Cephalopods of the Multiverse

Mark A. Carnall

Oxford University Museum of Natural History. Oxford, UK.

Email: mark.carnall (at) oum.ox.ac (dot) uk

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Magic the Gathering (MTG) is a popular trading and collectible card game, first published by Wizards of the Coast in 1993. Although the game now spans many formats and game types, the core concept pits two players “Planes-walkers” against each other, drawing power (mana) from plains, swamps, mountains, forests and islands to summon creatures and cast spells to battle and defeat opponents. The game has a complex and ever evolving set of rules. Wizards of the Coast regularly release new sets and blocks introducing new cards, mechanics and lore to the rich Multiverse, the planes of existence that Planeswalkers can travel between, that makes the games setting.

One aspect of the game which arguably underpins the continued success of MTG is the vibrancy and colour which gives flavour to the complex ruleset of the game. Storylines featuring several recurring characters, normally Planeswalkers, are told across novelisations, through flavour text and the beautiful artwork of the cards. The designers and artists liberally take inspiration for the denizens of the Multiverse from wider science-fiction, fantasy and of course the natural world.

Although your average game of MTG may feature battles between Inexorable Blobshammer wielding cat wizards and goblin bombers, more zoologically minded Planeswalkers may summon an AllosaurusHammerhead Shark or a Grizzly Bear or two to the fray. Of course, as numerous Journal of Geek Studies papers have highlighted (Salvador, 2014, 2018; Cavallari, 2015; Salvador & Cunha, 2016), cephalopod molluscs have also inspired the designers of MTG and this paper will look at known cephalopods from the Multiverse with some comments on differences between their biology and the cephalopods we’re more familiar with on our humble plane.


‘Squid’, octopuses and nautiluses have all featured in MTG so far on creature, other spell and even Planeswalkers cards. Krakens are also a creature type within the Multiverse but differ from the Kraken of historical and contemporary mythology, normally associated with giant squid or squid-like creatures. In MTG krakens are giant, island destroying, beasts which show a diversity of cetacean, arthropod and molluscan features amongst others. For this reason, krakens get an honourable mention here but won’t be examined as the mutating magical powers of the deep sea defy current systematic reasoning.

Mirroring trends in scientific research and literature on cephalopods, although they are culturally important organisms they make up a small niche of known creatures in the Multiverse. Unlike other creature types which have been a mainstay in MTG sets, cephalopod cards are comparatively rare. Cephalopod-themed cards were published as early as 1997 but it’s only comparatively recently that enough cards have been produced to attempt an all-cephalopod themed standard 60-card deck.

The different cards will be examined in a hybrid taxonomic and card type order starting with creature cards then moving onto enchantments, Planeswalkers and sorcery types. In total, excluding reprinted cards and art variants, there are 21 cephalopod-themed cards currently published for MTG: 14 creatures, 2 sorceries, 2 enchantments, 2 tokens and 1 Planeswalker.


In MTG the comparative power, strength and endurance of different creatures is expressed as a number on the bottom right hand of creature cards. The numerator represents the power of a creature (the amount of damage it can do by punching, slicing, psychically tormenting or oozing on a defending creature) and the denominator represents toughness (the amount of punching etc. it can take).

The power levels of various creatures of the Multiverse is the subject of much debate and mirth amongst players but for this paper the Grizzly Bear with the power/toughness 2/2 will be used as a baseline to make inferences about analogies between cephalopods from other planes and our own.


Perhaps unfairly maligned as hangers-on or ‘living fossils’ on our plane, today’s diversity of living species of nautiluses, the only externally shelled cephalopods, have inspired philosophers, artisans and scientists for centuries. The exact species diversity and relationships between them is still in flux, compounded by the difficulty in accessing and studying these organisms.

There are just two nautiluses in MTG, the Chambered Nautilus, which shares its name with a generic name used to refer to the whole living group, or sometimes, specifically Nautilus pompilius, and the Crystalline Nautilus (Fig. 1). Much like living nautiluses, which are nationally and internationally protected by law, the flavour text for chambered nautilus suggests that their shells are also exploited by jewellers on some planes at least:

“What’s merely a home for the nautilus can become exquisite jewelry in the hands of Saprazzan artisans.”

— Flavour text from Chambered Nautilus card.

Chambered nautiluses are 2/2 creatures in MTG and the card art shows one giving a merfolk an unwanted cuddle. The art and power level suggests that Magic’s nautiluses are significantly larger than living ones. Interestingly, they share a fleshy hood, numerous tentacles and a lenseless eye complete with iris groove for channelling mucus (Muntz, 1987).

Figure 1. The nautiluses. Source: Gatherer.

