TY - JOUR AU - Shelomi,, Matan AB - The video game industry has grown dramatically since its beginnings in the early 1950s. Games today have production values and expenses rivaling those of movies, and sales exceeding them. In the U.S. alone, consumers spent US$30.4 billion on video games in 2016 (Entertainment Software Association 2017), compared to only US$11.4 billion at movie theaters (Motion Picture Association of America 2017). Video games are thus a major player (no pun intended) in the entertainment industry. Like film and literature, video games both reflect and influence the cultures that produce and consume them, and can also be studied as a narrative medium (Frasca 1999), including cultural studies on the symbolism of living things depicted therein (Cassel 2016). This article is a cultural entomological review of insects in video games, first presented at the “Insects and the Global Human Experience” symposium at the 2016 International Congress of Entomology. The title incorporates the term “ludology” (from the Latin ludere, “to play”), meaning the study of games in general, such as rules of play or the functioning of software (Frasca 1999). My goal for this study was to review the myriad forms in which terrestrial arthropods appear in video games (Cassel 2016). How do societal views about different insects, positive or negative, shape whether and how they appear in games? This analysis, spanning video games from their origin to the present, highlights general trends and some iconic or significant representations of gaming’s entomology-related content. Read the Manual For this review, I compared all recorded video games from the time of their invention in the 1950s (two decades before 1974’s Pong) until 2018, identifying any with potential entomological references. I mined the video game listings on Wikipedia.org and the comprehensive database of nearly 100,000 games on MobyGames.com for any game with an entomological name or details suggesting entomological content. I checked all games to verify that they actually involved insects; games with “bug” in the title surprisingly often referred to the Volkswagen Beetle. When possible, I scanned online gameplay videos, transcripts, and screenshots for entomologically relevant content. I also found appropriate video games using the entomology-related tropes at tvtropes.org, a wiki-type website that compiles media tropes and attempts to list all examples of each. In this way, I could find games with entomological content even if their content was otherwise inaccessible. Mobile games (games for smartphones and tablets only) and browser games (Internet-only games) were excluded from this review, because their production is unregulated and they are far too numerous to analyze. I included arachnids and myriapods in the analysis, given their presence in entomological publications and their similar role in the layperson’s idea of a “bug”: a terrestrial, multi-legged arthropod of a potentially pestiferous nature (Cassel 2016). I chose not to include crustaceans because of their dissimilar cultural role and the sheer abundance of marine-themed video games. I’ll leave a review of the depictions of crustaceans in video games to aspiring carcinoludologists. In addition to games with “realistic” organisms, I included games with fictitious arthropoid creatures and arthropod-human hybrids. Lastly, I searched for games in which entomologists sensu lato were present, including beekeepers and amateur insect collectors or any character who explicitly “loves insects.” Thus, all examples could be separated into three categories: “realistic” arthropods, arthropoid entities, and entomologists. Insects were further classified according to their role in the game’s mechanics and their “alignment” on a spectrum from positive to negative characterization. I defined three groups: antagonists (enemies, “bad guys”), protagonists (player characters, “good guys”), and neutral or non-player characters (NPCs). Players are not typically interested in defeating NPCs; NPCs may either aid or be aided by the player, or NPCs may not be meaningfully interacted with at all. Some games had insects as items or in the background, which I counted as neutral. Video game franchises (multiple games set in the same universe with many of the same characters, like the Super Mario franchise) have many recurring elements; each game in the franchise may have the same example of an entomological manifestation. However, individual games within the franchise may have a unique example not present in the other games. I merged franchise-wide examples into single entries in my analysis, but I added single-game-specific examples as additional entries (Suppl. Table S1). This article contains screenshots and box art from certain games, which “should be deemed a ‘fair use’ under Section 107 of Title 17 of the United States Code” given their usage in a scholarly, non-commercial work for identification purposes, as expressed by the Digital Games Research Association (Lastowka and Ogino 2014). General Statistics I identified 679 games and game franchises with entomological content (Suppl. Table