dinosaurs (e.g. Far Cry: Primal; The Elder Scrolls V: Skyrim, 2011, Bethesda Game Studios; Syberia, 2002, Microids.).
However, in recent years, there has been a growing number of games that use a greater diversity of dinosaur species as an active selling point to market the game. Park building games such as Jurassic World: Evolution allow the player to send teams of scientists to real-world fossil sites to collect DNA from a host of lesser-known dinosaur species, such as Baryonyx, Huayangosaurus, Proceratosaurus, Carcharodontosaurus, Tsintaosaurus, etc. Dinosaur simulator games such as Saurian often feature lesser-known dinosaurs, while Ark: Survival Evolved features an abundance of relatively obscure dinosaurs (e.g. Yutyrannus) but also Permian therapsids (e.g. Moschops and Lystrosaurus), Mesozoic birds (e.g. Ichthyornis and Hesperornis), and Cenozoic mammals (e.g. Chalicotherium and Phiomia). However, many of these animals are still “monsterificated”, and the game proves the point of this section by referring to every ancient animal in the game as a dinosaur in the in-game information.
As new games seek to find a niche within the gaming ecosystem, there are a number of popular video games featuring and drawing attention to even more obscure palaeo-taxa and time periods. The hugely popular game, Animal Crossing: New Horizons tasks players with finding fossils for display within a local museum and, in so doing, introduces players to a variety of relatively niche groups such as Anomalocaris (a stem arthropod), Eusthenopteron (a sarcopterygian fish), Acanthostega (a basal tetrapod), and Myllokunmingia (a stem chordate). Interestingly, these fossils (and others found by the player) are displayed in a large phylogeny demonstrating evolutionary patterns in a simple but highly informative manner. Another game, Abzû, features a large number of obscure aquatic ancient animals such as ammonoids, Anomalocaris, Arandaspis (a jawless fish), Diplocaulus (amphibian), Archelon (giant sea turtle), etc., although there is no context or distinction for the player as to where/when these organisms lived. Ancestors: The Humankind Odyssey avoids this trope entirely by being specifically set in Miocene Africa, including a suite of time-specific organisms such as Stegotetrabelodon (a stem elephant), Machairodus (sabre-toothed cat), and Enhydriodon (giant otter). All these games introduce the player to organisms and concepts they would have had limited (if any) exposure to in other types of media.
3.3 “Did you know...?”: palaeotrivia and palaeodatabases
In order to help the player understand complex topics or to introduce elements of the story, many video games feature in-built encyclopaedias or glossaries containing fact files about characters, objects, and locations integral to the game's setting. In games featuring palaeontological themes, fact files, databases, and bonus trivia provide the ideal opportunity for optional, longer-form educational content with the potential to inform and provoke further investigation from players, especially those who were not necessarily attracted to the game for the science.
In-game encyclopaedias can be extremely informative. This is particularly common in park management games, such as Jurassic World: Evolution and the Zoo Tycoon series (Blue Fang Games, 2001–2017), that feature highly detailed databases about each ancient animal found in game, often incorporating facts pertaining to the organism, such as where and when the animal was discovered, as well as general information. This information is not necessary to play the game, but because such databases are detailed and well researched, they often add to the “scientific” aesthetic of these types of games and are useful educational tools.
More typically encountered is the usage of “palaeo-facts” within different elements of the game. One excellent example is Animal Crossing: New Horizons, where Blathers, the strigiform (owl) museum curator, will regale the player with light-hearted but informative trivia about each fossil presented to the museum (see Table S2). Another example commonly employed is the use of “palaeo-facts” to entertain the player on loading screens (i.e. a picture the player must watch while the game is initialising). LEGO Jurassic World (2015, TT Fusion) uses this tactic to deliver bite-sized, informative facts about many of the dinosaur species encountered in the game.
Generally speaking, the inclusion of palaeo-facts and databases in video games are good science communication if they are well researched. However, if factual information is presented alongside misinformation, it can be difficult for some players to discern the difference. For example, in The Sims 4, various fossils can be found throughout the game world – including trace fossils, trilobites, plants, and raptor dinosaur claws. However, other fossils items, ranging from the ridiculous (fossilised cow udders) to the bizarre (“perfectly preserved moustaches” – Fig.4h) are presented as genuine fossil remains. While clearly light hearted and for comedic effect, the indiscriminate mixing of real and fake fossils presents a challenge for science communicators.