By contrast the crystalline nautilus, masterfully depicted by artist Brad Rigney, suggests extreme adaptation unlike that of known nautiloid species. In the first instance, the crystalline nautilus is both a creature and enchantment and is shown with a vivid pearlescent shell similar to polished shells of nautiluses. The soft tissue anatomy is consistent with known species of Nautilus and Allonautilus; however, the crystalline nautilus is shown moving at speed over the surface of the water. This has never been documented in known species and furthermore, from the depiction, the hyponome plays no part in this high speed aquaplaning mode of locomotion. A power and toughness of 4/4 suggests that crystalline nautilus is significantly more durable and powerful than Magic’s chambered nautilus too.


As a general term, squid is often used for decapodiform cephalopods excluding cuttlefish which is not a natural grouping of these soft-bodied cephalopods. There are three squid creatures in MTG and two squid producing creatures. With the exception of Gulf Squid, the squid appear to have corneal membranes and are classified, albeit tentatively, here as myopsid squid.

The three squid creatures in MTG are the FylamaridSand Squid and the intriguing Gulf Squid (Fig. 2). Sand Squid appear the most similar to known myopsid species albeit significantly larger than any known decapodiform cephalopod, depicted embracing a human-sized creature with thick, flat arms. Fylamarids are flying squid which appear to have evolved true sustained flight beyond the shorter bursts of flight in species of flying squid (Muramatsu et al., 2013) with adaptations of large wing like projections underneath the siphon region, huge lateral fins and vampire squid-like filament arms alongside usual arm array. The tentacles appear to have been lost, but they can squirt ink.

Figure 2. MTG’s ‘squid’ cards including the presumably misclassified Omastar Gulf Squid. Source: Gatherer.

Although the Gulf Squid has been categorised as a squid by MTG (presumably informed by scholars from across the Multiverse), the gulf squid possesses a large ornamented spiral shell suggesting an ammonoid affinity or convergence. The direction of shell coiling with relation to the position of the aperture as well as the skin colour, suggests a close resemblance to another well-known fictitious cephalopod (Salvador, 2014). Further study of this group is required to confirm relationship with other known cephalopods from the Multiverse.

Likewise, Chasm Skulkers, categorised by MTG as a ‘squid horror’ also defies known relationships within Cephalopoda. Upon the death of a Chasm Skulker, a number of 1/1 squid creatures are created. It is unknown if these are symbiotic or parasitic cephalopods, who attack on the death of their ‘host’, or spontaneously created with magical forces. The last ‘squid’ card gives some insight into ecology in the oceans of different planes, summoning a Coral Barrier also brings with it a 1/1 squid creature consistent with reef species in our plane.


In terms of types of octopuses in MTG, which in some cases seems to be analogous to species, octopuses are the most speciose of known cephalopods from the Multiverse. There are six octopus creatures. Like cephalopods in our plane, the Multiverse also seems to be plagued with problematic naming conventions when it comes to octopus types.

In order of power, Crafty Octopus (Fig. 3) is the weakest octopus card, but like living species, makes up for it in terms of brain power. In addition to showing an advanced range of tool use, Crafty Octopus is also wearing glasses, steadfast evidence of intelligence in ethological studies.

Figure 3. The octopuses, with fourth wall breaking Jules Verne quote on this printing of the card. Source: Gatherer.

The next octopus in terms of power is the Giant Octopus (Fig. 3), depicted at a size larger than buildings and capable of destroying ships with their arms. Although certainly giant by comparison to the largest known species of octopuses in our plane, the name may be a misnomer as they are the second smallest type of octopus in MTG, and therefore not biologically giant as defined by Klug et al. (2015). The flavour text for the various reprints of this card tell us many things. Firstly, that calamari is appreciated across the Multiverses and secondly with a quote from Jules Verne’s Twenty Thousand Leagues under the Sea, that this influential volume has somehow also made its way across the Multiverse (or perhaps Verne walked the planes?).

Tied at 5/5 power and toughness are the ship-crushing Sealock Monster and multi-mouthed Godhunter Octopus (Fig. 4). Studying specimens of this size would have huge implications for understanding the evolution of colossal size in coleoid cephalopods. From a restricted glimpse of Godhunter octopuses, it appears they possess numerous toothed mouth-like openings, superficially similar to toothed sucker rings.

Moving up the power scale, the Elder Deep-Fiend (Fig. 4) is next, literally bursting from inside another creature which is handy in a pinch. The Elder Deep-Fiend shows some interesting anatomy similar to Godhunter Octopus with a toothed maw on the surface of the mantle rather than in the centre of arms. However, it’s important to note that this octopus is a physical manifestation formed from the ceaseless hunger of titans from the Blind Eternities so adherence to biological principles is not necessarily a given.

Figure 4. The octopod monsters, depicted destroying people, boats and mountains? Source: Gatherer.

The last of the octopus creatures is Lorthos, the Tidemaker (Fig. 5) a whopping and cephalopod-theme pleasing 8/8 legendary creature. Unfortunately, last seen being dismembered by an Eldrazi titan, this unique specimen is presumed lost to science (Digges, 2015).