S1). Of these, 423 used arthropods as antagonists, 143 as protagonists, and 140 as neutral. These numbers are not exclusive; some games had insects in two or three categories. Arthropoid/humanoid creatures appeared as antagonists in 61 games, as player-protagonists in 15 games, and as NPCs in 32 games. Entomologists or insect enthusiasts appeared as antagonists five times, as protagonists 22 times, and as NPCs 13 times. Given the extraordinarily large number of games produced (most with little to no available data about their content), my numbers are underestimations. The number featuring insects as antagonists alone is likely at least twice as high. The earliest video game to feature insects is probably 1976’s Dukedom, a text-based strategy game in which the player must manage resources for a kingdom periodically facing “severe crop damage due to seven-year locusts” (Ahl 1984). The 1978 Sega/Gremlin game Frogs featured edible flies (Frogger came out in 1981), and the 1978 arcade game Gee Bee was a pinball clone with the player as a bee. Possibly the earliest example of arthropoid alien creatures is the 1979 Galaxian franchise, including the classic 1981 Galaga. The famous arcade game Centipede came out in 1980. Insects as Scenery or Symbols Countless games have flying insects in the background to indicate time and/or mood: butterflies by day, moths or fireflies at night, spiders in spooky houses, etc. These can be as simple as a few floating pixels, and none would be mentioned in any game content resources. Some games allow player characters to get stung by wild bees, as in the Animal Crossing series (Fig. 1). Many games have insect themes for certain levels within the game, or sometimes are entirely insect-themed for no particular reason, such as Hive-Sweeper (a bee-themed version of Minesweeper; Fig. 2) and Mega-Bug (an insect-themed Pac-Man, complete with “La Cucaracha” as the theme music). Fig. 1. View largeDownload slide Screenshot from Animal Crossing: New Leaf, in which the player character has been stung. Copyright: Nintendo of America Inc., 2002. Fig. 1. View largeDownload slide Screenshot from Animal Crossing: New Leaf, in which the player character has been stung. Copyright: Nintendo of America Inc., 2002. Fig. 2. View largeDownload slide Official PlayStation Store promotional art for Hive Sweeper, a Minesweeper clone in which the insect theme is purely cosmetic. Copyright: Skylon Games, 2010. Fig. 2. View largeDownload slide Official PlayStation Store promotional art for Hive Sweeper, a Minesweeper clone in which the insect theme is purely cosmetic. Copyright: Skylon Games, 2010. Symbolic use of insects in video games frequently involves a butterfly or moth as a symbol of rebirth or resurrection. A hotel room decorated with suspended moth pupae appears in the Dutch production Darkfall: Lost Souls. Red butterflies in the Japanese horror game Fatal Frame II: Crimson Butterfly lead twin sisters into a forest containing the tortured souls of the dead. Blue butterflies appear throughout the games of the Japanese Persona franchise, often associated with a character named Philemon, drawn from the writings of Carl Jung (1963). Other cases of symbolism are less subtle. In Fable, flies will begin to buzz around player characters whose choices have turned them evil. Spiders are used to symbolize fears or phobias during a nightmare sequence in Dragon Age: Inquisition; selecting a spider reveals text that states which fear it represents, such as “Death,” “Abandonment,” and “Ironically, Spiders” (Fig. 3). Fig. 3. View largeDownload slide Screenshot from Dragon Age: Inquisition, in which a player must defeat symbolic spiders representing fears and phobias, including one that represents, “ironically,” fear of spiders. Copyright: BioWare Edmonton, 2014. Fig. 3. View largeDownload slide Screenshot from Dragon Age: Inquisition, in which a player must defeat symbolic spiders representing fears and phobias, including one that represents, “ironically,” fear of spiders. Copyright: BioWare Edmonton, 2014. Arthropod Antagonists Insects as enemies represent by far the most common use of insects in games. The trope is likely more widespread than I could count; any game featuring monsters or wilderness enemies likely has an arthropod (of varying biological accuracy, but usually giant-sized) to kill at some point. Some franchises have iconic insect enemies: the hymenopteran Zingers of Donkey Kong Country, Wigglers and Buzzy Beetles from the Super Mario franchise, and Bugzzy, the stag beetle enemy from Kirby. Although many games feature a progression in the enemies a player can defeat, I found arthropod enemies throughout the enemy hierarchy, from the small, easily defeated “mobs” that seem to endlessly spawn from everywhere, to large-sized “bosses” that must be defeated at the end of a level, to the most challenging “final bosses,” whose defeat is the goal and endpoint of the game. A giant bee named Hibachi (“fire bee”) is the final boss in every installment of the shooting game franchise DonPachi, whose punny name means “leader bee” and is Japanese onomatopoeia for gunfire. The final boss