Tangentially, many of the games mentioned in this section have large, dedicated player fan-bases who often form distinct online communities. In order to assist with completing aspects of the games, many players create, edit, and maintain sizable, publicly editable online “wikis” (an online database resource). These online encyclopaedias not only provide game-specific information but often contain further reading and palaeo-trivia for interested players, making them a useful educational resource; however, the degree of moderation and fact checking varies wildly.
3.4 The depiction of palaeontological science
The perception of palaeontology as a scientific discipline is difficult to gauge in computer games. Most palaeo-video games convey good palaeontological communication by explicitly integrating physical and biological processes into gameplay and world-building that will, either directly or indirectly, increase the knowledge of the player. However, most games typically focus on very specific aspects of palaeontology (e.g. fossil collection, genetics, evolution) and can take creative liberties in order to make engaging gameplay mechanics. This can mean that the player may be presented with an inaccurate or misleading representation of palaeontological science. We discuss some of these below.
3.4.1 Geological processes
Though it is often taken for granted, many open-world video games rely heavily on aspects of climate, topography, biomes, and even natural disasters to create realistic settings, the exploration of which can subtly communicate basic earth science to the player. Enormous effort is taken to accurately capture the surface geology, flora and fauna, and variable ecosystems in open-world games such as Red Dead Redemption 2, allowing players to learn tangentially (see Crowley et al., 2021). Other games, such as American Truck Simulator (2016, SCS Software), allow players to take virtual road trips across hyper-realistic (albeit scaled-down) sections of the United States, including many famous North American fossil localities (e.g. Dinosaur National Monument, Colorado, USA). Other games incorporate geological processes to build the game-world for the player to explore. Subnautica (2018, Unknown Worlds Entertainment) is an underwater survival/horror game based in the crater of a guyot (a submerged volcanic seamount) on an alien planet. The explorable map is separated into distinct biomes, some of which contain large, fossilised remains preserved due to the area's specific environmental conditions. The processes of fossil formation (e.g. taphonomy) are explained to the player by scanning the fossils with scientific equipment in their inventory.
As computing power increases, greater graphical fidelity allows more accurate geological features to become standard in computer gaming; as a tangential effect, this can be used by science communicators in engagement activities. During the recent COVID-19 pandemic, where travel was restricted, some academic institutions created virtual field trips for their students using technology commonly utilised by video gamers (e.g. VR headsets; Klippel et al., 2019). Furthermore, video game engines allow complex geological features to be explored in ways that would be difficult for actual students in the field (e.g. utilising aerial imagery; Klippel et al., 2020). This demonstrates that video games have great potential for geological engagement, especially for players who are unable to access key sites due to travel restrictions or accessibility issues, not to mention the reduced environmental cost associated with virtual fieldwork.
3.4.2 Depicting evolution
“Evolution” is a common theme in video games; however, while video games are potentially a great way to introduce players to the complex process of evolution, it should be remembered that COTS games must prioritise delivering engaging gameplay far above educational content. It should also be noted that the term “evolution” is commonly used in video games to encompass a host of different game mechanics that do not accurately represent biological processes, potentially skewing the player's understanding of the phenomenon. One example of the term “evolution” being misused is as a levelling up mechanic. In the Pokémon franchise, the Pokémon “evolve” into larger, more powerful forms as they gain more experience and level up despite the process actually being a type of metamorphosis.
Another common misuse of the term evolution applies to games where novel morphological traits are acquired via predation. In Spore (2008, Maxis), players must guide the “evolution” of their own custom creatures from microscopic organisms to an interstellar empire by acquiring new, evolutionary traits by scavenging them from the creatures they ingest (see Poli et al., 2012; Herrero et al., 2014). This depiction of “evolution” is also seen in E.V.O.: Search for Eden (1992, Enix), where the main character begins as a generic fish and must acquire teeth, jaws, limbs, and other features by eating prey items as they advance through levels inspired by different geological eras. This gameplay mechanic reinforces the trope that evolutionary changes are a result of a conscious decision by an organism as opposed to random, coincidentally beneficially traits with a greater likelihood of being passed to the next generation via natural selection.