Figure 5. Lorthos. Source: Gatherer.


In addition to summoning creatures to go head to head with each other in magical conflicts, Planeswalkers can also use a variety of spells to tip the table in their favour and control the field of play. They can also summon other Planeswalkers to assist in battles. There are a number of cephalopod spells in MTG but unfortunately, their magical and ethereal nature defies existing classification systems and biological concepts.

Crush of Tentacles (Fig. 6; although crush of cephalopod arms appears to be more accurate) is a powerful sorcery spell that makes all other creatures disappear and, if you’ve got the mana to spare, summons an 8/8 octopus to boot. Octopus Umbra (Fig. 6) is an enchantment aura that can be used to give other creatures ‘the power of Octopus’ boosting them to 8/8 power and toughness with the ability to shut down creatures with a power less than 8 (see what they did there?).

Then there are two spells and one creature which cause pause for thought on cephalopod taxonomy. Quest for Ula’s Temple (Fig. 6), Whelming Wave and summoning Slinn Voda all affect creature types. Quest for Ula’s Temple becomes a tidal wave of creatures and the other two remove certain creatures from play. Interestingly, octopuses are the only cephalopods affected by these alongside aforementioned Krakens, Leviathans and Serpents. Quite why it’s only octopuses and not all cephalopods which are affected is currently unknown. Interestingly, Whelming Wave summons a… err… whelming wave, but octopuses are spared from its destructive power. This then allows them to take over the land presumably as happened recently in Wales (Ward, 2017).

Figure 6. Cephalopod flavoured spells: Quest for Ula’s Temple, Octopus Umbra, Crush of Tentacles [sic]. Source: Gatherer.

The last cephalopod-themed card worth mentioning is Planeswalker Kiora. A merfolk Planeswalker, she has the power to summon 8/8 octopuses into battle and is depicted in both her Master of the Depths and Crashing Wave (Fig. 7) as keeping a suckered beast or two on hand at all times. A must-have ally for those wanting to literally bring more arms to the fight.

Figure 7. Both depictions of Planeswalker Kiora A.K.A. ‘The one with all the fan art’. Source: Gatherer.


As of the time of writing, these are all the known cephalopod and cephalopod-related creatures, spells and Planeswalkers from the MTG Multiverse. In this examination there is some biological conservatism across planes of existence when it comes to cephalopod biology, anatomy and ecology. There are also some marked differences, which although may be biologically questionable, implausible or indeed impossible, they make for a fun game. There are still plenty of cephalopods yet to draw inspiration from including early fossil forms, cuttlefish, ram’s horn squid and bobtail squid. Here’s hoping that many more cephalopods will be making their way to a card table soon.


Cavallari, D.C. (2015) Shells and bytes: mollusks in the 16-bit era. Journal of Geek Studies 2(1): 28–43.

Digges, K. (2015) The Rise of Kozilek. Wizards of the Coast. Available from: https://magic.wizards. com/en/articles/archive/uncharted-realms/rise-kozilek-2015-12-09 (Date of access 12/10/2018).

Gatherer. (2018) Wizards of the Coast. Available from: http://gatherer.wizards.com/Pages/De fault.aspx (Date of access 12/10/2018).

Klug, C.; De Baets, K.; Kreoger, B.; Bell, M.A.; Korn, D.; Payne, J.L. (2015) Normal giants? Temporal and latitudinal shifts of Palaeozoicmarine invertebrate gigantism and global change. Lethaia 48: 267–288.

Magic: The Gathering. (2018) Wizards of the Coast. Available from: https://magic.wizards.com/en/ new-to-magic (Date of access 12/10/2018).

Muntz, W.R.A. (1987) A Possible function of the iris groove of Nautilus. In: Saunders, W.B. & Landman, N.H. (Eds.) Nautilus: The Biology and Palaeobiology of a Living Fossil. Plenum Press, New York. Pp. 245–247.

Muramatsu, K.; Yamamoto, J.; Abe, T.; Seikiguchi, K.; Hoshi, N.; Sakurai, Y. (2013) Oceanic squid do fly. Marine Biology 160(5): 1171–1175.

Salvador, R.B. (2014) Praise Helix! Journal of Geek Studies 1(2): 9–12.

Salvador, R.B. (2018) One squid to rule them all. Journal of Geek Studies 5(1): 23–32.

Salvador, R.B. & Cunha, C.M. (2016) Squids, octopuses and lots of ink. Journal of Geek Studies 3(1): 12–26.

Verne, J. (1872) Twenty Thousand Leagues under the Seas: A Tour of the Underwater World. Pierre-Jules Hetzel, Paris.

Ward, V. (2017) Octopus invasion on Welsh beach blamed on effects of recent storms. The Telegraph: 29/Oct/2017. Available from: https:// http://www.telegraph.co.uk/news/2017/10/29/octopus-invasion-welsh-beach-blamed-effects-recent -storms/ (Date of access 01/12/2018). 