of the highly successful Final Fantasy X is a tick that wears armor made from harvested souls. The most inexplicable arthropod final boss appears in Growl: the player is a pugilistic forest ranger who punches poachers, and the final boss is a giant myriapod disguised as a man. By far the most common enemy arthropods are stinging hymenopterans, plus whole beehives and generic “swarms” of undefined creatures. Almost as common are arachnids, such as spiders, scorpions, and even solfugids (“canyon spiders” in the Guild War series), especially in games where the player is an insect. The next most common enemy arthropods, somewhat unexpectedly, are antlions (Myrmeleontidae), always as their eponymous nymphal stage and almost always present in desert levels within an overly large sand pit (Fig. 4). Mantids, moths, roaches, and centipedes are also prevalent as antagonists. The post-apocalyptic American Fallout franchise deserves recognition for including the esoteric Panorpidae (scorpionflies) among its realistically rendered arthropod enemies in the form of giant, radiation-mutated Stingwings (though real Panorpidae are stingless). Fig. 4. View largeDownload slide Screenshot of an “earwig scorpion,” likely inspired by larval antlions (Family Myrmeleontidae), from Gradius III. Copyright: Konami, 1989. Fig. 4. View largeDownload slide Screenshot of an “earwig scorpion,” likely inspired by larval antlions (Family Myrmeleontidae), from Gradius III. Copyright: Konami, 1989. The names of some of these enemies deserve praise for entomological effort, including the Neopterons of Monster Hunter, the Phantasmaraneae from Bayonetta, and the mantis/human character Empusa from God of War: Ascension. (Empusa was a Greek demigoddess, fitting that game’s Hellenic theme, but the character’s mantis limbs may have been inspired by the real-world mantis genus Empusa.) Perhaps the most descriptively named character is the termite-inspired enemy from Earthworm Jim: The Evil Queen Pulsating, Bloated, Festering, Sweaty, Pus-Filled, Malformed Slug-for-a-Butt. Pestiferous insects can plague game worlds in varying states of fidelity to their real-world counterparts. The life simulation franchise The Sims features cockroach infestations in homes. Abandoned houses in Constructor are similarly infested, albeit by eight-foot-tall cockroaches. Agricultural pests and crop-dusters feature in SimFarm. In Plague Inc.: Evolved, in which players direct the evolution of a pathogen to eradicate humanity, choosing to have insects vector the germ provides bonus infectivity in warm climates. Less realistically, the medical simulation game Trauma Team [spoiler alert!] features a devastating viral plague spread by the falling scales of migrating monarch butterflies, which had become infected by eating milkweeds that grew near Patient Zero’s corpse. Some games (usually simple games from the 1980s with a focus on mechanics and no real plot) are entirely devoted to pest eradication. The list includes such inspired titles as Bug Busters, Bug Butcher, Bug Bomb, Bug Spray, Bug Attack, Bug Blaster, Bug Blasters: the Exterminators, Bug Hunt, Bug Hunter, Bug Hunter In Space, Bug-N-Out, Bug Bash, BUGS, Splat!, Squash!, and Swat!. In 1983’s Donkey Kong 3, players control Stanley the Bugman, who uses a Flit sprayer to rid his greenhouse of pesky bees and the titular giant gorilla. Games in which the player is explicitly a certified exterminator include Exterminator, Arachnophobia, and Pesterminator: The Western Exterminator, in which the player assumes the role of the mascot of the Western Exterminator Company of Anaheim, California. Impersonating an NPC pest exterminator in 2016’s Hitman allows the player to use insecticide to incapacitate a (human) target. In 2015’s Fly in the House, eradicating a pest is a MacGuffin—a pointless plot device—as the actual gameplay involves smashing as much of the house’s furniture as possible in the process. Arthropod enemies in games are often robots (25 games) and/or aliens (65 games). Usually the plot is a simple “bug war” in which the earthling player must defeat the evil, arthropoid aliens. Note that alien insect races are sometimes protagonists or even playable characters, such as the Zerg in Starcraft or Hivers in Sword of the Stars. Other arthropod-inspired beings in video games include demons and mutants. Special merit for taxonomic diversity goes to the Resident Evil series, whose infected/mutated/undead antagonists borrow traits from fleas, silkworms, stag beetles, cicadas, sea lice, and caterpillars, along with the usual suspects such as roaches, bees, mantids, and spiders. Eusociality Heavily overlapping with science-fiction elements were tropes involving eusocial insects. Arthropoid alien races in games usually have a queen or other central hive mind whose destruction causes their entire society to collapse, such as the titular Metroid. Members of a hive may speak in a plural or third-person voice, such as Mortal Kombat X’s hymenopteran-humanoid D’vorah (Fig. 5), who refers to herself as “This One” (and whose name is Hebrew for “bee”). Joining a hive mind when one is not born into it is usually