Other games tackle evolution as a gradual process with greater nuance; however, many of these games are limited by the need for fun game mechanics. In Cell to Singularity: Evolution Never Ends (2018, Computer Lunch), the player clicks or taps the screen to acquire currency, which is spent to advance Earth history in short steps. The game is presented as a branching tree, with each node representing a new species or event; however, the order in which evolutionary milestones are achieved is entirely arbitrary, presenting evolution as a simplified, linear checklist of sequential events. Even games accurately depicting ancient organisms and past environments often portray the processes of evolution poorly. Ancestors: The Humankind Odyssey depicts the evolution of early humans as distinct leaps from one species to the next as opposed to multiple species coexisting at the same time (see Snyder, 2022). Similarly, the game also refers to the levelling-up mechanism of learning new skills and abilities, such as being able to use tools, as “evolution”. By presenting evolution as an oversimplified, linear process, it also reinforces the outdated notion that humans are the pinnacle product of evolution.
Conversely, some games approach evolution from a palaeontologist's perspective – using fossils to infer evolutionary patterns. In the puzzle game Fossil Corner (2021, Overfull Games), the player is given a box of procedurally generated fossils and is tasked with resolving their family tree by observing morphological character changes through each generation. This introduces the player to the concepts underlying maximum parsimony, a method commonly used by palaeontologists to generate phylogenetic trees for extinct organisms. As such, Fossil Corner is a good example of digital game-based learning (DGBL).
3.4.3 Cloning and resurrecting extinct animals
One aspect of palaeontological science that consistently arises in video games is cloning/de-extinction. The Jurassic Park (Crichton, 1990) franchise popularised the concept of extracting genetic information from fossiliferous material in order to clone ancient animals (for a review of this topic see Jones, 2018). This concept pervades video games based on the franchise, such as Jurassic World: Evolution, but is also prevalent in many other video games. For example, in Terraria (2011, Re-Logic), ancient amber is one of many natural resources that can be collected by the player, and each piece has a tiny chance of containing a fossil mosquito. The player can then process the fossiliferous amber using an “extractinator” to resurrect a pet baby dinosaur. In Pokémon Red/Blue, the player receives a fossil as a reward and can take it to a laboratory on Cinnabar Island, where a scientist will resurrect the extinct Pokémon. This is taken to the extreme in Pokémon Sword and Pokémon Shield (2019, Game Freak), where the palaeontologists take the player's fossils, splice them together, and resurrect them into horrific, often unviable, chimaeras.
Because the trope of DNA extraction from fossiliferous material is so prevalent in media, video game players may not be aware that, currently, we are not able to extract viable genetic material from fossilised remains to “resurrect” extinct animals (although fragmentary fossil DNA can be extracted from some more recent fossils and has allowed important scientific advances in identifying these fossils and placing them in the tree of life (e.g. Buckley and Collins, 2011; Orlando et al., 2013; Perri et al., 2021)). Generally, there is very little nuance in video games regarding the depiction of the growing field of palaeo-proteomics (i.e. the study of ancient proteins) and the recent advances made in the last decade (see Buckley et al., 2018). Furthermore, there is very little discussion in any video games about the ethical considerations of de-extinction (bioethics; see Attwood, 2021), and this should be considered by science communicators when discussing this popular and exciting field of science.
3.4.4 Keeping up with science
Most media is dated by the science it is based on. A classic example is the Jurassic Park franchise, which was lauded for its exciting and scientifically accurate dinosaurs when the first film was released in 1993. However, as the series has progressed, it has chosen to maintain a distinct aesthetic rather than keeping up to date with the current scientific understanding (Polo, 2015). Similarly, while video games can be limited by graphical constraints (due to the computing power of the consoles or graphics cards they are designed for; see Fig. 5), many games choose to be scientifically inaccurate because of the need for recycled assets or because they have a specific aesthetic (e.g. Jurassic World: Evolution). However, unlike other traditional media, PC and (most) game consoles are connected to the internet, meaning game developers (and even fan communities) can issue updates and “mods” (modifications) that update gameplay, graphics, or visual assets indefinitely after the games are released. Therefore, new fossil discoveries can be incorporated into game updates to keep games scientifically up to date. However, this relies on the game developer issuing updates, which may not be viable due to financial or time constraints. One example of this is Animal Crossing: New Horizons, which was released in March 2020. The game contains a skeleton of Spinosaurus that the player can find, accession, and mount in their museum (similar to Fig. 4t). Three weeks after the games release, however, Ibrahim et al. (2020) was published, detailing a new Spinosaurus specimen from Morocco with a previously unknown tail section of significant scientific and public interest. This hype caused considerable media interest in whether Nintendo Entertainment Analysis & Development Division (EAD), the game developers of Animal Crossing, would update the visual asset in line with this new discovery (Watts, 2020; Rochlin, 2020). As of publication, the Spinosaurus model in Animal Crossing has not been updated despite multiple updates to the game.