I’d like to thank ‘Worm Tongue’ Murphy, ‘Tap to Block’ Nick, ‘Read the Cards’ Andy and ‘Bobby’ Big Balls for hours of field testing these ideas and concepts. Special thanks go to the staff of Dark Sphere London for their patience in cephalopod card hunting. 


Mark Carnall is a natural history curator specialising in all living things across time which isn’t really a specialism. As a museum curator he knows better than most that there is no prying apart popular culture and science as they both feed on and into each other. All animals are the best but cephalopods are more best.

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One squid to rule them all

Rodrigo B. Salvador

Museum of New Zealand Te Papa Tongarewa. Wellington, New Zealand.

Email: salvador.rodrigo.b (at) gmail (dot) com

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When it was released in 2014, Middle-earth: Shadow of Mordor (Warner Bros. Interactive Entertainment) proved to be the game all Tolkien fans had been waiting for. Its sequel, Middle-earth: Shadow of War, released in 2017, improved and expanded the first game. Besides all the orc-slaying action, the game has a bunch of other activities, including the most staple of gaming side quests: collectibles.

Simply put, collectibles are items scattered throughout the game and completionist gamers go crazy hunting them. In most games, collectibles do very little or even nothing at all, but in Shadow of War, they reveal little tidbits of the game’s lore. When dealing with any Tolkien-related story, we fans are always happy to have more information about the setting and this makes the collectibles in Shadow of War rather enjoyable.

One of these collectibles, a fossilized squid’s beak, immediately and inevitably caught my attention. Since this fossil deserves more time in the spotlight than what it got in the game, I have devoted this article to analyze it more thoroughly.


The fossil in Shadow of War can be found in Mordor and it represents a squid’s beak (Fig. 1). In the game, the item is called “Kraken Beak Fossil” and is accompanied by the following comment by Idril, the non-player character responsible for the treasury of the Gondorian city Minas Ithil: “Our patrols found this fossilized squid beak years ago. If it is proportional to the smaller squids that fishermen sometimes catch, the sea creature would be several hundred feet long.

Figure 1. The fossilized squid beak found in Middle-earth: Shadow of War. Credit: Monolith Productions / Warner Bros. Interactive Entertainment; screenshot from the game.

The item is named a “Kraken beak” in allusion to the well-known fact that real-life giant squids were the origin of the Kraken myth (Salvador & Tomotani, 2014). So the characters in the game recognize they are dealing with a “giant version” of their common squids. But what exactly is a squid’s beak? And can fossil beaks really be found in our planet’s rocks? To answer these questions, we will need a little primer in squid biology.


Squids are animals belonging to the Phylum Mollusca, the mollusks, and more specifically to the Class Cephalopoda. Cephalopods are very diverse creatures and the group includes not only squids but also octopuses, cuttlefish, nautiluses and two completely extinct lineages: the belemnites and the ammonoids. Cephalopods live in seas worldwide (from the surface to 5,000 m deep) and are represented by over 800 living species; the fossil record, on the other hand, counts with 17,000 species (Boyle & Rodhouse, 2005; Rosenberg, 2014).

The first cephalopods appeared over 450 million years ago during the late Cambrian (Boyle & Rodhouse, 2005; Nishiguchi & Mapes, 2008). They achieved an astounding diversity of species during the Paleozoic and Mesozoic eras, but some lineages (ammonoids and belemnites) are now extinct (Monks & Palmer, 2002). Today, we have two distinct groups of cephalopods: the nautiluses, a relict group with just a handful of species, and the neocoleoids, a latecomer group that appeared during the Mesozoic and includes cuttlefish, octopuses, and squids (Boyle & Rodhouse, 2005; Nishiguchi & Mapes, 2008).

Squids are soft-bodied animals and their body is divided into three parts (Fig. 2): (1) the mantle, where most organs are located; (2) the head, where the eyes, brain, and mouth are located; and (3) the eight arms and two tentacles (the latter usually look different from the arms and can be much longer).

Figure 2. Diagram of a squid, with the names of their body parts. Credit: Barbara M. Tomotani; image modified from Salvador & Tomotani (2014: fig. 7).

The mouth of the squid is on the center of the circle formed by the arms. It contains a pair of chitinous mandibles, which together are called a “beak” because of their resemblance to a bird’s beak (Fig. 3). Squids hold their prey with their arms, draw it towards the mouth, and take small bites off it using the beak. The beak and mandibles move by muscular action – they are connected by jaw muscles within a globular organ called “buccal mass” (Nixon, 1988; Tanabe & Fukuda, 1999).