considered a negative or “Game Over” scenario, as in System Shock. In Combat of Giants: Mutant Insects, the player is an insectoid trying to resist and escape from a eusocial hive mind. Some games’ mechanics are inspired by eusocial insect behaviors, such as colony management in Pikmin (Fig. 6). An interview with game creator Shigeru Miyamoto confirmed that he was inspired by looking at a line of ants carrying leaves and that he “started thinking about a game about lots of small people carrying things in a line, following a leader, with everyone going in the same direction” (LeJacq 2014). Fig. 5. View largeDownload slide Official Steam promotional art for Mortal Kombat X depicting D’vorah, a hymnopteran humanoid. Copyright: NetherRealm Studios & Warner Bros. Interactive Entertainment, 2015. Fig. 5. View largeDownload slide Official Steam promotional art for Mortal Kombat X depicting D’vorah, a hymnopteran humanoid. Copyright: NetherRealm Studios & Warner Bros. Interactive Entertainment, 2015. Fig. 6. View largeDownload slide Official Nintendo eShop promotional art for Pikmin 3, depicting the ant-inspired Pikmin creatures being led in a line. Copyright: Nintendo EAD, 2013. Fig. 6. View largeDownload slide Official Nintendo eShop promotional art for Pikmin 3, depicting the ant-inspired Pikmin creatures being led in a line. Copyright: Nintendo EAD, 2013. Eusociality in games is not limited to Hymenoptera or Isoptera; eusocial mantids appear in Turok 2: Seeds of Evil, eusocial antlions in Halflife 2, and eusocial reptilian-humanoids called “Locusts” in the Gears of War franchise. Eusocial spiders with queens and/or kings exist in at least 10 games, and two games featured eusocial scorpions with kings. Bee kings are present in Donkey Kong Country, Conker’s Bad Fur Day, and Bugdom. Special mention goes to the game Entomorph: Plague of the Darkfall, in which one must manage the politics among several insect races, including a hive of eusocial panorpids (though real panorpids are solitary). Insect Items and Arthropod Allies Weaponized arthropods are wielded by enemies in at least 16 games and by the player or an ally in 49 games. Nearly all involve swarms of Hymenoptera, although locust swarms are used in Diablo III and biocontrol insects in Garden Defense. Swarms are often “summoned” with magic or some other ability, such as the gene-based “plasmids” in Bioshock. Other games feature creatively named mechanical bee-launching weapons, such as AdventureQuest’s Bee Bee Gun or the Ratchet & Clank franchise’s Bee Mine Glove. In other games, the attacks are less elegant, involving the hurling of beehives or jars full of bees. I found two combat apiculturists: in Fist Puncher, The Beekeeper throws bees at her enemies, and in Vigilante 8, Beeswax destroys cars with them. In Metal Gear Solid 3: Snake Eater, an enemy boss called “The Pain” controls hornets, soaks the player in bee pheromones, and can somehow produce metal Tommy guns and grenades made entirely of bees. Some games’ characters can transform themselves into arthropods, willingly or otherwise. Even the famous Mario gets to wear a bee suit to fly and climb on honeycombs in Super Mario Galaxy (Fig. 7). Gigantic arthropods can also be mounts, e.g., giant scorpions in World of Warcraft. At least one insect-racing game exists: Bugriders: the Race of Kings, which takes place on the planet Entymion. Fig. 7. View largeDownload slide Illustration of “Bee Mario” from Super Mario Galaxy. Copyright: Nintendo EAD Tokyo, 2007. Fig. 7. View largeDownload slide Illustration of “Bee Mario” from Super Mario Galaxy. Copyright: Nintendo EAD Tokyo, 2007. The gaming mechanic of edible insects is rare, except in games where the player is also an animal. Metal Gear Solid 3 features entomophagy: Emperor Scorpions are a food item that can restore health. Magical butterflies will restore one’s health in EarthBound, though likely not through ingestion. Helpful arthropoid NPCs exist, often as denizens of an insect-themed level, or as a friendly insect race seeking your help against an unfriendly insect race. Insects often serve as what TV Tropes calls “exposition fairies”: a game mechanic in which the player carries or is followed by a diminutive companion that provides exposition, tutorial information, and hints, and that interacts with other NPCs for voiceless protagonists. Examples include Sparx the Dragonfly from the Spyro series, Issun the Bug (actually a tiny man in a beetle-shell hat) in Ōkami (Fig. 8), and Naggi the Patronizing Firefly in the metafictional Angry Video Game Nerd Adventures. Fig. 8. View largeDownload slide Official PlayStation.com screenshot from Okami, depicting the main character (a wolf goddess) and her “exposition fairy” companion, Issun, a tiny human wearing a beetle exoskeleton. Copyright: Clover Studio & Capcom, 2006. Fig. 8. View largeDownload slide Official PlayStation.com screenshot from Okami, depicting the main character (a wolf goddess) and her “exposition fairy” companion, Issun, a tiny human wearing a beetle exoskeleton. Copyright: Clover Studio & Capcom, 2006. Games that feature multiple anthropomorphic animals also