Inevitably, any released COTS video game will be overtaken by palaeontological advances and rendered outdated, especially if developers are no longer working on updates for the game. From a science communication perspective, outdated game design can still be used to engage with the public, allowing science communicators to communicate up-to-date science and also the history of scientific thought as well as to explore the ever-shifting process of scientific discovery.
3.4.5 Depicting palaeontological fieldwork
Collecting raw materials is an extremely common game mechanic, and games often use fossils as a resource. In most of these games, fossils are acquired by approaching specific types of rock and breaking them with a tool (e.g. The Sims 4; Stardew Valley, No Man's Sky) or by digging holes in specific locations (e.g. Animal Crossing: New Horizons; see Fig. 3c). In Red Dead Redemption 2, fossils can just be collected off the ground (albeit often in hidden or difficult-to-reach locations), while fossils can be found in Stardew Valley by fishing in streams in fossiliferous areas. Because fossil extraction is so common and often simple in video games, it can give players the impression fossil extraction is effortless, unmethodical, and skill-less – quite the opposite of the time-consuming, laborious, and often hazardous excavations that are often required to extract fossils.
Video games can also create the incorrect perception that fossils are a common occurrence in all types of rocks and that a destructive approach is required to extract fossil material. This can be problematic, as it may not be obvious to amateur fossils hunters that using hammers on fossiliferous rock faces may cause irreparable damage in addition to being potentially dangerous and, in some areas, illegal. Recently, local government organisations have started to introduce ethical rock collection policies (see Scottish Geodiversity Forum 2017), which can be disseminated to the public by science communicators.
Some games do take a more nuanced approach to fossil collection. In Starbound, fossils can only be uncovered with specialised tools, such as brushes and rock hammers, and failure to carefully extract the fossil-bearing rock permanently destroys the fossil. Once collected, fossils can be placed in customisable display cabinets at the player's base, which features a placard with basic trivia. Tap! Dig! My Museum! (2019, Oridio) is a mobile game where the player manages a museum but also takes part in prospecting and excavating dinosaur skeletons. By tapping the screen, the player can remove layers of rocks piece by piece to find individual bone elements. Although simplified and highly stylised, the game does represent the processes of uncovering and transporting fossils in a fieldwork setting.
Another aspect of palaeontological fieldwork that is not commonly addressed in video games is ethical issues, which are highlighted below.
3.4.6 Representation of ethics in palaeontological video games
Palaeontology has a long, colonial history with deep-rooted extractive and exploitative practices (Monarrez et al., 2021). Because many harmful practices (e.g. the erasure of indigenous contributions, the illegal acquisition/removal of fossil material from their country of origin by higher income countries, the refusal to repatriate fossil material – see Monarrez et al., 2021; Raja et al., 2021; Cisneros et al., 2022) are so ingrained within palaeontological science, they often appear within video games unchallenged, with little objective analysis. For example, to clone more dinosaurs and expand your park in Jurassic World: Evolution, the player must send dig teams to sites across the globe to extract more fossil genetic material. These dig sites are based on real-world localities, which is a good educational tool. However, many of these sites have strict laws governing the removal and sale of fossils, such as in Mongolia, where it has been illegal to remove fossils from the country for almost 100 years. This is ignored in gameplay, obscuring a complex legal and ethical situation players could learn from.
While finding and collecting fossils is an integral part of the enjoyment of palaeontology and is important scientifically, one of the most contentious ethical issues facing palaeontology is the buying and selling of fossils (Shimada et al., 2014). The commercialisation of fossil material, especially over the internet and in high-profile public auctions (Shimada et al., 2014), directly leads to a myriad of issues, including the destruction of fossiliferous sites by illegal fossil hunters (DeMiguel et al., 2021; Murphy, 2007), samples being lost to science (Shimada et al., 2014), and in the worst case, the illegal exportation and smuggling of fossil material (e.g., Pérez Ortega, 2021), which has dire real-world consequences, e.g. fuelling humanitarian crises, such as in Myanmar (Dunne et al., 2021; Raja et al., 2021). There is increasing awareness of the problems of fossil commercialisation; yet, in virtually every game featuring fossils as collectibles, excess fossils exist purely to be sold for profit. For example, in Jurassic World: Evolution, the game is designed so that any non-dinosaur fossil found by the player must be sold instantly to raise funds. In the space exploration game No Man's Sky (2016, Hello Games), the players can journey between planets and find fossils on the worlds they visit; however, these fossils serve no function other than to be sold for in-game currency. Even in games that otherwise represent museum curation and fossil finding in a positive and educational manner, such as Animal Crossing: New Horizons, fossils are among the most financially valuable natural resources available. Once a fossil has been donated to the museum by the player, any duplicates found can be sold at the local market for in-game currency. These games do little to challenge unethical practices and normalise the commercialisation of fossiliferous material as a standard procedure within the palaeontological community. We therefore suggest that it is vital for science communicators to be aware of the impact of the illegal trade in fossils and to incorporate issues such as ethical fossil extraction and fossil ownership into public engagement to counter this narrative where possible.