Figure 3. Example of a squid: a (dead) specimen of Doryteuthis sanpaulensis (Brakoniecki, 1984). Top: whole animal. Bottom left: mouth region (in the center of the ring of arms). Bottom right (upper inset): close-up of the mouth; the beak is barely visible. Bottom right (bottom insets): beak (removed from the specimen) in frontal and lateral views. The specimen is deposited in the scientific collection of the Museu de Zoologia da Universidade de São Paulo (São Paulo, Brazil) under the record number MZSP 86430. Photos by Carlo M. Cunha; image reproduced from Salvador & Cunha (2016: fig. 6).

Usually, the only parts of an animal to become fossils are the mineralized (and thus hard) skeletal structures, such as bone, teeth, and shells. Squids are almost completely soft-tissue animals and so are only preserved in the fossil record in exceptional circumstances. The beak of a squid is not mineralized; rather, it is composed only of organic compounds such as chitin (the same substance found on insects’ exoskeleton) and proteins (Miserez et al., 2008). Nevertheless, the beak is reasonably tough and thus, it can become a fossil under the right circumstances. Indeed, several fossil squids (and neocoleoids in general) are known only from their beaks (Tanabe, 2012; Tanabe et al., 2015; Fig. 4) or their internal vestigial shell[1].

Therefore, it is plausible that a fossil beak of a squid could be found in Mordorian rocks. It could be argued that the fossil presented in the game is not morphologically accurate, especially the frontal part of the beak, which seems to be a single piece instead of two (Fig. 1), but we can disregard this here and accept the Mordorian fossil for what the game says it is: the remains of a squid that lived long ago. The game’s description of the fossil implies that the animal would be huge – but how can we know the size of the animal only from its beak? And how big can a squid get anyway? I will try to answer those questions now.


Besides Idril’s comments about the fossil in Shadow of War and how large the actual animal must have been (“several hundred feet”), we have no real indication of the fossil’s size – no scale bar alongside its depiction, for instance. Knowing the actual size of a squid’s beak allows scientists to estimate the animal’s size, based on data from recent species. For instance, Tanabe et al. (2015), described a new squid species based on a fossilized beak (Fig. 4). They named it Haboroteuthis poseidon and, by its lower beak of roughly 7 cm, estimated it to be the size of a Humboldt squid (Dosidicus gigas d’Orbigny, 1835), with a mantle length of 1.5 m – a giant in its own right. However, nature does not disappoint us in this regard and we have two amazingly huge species, aptly named Colossal squid and Giant squid.

Figure 4. The fossil beak (lower jaw, viewed from several angles) of Haboroteuthis poseidon Tanabe, Misaki & Ubukata, 2015, a squid from the late Cretaceous period (roughly 85 million years ago) of Japan. Image reproduced from Tanabe et al. (2015: fig. 7).

The Colossal squid, Mesonychoteuthis hamiltoni Robson, 1925, is the largest living cephalopod species in terms of body mass. It is very bulky, weighing up to half a ton and maybe even more. The Giant squid, Architeuthis dux Steenstrup, 1857, is actually the largest invertebrate alive – it can reach up to 20 meters (about 65 feet) in length, from the tip of its mantle to the tip of its long tentacles. However, Architeuthis has a slender build and even though it is larger, it weighs less than Mesonychoteuthis. Centuries ago encounters on the open sea with Architeuthis left Nordic seafarers in awe, giving rise to the legend of the Kraken (Salvador & Tomotani, 2014).

But since Idril did not take her time to actually measure the fossil, we cannot estimate the body size of the Mordorian squid. Her estimate of several hundred feet is way larger than the “modest” 65 feet of Architeuthis and extremely unrealistic for any kind of animal (both soft-bodied and with a hard internal skeleton); thus, it can be dismissed as a guesstimate of someone without training in zoology. However, given the large “prehistoric” proportions of other animals in Tolkien’s legendarium, such as wargs and oliphaunts, we could expect the Mordorian squid to be really big – but good old Biology would not allow a much larger size than Architeuthis.

But what about the Middle-earth canon? Did Tolkien provide us with some nice Kraken-like legends to settle this matter?


Judging by videos and forum discussions on the Internet, most of the players that found the fossil in Shadow of War just considered it to belong to a monster akin to the “Watcher in the Water” from The Fellowship of the Ring (Tolkien, 1954a). Of course, that simply cannot be, because the Watcher is not a cephalopod; for starters, he is watching from a pool of freshwater. Its physiology and behavior do not really match those of cephalopods. The Watcher’s physical description (Tolkien, 1954a) is vague enough to match virtually any kind of “tentacled” monster; people just assume it is a cephalopod because of the tentacles[2] (e.g., Tyler, 1976).

In his Tolkien Bestiary, Day (2001) took a huge liberty and gave the name Kraken to the Watcher.[3] Tolkien, however, never mentioned a Kraken (or cephalopods) in his writings – and surely did not relate that name to the Watcher[4] (even in manuscript; C. Tolkien, 2002a).