sometimes include insects. Special mention goes to the licensed games in the Sam and Max franchise for Sal, an upright-walking cockroach NPC, whose passionate in-game kiss with a human NPC (complete with saxophone riff) may be the first and only example of a human–arthropod romantic relationship in a video game. Arthropod Player-Protagonists Of the games in which the player controls an arthropod explicitly, rather than a member of an arthropoid alien race, many were brightly colored children’s games (Bug!, Bugdom), educational games (Bee Active), game adaptations of insect-themed licensed products (Maya the Bee, A Bug’s Life, Bee Movie), or early games with simple sprites in which the insect theme is purely cosmetic (Funky Bee, Mr. Butterfly, Bee & Flower). Though Hymenoptera again dominate in these roles, other arthropod protagonists include crickets (Zapper), isopods (Bugdom), moths (Space Moth DX), and fleas (Bugaboo). In the 1997 PC game Koala Lumpur: Journey to the Edge, the cursor is a fly, serving as the only way the player interacts with the game. An important sub-category is the ant colony simulator. The genre-defining game is 1991’s SimAnt: The Electronic Ant Colony, based on B. Hölldobler and E.O. Wilson’s The Ants (Seabrook 2006). In the game, players control the “yellow ant,” one individual in a colony of black ants in a human’s backyard. The ant must collect food, expand the nest, and avoid predators and the human’s lawnmower. When they are hungry, player ants can receive oral trophyllaxis of food from another ant (Fig. 9). The player ant also can lay pheromone trails and recruit other nestmates to follow it to help take down prey and to attack the enemy red ant nest. Players also control the caste ratio of eggs (worker, soldier, reproductive) produced by the queen and how many workers are devoted to foraging, digging, and nursing. Players can expand the colony to other parts of the yard and eventually invade the house, with the goal of eradicating the red ants and driving the human to move out, winning the game. No ant simulator since has been so realistic or entomologically correct; most focus on only one aspect of ant behavior, such as inter-colony combat (Ant General, Ant Raid, Ant War: Domination, Empire of the Ants) or collecting food items (Ant Nation, Ant Queen, Ant Hill, Microsoft Ants). Fig. 9. View largeDownload slide Screenshot from the Amiga version of SimAnt: The Electronic Ant Colony, in which the player protagonist yellow ant receives oral trophyllaxis from a nestmate. Copyright: Ocean Software Ltd. & Maxis Software Inc., 1991. Fig. 9. View largeDownload slide Screenshot from the Amiga version of SimAnt: The Electronic Ant Colony, in which the player protagonist yellow ant receives oral trophyllaxis from a nestmate. Copyright: Ocean Software Ltd. & Maxis Software Inc., 1991. At least one bee simulator, Bizzy Bee, exists, challenging a player to race other bees to collect pollen. Spider simulators also exist, usually as puzzle games involving web-swinging and jumping (Along Came A Spider, Spider: Rite of the Shrouded Moon). In 1996’s Spider, the player is a scientist whose consciousness is inside a cybernetic spider. In some games, such as Eco and Banzai Bug, the player is a nondescript insect. Special mention goes to the Japanese game Mister Mosquito (or Ka in Japanese) for its slew of entomological errors, starting with how the four-limbed male protagonist must stock up on blood so that his family will have enough to last through winter. I should also mention Sim Brick, a Sim Ant parody, in which the player causes a simulated brick to “exist”—a brick appears in mid-air, falls, and squishes an ant. Insect Collectors and Entomologists Collecting insects is the entire point of some games. In the 1985 German game Butterfly, the player is “Otto den Schmetterlingsjaeger” (Otto the Butterfly-Hunter), who uses his net to catch creatures to complete his collection (Fig. 10). Catching a fictional “Technoloptera” to win the 10th Annual Insectathon is the goal of the educational game Elroy Goes Bugzerk. In other games, insect collecting is a side activity, either for the player’s own enjoyment (The Sims 2, Animal Crossing; Fig. 11) or to give to NPCs in exchange for items or currency (Harvest Moon, MySims Agent, Twilight Princess). Amateur entomologists appear in games sporadically, usually as girls or young women. The player’s first target in Hitman: Blood Money is a Chilean drug lord who happens to collect butterflies. In his obituary, he is quoted as saying, “They call the study of butterflies lepidopterology…and that’s enough to scare anyone away from butterflies for good.” Fig. 10. View largeDownload slide Screenshot from Butterfly. Copyright: Eurogold, 1985. Fig. 10. View largeDownload slide Screenshot from Butterfly. Copyright: Eurogold, 1985. Fig. 11. View largeDownload slide Screenshot from The Sims 2, in which a character is collecting an insect. Copyright: Maxis & Electronic Arts, 2004. Fig. 11. View largeDownload slide Screenshot from The Sims 2, in which a character is collecting an insect. Copyright: Maxis & Electronic Arts, 2004. I found only one