3.5 Male, pale, and stale: the depiction of palaeontologists in video games
COTS video games introduce the public to palaeontologists and allied workers, however; the diversity and representation of palaeontologists in video games is very limited.
Often scientists and/or palaeontologists are portrayed in video games as old, white men that often resemble Charles Darwin (e.g. Professor Snail in Stardew Valley). Another common trope is the perpetuation of the “Indiana Jones stereotype” (Fitzpatrick, 2020). For example, Dinosaur Fossil Hunter (demo released 2020, Pyramid Games) is a “palaeontologist simulator” that aims to simulate fossil discovery, digging, and extraction (even tasking players with jacketing fossils in plaster). However, at the time of publication, the game uses only a white male character model to market the game – complete with stereotypical fedora. Not only does the reliance on these characters reinforce the perception of palaeontology as a white-male-dominated science (Panciroli, 2017), it also propagates the “brilliant lone scientist” trope (Black, 2019), failing to illustrate the requisite collaborative nature of actual palaeontological science and fieldwork.
Some palaeo-video games do have more diverse casts. However, these characters are always able-bodied, often poorly fleshed-out characters and are typically plagued with questionable motives and ethics. For example, in Jurassic World: Evolution, the player must liaise with Dr. Kajal Dua, the head of the science division. However, her character has almost no backstory and often undermines her position as a scientist by tasking the player to undertake ethically dubious missions, such as cloning dinosaurs specifically bred to be hyper aggressive when fighting other dinosaurs. Likewise, the cowboy simulator Red Dead Redemption 2 has the player meet Deborah MacGuiness, an older white woman, digging up fossils on the American frontier of 1899. The player learns that she, like many women in the nineteenth century, was rejected from several universities and that no other academics take her scientific ideas seriously. She resembles, and is most likely inspired by, Mary Anning, the most famous of the multitude of female palaeontologists overlooked by their male peers. After the player collects a number of fossils from across the game world to help her with her research, she assembles a biologically impossible chimaera and reveals that she was a terrible scientist all along, thus reinforcing the ”unethical genius” and ”incompetent woman” tropes (see Weingart et al., 2003). The game could have included some genuine information about the infamous “Bone Wars” of early American palaeontology and about women fighting for their place in science; however, by including a female character who validates every rejection she was served, a potentially interesting characterisation and story (and the potential to educate players on such) was lost.
Historically and stereotypically, non-disabled white men have dominated the narrative of palaeontology, and this in turn has shaped the depiction of palaeontologists in media, especially video games. The systematic lack of diversity in the earth sciences, especially palaeontology, is well known (Bernard and Cooperdock, 2018; Warnock et al., 2020). One potential aspect (amongst many) contributing to the lack of diversity in palaeontology is the lack of minority role models (Panciroli, 2017). The poor representation seen in COTS video games fails to counteract this – if minority players, who may be prospective students and future scientists, do not see role models represented in games and only see white male faces, they may be discouraged from pursuing their interest in earth science. This is especially relevant to games that are marketed as “palaeontologist simulators” but only contain white, non-disabled, male characters.
3.6 Perpetuation of harmful, misogynistic, and racist tropes in palaeo-games
Video games have a history of perpetuating problematic tropes, especially ableism, sexism, misogyny, and racism (Nakamura, 2019). Although these tropes are not specific to palaeo-video games, they do warrant attention from science communicators, as they can have an impact on how palaeontology may be perceived by players.