As Tolkien scholarship is very complex, I reached out to the American Tolkien Society just to be safe. They confirmed the absence of krakens and squid-like beasts in Tolkien’s works (A.A. Helms, personal communication 2017).

We must remember, however, that the video games (including Shadow of War) are not part of the accepted Tolkien’s canon, which includes only the published writings of J.R.R. Tolkien and the posthumous works edited and published by his son Christopher. Games like Shadow of War are thus allowed to deviate from the core works and invent new things to amaze and surprise its players. And one of these things seems to be the fossil giant squid.

Therefore, we can think of Shadow of War’s squid as a new discovery: a new species hitherto unknown to Science. New species discoveries always get the public’s attention, but few people actually know how scientists are able to recognize a species as new and what they do to formally describe and name it. So let us take a closer look at the whole process.


The beaks of recent cephalopods have been widely studied by zoologists (e.g., Clarke, 1962; Nixon, 1988) and so they provide a good basis for comparison when someone finds a new fossil. By comparing the morphological features of the new find with previously known species, it is possible to decide if it belongs to one of them or if it represents a new species.

Now let us imagine that the Mordorian fossil was compared to all known cephalopods and we discovered it is, in fact, a new species. How do scientists formally describe a new species and give it one of those fancy Latin names?

The science of defining and naming biological organisms is called Taxonomy and it deals with all types of living beings, from bacteria to plants to animals. Zoologists have long ago come up with a set of rules for describing new species; it is called the International Code of Zoological Nomenclature, or ICZN for short.[5] We are now in the 4th edition of the ICZN, from 1999. The “Code” gives us guidelines for naming species and for what is considered a good (or valid) species description. For a new species to be recognized by the scientific community, its authors (i.e., the scientists describing it) have to provide a set of crucial information: (1) a description or a diagnosis of the species; (2) a holotype specimen; (3) the type locality; and (4) a scientific name. So let me explain each of these.

The description of a species is very straightforward: the researcher lists all the features (called “characters”) from the species. This includes morphology (e.g., shape, color), anatomy (e.g., internal organs), behavior (e.g., feeding habits, courtship), ecology (e.g., preferred prey), habitat, etc. As Mayr et al. (1953: 106) put it, the characters listed in the description are limited “only by the patience of the investigator”.

The diagnosis, on the other hand, is a list of just those characters that distinguish the new species from all the other species in the same group (like a genus or family). The word “diagnosis” comes from the Greek and originally means “to distinguish between two things” (Simpson, 1961). Both description and diagnosis are written in a peculiar telegraphic way, which will seem very odd for people not used to it.

The holotype is a single physical specimen chosen by the author to be the name-bearing specimen of the given species. That means the scientific name of the species is forever linked with that specimen and this will form the basis for the definition of the species. The holotype should ideally represent the species well, but this is not always the case: it can be an entire animal, such as a squid preserved in a jar of ethanol, or just part of the animal, such as the squid’s beak. The latter case is especially true for fossils, where the whole animal is not preserved. Finally, the holotype should be preserved and kept in a museum or university collection, thus allowing access to anyone interested in studying it.

The type locality is the place where the holotype comes from; the more precise the locality (e.g., GPS coordinates), the better. For fossils, it is also common to indicate the type stratum, that is, the layer of rock where the holotype was found.

Finally, the author gets to choose a scientific name for the species. The scientific names of species are formed by two parts; let us have as an example the species Corvus corax, the common raven. The first part is actually the name of the genus, Corvus, which includes not only ravens but also species of crows, rooks, and jackdaws. The second part of the name (corax) is called the “specific epithet”. However, one should always remember that the species name is not simply corax. The word corax by itself means nothing unless it is accompanied by the genus name. Thus, the complete name of the raven species is Corvus corax.

When choosing the specific epithet, the author can use anything he wants, but most commonly people use a word that denotes: (1) a morphological feature, such as Turdus rufiventris, the rufous-bellied thrush (naturally, rufiventris means “rufous-bellied”); (2) the place where the species can be found, such as the Abyssinian thrush, Turdus abyssinicus (Abyssinia is a historical name for Ethiopia); (3) an ecological or behavioral trait, like the mistle thrush, Turdus viscivorus (viscivorus means “mistletoe eater”); or (4) a homage to someone, like Naumann’s thrush, Turdus naumanni, named in honor of the German naturalist Johann Andreas Naumann (the suffix “-i” in the specific epithet is the Latin masculine singular form of the genitive case). The explanation of where the name comes from is called etymology.

Furthermore, when writing a scientific name, it is good practice to also include the authorship of the species; this means including the name(s) of the author(s) who originally described it. In the example above, the complete species name would be Corvus corax Linnaeus, 1758. Linnaeus is the scientist who first described the species and 1758 is the year he published the description.