game in which the player is explicitly an academic entomologist: 1996’s cult classic Bad Mojo. The player is Professor Roger Samms, an unsavory and unethical entomologist who has embezzled millions in research funding, including a generous donation from “The Blattidae Group” of Swiss chemical manufacturers, from his position at the California University at the Barbary Coast (all fictional). As readable documents in his rather filthy apartment reveal (Fig. 12), he’s supposed to be developing a pesticide called “Blataproprid” for eradication of Blattella germanica and Periplaneta americana, but instead he’s planning to abscond to Belize. Samms is the product of a lifetime of negative experiences, starting with his father abandoning him in an orphanage after his mother died during childbirth, and including his thesis advisor dismissing his research: “Your nicotine derivative shows imagination,” she says, “but it is entirely impractical. It won’t work.” However, a magical locket that belonged to his mother transforms him into a cockroach; hence the pun in his name, referring to Gregor Samsa of Kafka’s Metamorphosis. The gameplay involves the player as a cockroach wandering between Roger’s grimy apartment and that of his cantankerous landlord as he tries to reverse the curse. Other insects and animals are allies or hazards in the game, including a roach nymph, a spider, a rideable moth, and a hungry cat. The game’s plot features a soap-opera-worthy twist as Roger’s newfound perspective as a cockroach helps him discover his origin story, plus multiple endings that depend on the player’s choices and successes. Fig. 12. View largeDownload slide Screenshot from Bad Mojo, depicting the player character as an entomologist transformed into a cockroach. Copyright: Pulse Entertainment, 1996. Fig. 12. View largeDownload slide Screenshot from Bad Mojo, depicting the player character as an entomologist transformed into a cockroach. Copyright: Pulse Entertainment, 1996. In a “making of” featurette for Bad Mojo, director Vinny Carrella describes how the creators’ desire for rich, detailed graphics combined with the slow loading limitations of CD-based machines to give rise to the entomological theme. The game called for a small sprite that could move around the detailed backgrounds, and insects fit the requirement. Given the general stylistic themes of the 1990s, a cockroach was a natural choice. “Obviously we weren’t going to do ladybugs or grasshoppers,” says Carrella. “We were thinking of things that had some edge to it.” Once the creative team realized that all of them had had personal “harrowing” experiences with cockroach infestations, they determined that cockroach experiences are “universal” to humans and would produce similar visceral reactions in most players. To achieve the game’s remarkably realistic look, thousands of roaches were reared in a terrarium for a year for the artists to study. Scans of entomological texts seem to have been used in the game as part of the artwork (Fig. 12). The game also prominently features entomologists in the credits. At the top of the “Special Thanks” section are acknowledgments to “Dr. Tom Turpin (Entomologist at Purdue University)” and “Dr. Byron Reid (for his vast insight about roach behavior and physiology),” along with acknowledgments to the University of California at Berkeley Entomology Library and Whitmire Research laboratories. Entomology: The Game(s) Collecting real insects is a relatively popular pastime in Japan, where live beetles kept as pets are traded and/or pitted against each other in fights. These linked hobbies spawned two wildly successful—and lucrative—video game franchises. The combination arcade and card-collecting game franchise Mushiking: The King of Beetles (Fig. 13) was developed by Sega in 2003. Players collect physical cards (either bought in packs or obtained from playing the arcade game), each of which features a real photograph of a beetle and its Japanese common name, Latin scientific name, description, and in-game statistics (e.g., power, stamina, favored move). The beetles are mostly stag and rhinoceros beetles, whose role in Japanese culture has been described elsewhere (Hoshina and Takada 2012). The cards, along with “skill cards” and “title cards” (awarded in real-world competitions organized by Sega that provide embellishing titles to the player’s character name), are scanned into the arcade machines for gameplay. Players use their scanned beetle cards as combatants against computer-controlled beetle enemies or human players, with a rock-paper-scissors style combat system (hitting-pinching-throwing in this case) in a tournament or tag-team system. Mushiking has been re-released several times, spawned a manga (Japanese comics) series, and inspired two Japanese pro-wrestling gimmicks. The game was officially entered into the Guinness Book of World Records on 30 November 2007 for the most official tournaments (>100,000) for a game. Fig. 13. View largeDownload slide Mushiking arcade cabinets. Photo released to the public domain by “Mushiking” on English Wikipedia. Fig. 13. View largeDownload slide Mushiking arcade