A common problematic video gaming trope is the hypersexualisation of women (Beasley an Standley, 2002). Evidence shows that the negative representation of women in video games can have severe negative effects for female gamers, including self-objectification and low levels of self-efficacy (see Gestos et al., 2018; McCullough et al., 2020). One of the most egregious examples of casual hypersexualisation seen in a palaeo-video game is found in Trespasser (1998, DreamWorks Interactive), where the player's character, Anne, is stranded on a dinosaur-infested island. The game does not feature any visual cues for the player (i.e. health or ammunition counters), and so, in order to check the health of the character, the player must actively look down to check a heart-shaped tattoo on her breast that fades as the player is closer to death. Analysis has shown that the hypersexualisation of women video game characters has decreased since the 1990s (Lynch et al., 2016), with a notable example being the de-sexualisation of Lara Croft, the protagonist of the Tomb Raider franchise, in recent games. While female lead characters are becoming more commonplace (e.g. Aloy, the lead character in Horizon Zero Dawn), it should be noted that Lynch et al. (2016) found COTS video games still typically depict female characters in secondary roles and that these characters are more sexualised than their male counterparts. Furthermore, within palaeontological video games, sexist and misogynistic tropes can also be coupled with racism, often thanks in part to the perpetuation of the “lost world” fantasy genre, which pervades palaeo-adjacent culture. Based on Arthur Conan Doyle's book The Lost World (1912), wherein an ancient ecosystem is found in the Amazon basin by white explorers, this genre combines palaeontology, colonialism, and the mystification and exotification of indigenous cultures (see Harrer, 2018). Many aspects of this sub-genre are harmful, propagating damaging stereotypes, economic oppression, and cultural appropriation. Care should be taken by scientific communicators to not disseminate these damaging tropes if using video games as part of their engagement and also to take further action by actively highlighting and challenging these practices within palaeontology-themed media.
Video games are culturally prolific, and this sector is rapidly becoming one of the largest entertainment markets in the world. Palaeontological themed video games are extremely popular, and because of this, increasing numbers of the public are exposed to ancient animals and palaeontological science, far eclipsing engagement efforts undertaken by scientists and allied palaeo-workers. Many COTS video games contain elements of good science communication – and some games, especially dinosaur simulators, strive for scientific accuracy. Indeed, aspects of palaeontological-themed COTS video games can be used by science communicators to highlight, engage, and educate the public regarding core concepts of palaeontological science. However, as with most types of media, palaeontological video games can contain a suite of negative and potentially damaging tropes. Many of these tropes are widespread issues in the gaming industry (e.g. poor representation, monsterification, hypersexualisation), but their presence in palaeontological video games specifically detracts from their use as science communication tools. Science communicators, therefore, should undertake careful examination of COTS video games before using them as educational tools. Furthermore, because palaeontological-themed video games are so popular, science communicators should be aware of and challenge harmful tropes in their engagement and outreach efforts to the public and, if possible, to COTS video game developers. By raising awareness of these damaging tropes, we can relegate them into extinction.
Data regarding the number of PC games depicting palaeo-science, their pricing and popularity were obtained from SteamDB (https://steamdb.info/, last access: February 2022, SteamDB, 2022) or by accessing the Steam Store directly (https://store.steampowered.com/, last access: February 2022, Valve, 2022). These are free, online databases and storefronts. The remaining data were collected by directly playing individual games, which cannot be shared freely; however, all games used in this study are publicly available to purchase and download.
The supplement related to this article is available online at: https://doi.org/10.5194/gc-5-289-2022-supplement.
TC and JA conceptualised the project and led the investigation, formal analysis, data curation, project administration, production of figures and writing of the original draft. All authors (TC, JA, TC, RPD, and VR) developed the methodology, resources, and contributed to the reviewing and editing process.
The contact author has declared that none of the authors has any competing interests.
The work presented is original and reflects the authors’ views. Ethics approval and informed consent were not sought; this study does not deal with sensitive data or human participants.
Publisher’s note: Copernicus Publications remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
We would like to thank Edward McGowan and Jazmin Scarlett for inspiring this paper with their recent work on volcanoes in video games, and also to thank Jazmin for the fruitful discussions during the early draft phases of this paper. We would also like to thank the Palaeocast Gaming Network (PGN) Group for the feedback and ideas. A. Clements and E. Dunne are thanked for proofreading. Thanks to Flavia Strani, Elsa Panciroli, and Andrea Villa for their thoughtful reviews, which have enhanced this manuscript.
Funding acquisition: This project is not directly funded; however, Thomas Clements is funded by a Leverhulme Early Career Fellowship (ECF-2019-097). Jake Atterby is supported by a Royal Society Research Grants for Research Fellows RGFR̃11̃81021. Valentina Rossi was supported by a Research Südtirol/Alto Adige grant (CUP H32F20000010003).
This paper was edited by Leslie Almberg and reviewed by Elsa Panciroli, Flavia Strani, and Andrea Villa.
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