So now that the formalities of taxonomy were presented, let us see how our new Mordorian species could be described. If the species in question cannot be placed in an existing genus, a new genus might be described and the same ICZN rules above apply. So let’s start by naming the genus Mordorteuthis n. gen.[6], which reflects the place where the fossil was discovered (“teuthis”, from the Greek, means “squid”).

The new species could then be formally described as Mordorteuthis idrilae n. sp.[7], named in honor of Idril (the suffix “-ae” in the specific epithet is the Latin feminine singular form of the genitive case).[8] The holotype would be the specimen recovered by Talion (Fig. 1) that originally belonged to the treasury of Minas Ithil. For safekeeping, the holotype should then be handed over to a decent academic institution, like the Royal Museum of Minas Tirith (yes, I just invented that). The type locality would be Mordor, close to the Sea of Núrnen; the type stratum, however, remains unknown, as this information is not provided in the game (it is suggested, however, that the fossil was found on a beach of the Sea of Núrnen). The diagnosis should give a list of features (such as its large size) that can distinguish it from other fossil squids from Middle-earth; a hard task, given that this is the very first fossil squid described from Middle-earth. The description would be a full account of the fossil’s shape, proportions, and fine structures; this can be boring even for trained taxonomists, so I won’t do it here (for an actual example, see Tanabe & Hikida, 2010).

Finally, we might glimpse some information about the squid’s habitat: the fossil was found close to the Sea of Núrnen, which is an inland saltwater lake, like our Dead Sea (Tolkien, 1954b). Both the Sea of Núrnen and the Sea of Rhûn to the north are thought to be remnants of the old Sea of Helcar from the First Age (Fonstad, 1991; C. Tolkien, 2002b).[9] The Sea of Helcar would be much larger and thus, perhaps a fitting place for large squids to thrive. Besides, its old age makes it a likely point of origin for a fossil.

Of course, a new species description is only valid if published in the scientific literature. Therefore, our little flight of fancy with Mordorteuthis idrilae here is not a valid species description, but it can sure serve as a nice introduction to taxonomy and to how scientists describe new species.

Finally, it is always worthwhile to mention that several taxonomists have paid homage to Tolkien by naming their genera and species after characters and places from his writings (Isaak, 2014). For instance, we have the genera Smaug (lizard), Beorn (tardigrade), and Smeagol (snail), and the species Macropsis sauroni (leafhopper), and Bubogonia bombadili and Oxyprimus galadrielae (both fossil mammals). But there are many others. That may be inevitable in a sense, as several nerds end up becoming scientists. In any event, geeky names such as these sure make an otherwise arid science a little bit more colorful.


Boyle, P. & Rodhouse, P. (2005) Cephalopods: Ecology and Fisheries. Blackwell Science, Oxford.

Clarke, M.R. (1962) The identification of cephalopod “beaks” and the relationship between beak size and total body weight. Bulletin of the British Museum (Natural History), Zoology 8: 419–480.

Day, D. (2001) Tolkien Bestiary. Random House, New York.

Fonstad, K. (1991) The Atlas of Middle-earth, Revised Edition. Houghton Mifflin Harcourt, New York.

International Commission on Zoological Nomenclature. (1999) International Code of Zoological Nomenclature, 4th ed. The International Trust for Zoological Nomenclature, London.

Isaak, M. (2014) Curiosities of Biological Nomenclature. Etymology: Names from Fictional Characters. Available from: http://www.curioustaxonomy.net/etym/fiction.html (Date of access: 11/Jan/2018).

Mayr, E.; Linsley, E.G.; Usinger, R.L. (1953) Methods and Principles of Systematic Zoology. McGraw-Hill, New York.

Miserez, A.; Schneberk, T.; Sun, C.; Zok, F.W.; Waite, J.H. (2008) The transition from stiff to compliant materials in squid beaks. Science 319(5871): 1816–1819.

Nishiguchi, M. & Mapes, R.K. (2008) Cephalopoda. In: Ponder, W.F. & Lindberg, D.R. (Eds.) Phylogeny and Evolution of the Mollusca. Springer, Dordrecht. Pp. 163–199.

Nixon, M. (1988) The buccal mass of fossil and Recent Cephalopoda. In: Clarke, M.R. & Trueman, E.R. (Eds.) The Mollusca, Vol. 12, Paleontology and Neontology of Cephalopods. Academic Press, San Diego. Pp. 103–122.

Rosenberg, G. (2014) A new critical estimate of named species-level diversity of the recent Mollusca. American Malacological Bulletin 32(2): 308–322.

Salvador, R.B. & Cunha, C.M. (2016) Squids, octopuses and lots of ink. Journal of Geek Studies 3(1): 12–26.

Salvador, R.B. & Tomotani, B.M. (2014) The Kraken: when myth encounters science. História, Ciências, Saúde – Manguinhos 21(3): 971–994.

Simpson, G.G. (1961) Principles of Animal Taxonomy. Columbia University Press, New York.