cabinets. Photo released to the public domain by “Mushiking” on English Wikipedia. In Mushiking, the player is a forest fairy named Popo, friend to all insects, the strongest of which is a powerful Japanese rhinoceros beetle (Scarabaeidae: Allomyrina dichotoma) he calls Mushiking, although the beetle is too modest to accept the title. The antagonist is Adder, an evil wizard who raises an army of invasive, mutant beetles “from another world.” Besides A. dichotoma, the game includes the Thai Rhaetulus crenatus speciosus, the continental Asian Eupatorus gracilicornis, the African Mesotopus tarandus, and the South American Megasoma actaeon. Other insects such as swallowtail butterflies and honey bees can occasionally help the player. The insects in the game are anatomically realistic and entomologically correct, with great attention to detail, although their roaring sounds and combat abilities have been considerably embellished. No mention of entomological gaming is complete without a reference to the Pokémon franchise (Schmidt-Jeffris and Nelson 2018). What started as a pair of video games for the Game Boy (Nintendo’s portable, hand-held gaming hardware) in 1996 became a massive media franchise, with multiple video games in multiple formats, as well as associated trading card games, manga, television, and movies. It is the second best-selling gaming franchise after Mario, and the highest-grossing media franchise in the world (worth an estimated US$92 billion as of December 2018), beating out the entire Mickey Mouse & Friends media franchise by more than US$20 billion. Briefly, players called “trainers” catch wild creatures called “Pokémon,” usually at the behest of a “Professor” character, with the stated goal of catching them all from many different ecosystems and training them for non-lethal combat. Battling facilitates the capture of wild Pokémon by weakening them and is also done for sport against NPC trainers. The Pokémon do not seem to mind this and form deep bonds of friendship with their trainers. In the real world, players not only battle each other (professional Pokémon game tournaments exist), but also trade Pokémon they have collected in the games with other players through various means, depending on the hardware of the game. Trading is an integral part of gameplay, as some Pokémon can only be obtained in this way. The similarity of this mechanic of collecting and trading to how insects are collected and traded by enthusiasts in Japan is intentional. Pokémon is an insect-collecting simulator for urban youth (Fig. 14); it is entomology as a game (Schmidt-Jeffris and Nelson 2018). The franchise creator, Satoshi Tajiri, has attested that as a child in what was then rural Machida, Tokyo, he loved collecting insects and sharing his finds with friends, who also were collectors. He was the most successful and passionate of his peers, earning him the nickname “Dr. Bug.” His hobby would become more difficult as Japan became urbanized and the habitats where he once collected were turned to lifeless concrete. He was an aspiring entomologist until he discovered video games in the late 1970s and became fully engrossed in them, eventually building a reputation as a game designer. In 1990, Tajiri came up with Pokémon, inspired by the Game Boy and its link cable, which was meant to allow two players to connect their devices and play against each other. The apocryphal story is that Tajiri imagined insects crawling across the cable from one Game Boy to another and was inspired to create a game in which players used the link cable to trade rather than to fight. He successfully pitched the idea to Nintendo, and after six years the game was released, eventually becoming what it is today. In interviews, he describes his insect-collecting childhood as the driving force behind Pokémon (Larimer 1999): “Places to catch insects are rare because of urbanization. Kids play inside their homes now, and a lot had forgotten about catching insects. So had I. When I was making games, something clicked and I decided to make a game with that concept. Everything I did as a kid is kind of rolled into one—that’s what Pokémon is.” Fig. 14. View largeDownload slide Official art by Ken Sugimori of insect-inspired “bug type” Pokémon (obtained from the Pokemon.co.jp Pokédex under fair use): A) Caterpie; B) Beedrill; C) Heracross; D) Ribombee; E) Ninjask; F) Yanma; G) Durant; H) Pheromosa; I) Leavanny; J) Ledian; K) Buzzwole. Copyright: The Pokémon Company International. Fig. 14. View largeDownload slide Official art by Ken Sugimori of insect-inspired “bug type” Pokémon (obtained from the Pokemon.co.jp Pokédex under fair use): A) Caterpie; B) Beedrill; C) Heracross; D) Ribombee; E) Ninjask; F) Yanma; G) Durant; H) Pheromosa; I) Leavanny; J) Ledian; K) Buzzwole. Copyright: The Pokémon Company International. Summary The representations of arthropods in video games are highly diverse, as in the real world, but trends exist. Early games with limited graphics or little room for plots mostly used insects as cosmetic themes. We see arthropods predominantly used as adversaries, especially the clades that most evoke fear and panic: aposematic, stinging Hymenoptera and venomous, widely reviled arachnids. Just as the creators of Bad Mojo chose roaches because the entire design team had had bad experiences with them, Hymenoptera and arachnids are widely used because many people are aware of their venomous nature and fear them. Striking these visceral fears is effective, almost too much so. Arachnophobic players of the games Dragon Age, System Shock, and The Elder Scrolls V: Skyrim requested modifications and patches to remove the spiders because they were too frightening. I would hypothesize that, should a herpetoludology review ever be done, snakes will appear in games far more frequently than lizards or frogs. Not all examples are negative, however. Arthropods or arthropoid creatures can be allies, friends, or playable characters themselves. Hymenoptera are the most common friendly NPCs as well as the most common antagonists, because people are as familiar with bees’ importance as pollinators and garden visitors as they are with their sting. Several game characters are explicitly described as having positive views of insects, even if it has little bearing on gameplay or plot. The pro-entomology examples are not all from educational or niche games, nor do they appear to correlate with any particular country or decade of origin; they can appear anywhere, at any time. In fact, some of the most commercially successful and critically acclaimed games are those that draw upon insects as a positive inspiration for their game mechanics, such as SimAnt, Pikmin, and Pokémon. The latter could be considered the crown jewel of all of cultural entomology, having packaged the niche hobby of collecting insects into a popular and profitable form beloved by millions. Many real-world biologists are Pokémon fans. Pokemon (without the accent) was the original name for the cancer-causing gene Zbtb7—short for Poxvirus and zinc finger and Krüppel [POK] domain erythroid myeloid ontogenic factor—until Pokémon USA, Inc., threatened the scientists with legal action (Simonite 2005). A non-oncogenic protein, EGFLAM, is still known by the name Pikachurin, after the franchise mascot Pikachu (Sato et al. 2008). Entomologist Spencer Monckton named a species of bee Chilicola (Heteroediscelis) charizard (Hymenoptera: Colletidae) after the fire-breathing dragon Pokémon Charizard (Monckton 2016). Entomologists studying African parasitic wasps named a species Stentorceps weedlei (Hymenoptera: Figitidae) after Weedle, a wasp-larva Pokémon (Nielsen and Buffington 2011). Outside entomology, a genus of flying reptiles, Aerodactylus, is named after the pterodactyl-like Pokémon Aerodactyl (Vidovic and Martill 2014). A recent paper compared Pokémon spawning rates in the mobile game Pokémon GO to cancer initiation in the body for no apparent reason other than to attract readers and/or as a fan homage (Ma and Moraes 2016). Other authors wrote a thoroughly referenced paper titled “The Ichthyological Diversity of Pokémon” for The Journal of Geek Studies, using Pokémon to share fish facts with general science enthusiasts (Mendes et al. 2017). I wrote a satirical paper on Pokémon phylogenetics for the science humor journal Annals of Improbable Research more than five years ago (Shelomi et al. 2012), and I still get letters requesting copies of its phylogeny for use in teaching evolution or as gifts for scientist fans. Arguably, no game has touched the hearts of biologists so strongly, and it all started with a young boy collecting beetles who happened to later pursue a career in game design. Within entomoludology, what future questions could we ask? What percentage of games correctly show insects with six pairs of legs? Do “bug war” games make people hate insects, or do people who hate insects play “bug war” games? Is Bad Mojo’s Roger Samms a hurtful stereotype, or should entomologists be proud to have any representation in games at all? I won’t speculate on these mysteries for now, but I hope this review can serve as a start for any further forays into the field. Let the games begin! Acknowledgments I thank Dr. Gene Kritsky for encouraging the study of cultural entomology within ESA and organizing the “Insects and the Global Human Experience” symposium at ICE 2016, Dr. May Berenbaum for her strong encouragement to publish this work, and all those who post gameplay videos and data online for the common good. References Cited Ahl , D.H. 1984 . Big computer games . Creative Computing Press , Morris Plains, NJ . Cassel , C.S. 2016 . Bugs after the bomb: insect representations in postatomic American fiction and film. 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This article is published and distributed under the terms of the Oxford University Press, Standard Journals Publication Model (https://academic.oup.com/journals/pages/open_access/funder_policies/chorus/standard_publication_model) TI - Entomoludology: Arthropods in Video Games JF - American Entomologist DO - 10.1093/ae/tmz028 DA - 2019-06-07 UR - https://www.deepdyve.com/lp/oxford-university-press/entomoludology-arthropods-in-video-games-eSmwxas0ih SP - 97 VL - 65 IS - 2 DP - DeepDyve ER -