Tanabe, K. (2012) Comparative morphology of modern and fossil coleoid jaw apparatuses. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 266(1): 9–18.

Tanabe, K. & Fukuda, Y. (1999) Morphology and function of cephalopod buccal mass. In: Savazzi, E. (Ed.) Functional Morphology of the Invertebrate Skeleton. John Wiley & Sons, London. Pp. 245–262.

Tanabe, K.; Misaki, A.; Ubukata, T. (2015) Late Cretaceous record of large soft-bodied coleoids based on lower jaw remains from Hokkaido, Japan. Acta Palaeontologica Polonica 60(1): 27–38.

Tennyson, A.L. (1830) Poems, chiefly lyrical. University of Pennsylvania Press, Philadelphia.

Tolkien, C. (2002a) The History of Middle-earth II. HarperCollins, London.

Tolkien, C. (2002b) The History of Middle-earth III. HarperCollins, London.

Tolkien, J.R.R. (1954a) The Fellowship of the Ring. George Allen & Unwin, London.

Tolkien, J.R.R. (1954b) The Two Towers. George Allen & Unwin, London.

Tyler, J.E.A. (1976) The Complete Tolkien Companion. St. Martin’s Press, New York.


Brown, R.W. (1956) Composition of scientific words. Revised edition. Smithsonian Books, Washington, D.C.

Mayr, E. & Ashlock, P.D. (1991) Principles of Systematic Zoology, 2nd ed. McGraw-Hill, New York.

Salvador, R.B. (2014) Geeky nature. Journal of Geek Studies 1(1-2): 41–45.

Winston, J.E. (1999) Describing Species: Practical Taxonomic Procedure for Biologists. Columbia University Press, New York.

Wright, J. (2014) The Naming of the Shrew: A Curious History of Latin Names. Bloomsbury Publishing, London. 


I am deeply grateful to the people from the American Tolkien Society (Amalie A. Helms, Connor Helms, and Phelan Helms) for the information about “krakens” in Tolkien’s works; to Dr. Philippe Bouchet (Muséum national d’Histoire naturelle, Paris, France) for the help with ICZN articles; and to Dr. Barbara M. Tomotani (Netherlands Institute of Ecology, Wageningen, The Netherlands) and Dr. Carlo M. Cunha (Universidade Metropolitana de Santos, Santos, Brazil) for the permission to use, respectively, Figures 2 and 3 here.


Dr. Rodrigo Salvador is a malacologist who has made his peace with the fact that virtually no one knows what a malacologist is. In case you’re wondering, it means “a zoologist specializing in the study of mollusks”. Despite being a Tolkien fan through and through, he does think that Middle-earth could use more zoological diversity.

[1] Called “cuttlebone” in cuttlefish and “gladius” or “pen” in squids and octopuses, although some lineages have completely lost the shell. Other cephalopods, like the nautilus, have very prominent external shells, as is the norm for other mollusks (e.g., snails, clams, etc.).

[2] Since people always get this wrong, just let me clear things up: squids have 8 arms and 2 tentacles, while octopuses have 8 arms and no tentacles whatsoever.

[3] Day also took another huge liberty in using the opening verses of the poem The Kraken (Alfred Lord Tennyson, 1830) without giving proper credit to the poet.

[4] Being stricter, the Watcher, like the Nazgûl’s flying mounts, remained nameless.

[5] Botanists (and mycologists) have their own code, the International Code of Nomenclature for Algae, Fungi, and Plants. Bacteriologists have their code as well, the International Code of Nomenclature of Bacteria.

[6] The abbreviation “n. gen.” after the name means “new genus” and indicates that the genus is being described here for the first time.

[7] Likewise, “n. sp.” means “new species” and indicates that the species is being described here for the first time.

[8] The nomenclatural acts on this article are presented simply for hypothetical concepts (a Middle-earth squid) and are disclaimed for nomenclatural purposes, being thus not available (ICZN Articles 1.3.1 and 8.3).

[9] In earlier writings, the names are usually spelled Nûrnen and Helkar.

Check other articles from this volume


Squids, octopuses and lots of ink

Rodrigo B. Salvador1 & Carlo M. Cunha2

1 Staatliches Museum für Naturkunde Stuttgart; Stuttgart, Germany. Eberhard Karls Universität Tübingen; Tübingen, Germany. Email: salvador.rodrigo.b (at) gmail (dot) com

2 Museu de Zoologia da Universidade de São Paulo; São Paulo, Brazil. Email: carlomagenta (at) gmail (dot) com

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Splatoon was recently released (second quarter of 2015) for the Wii U, receiving a warm welcome by Nintendo fans (it’s nigh unthinkable for the company to launch a new IP like this) and generating a flood of fan art on the Internet. The game is a third-person shooter with ink instead of bullets. It features two races, inklings (the playable one) and octarians (the enemies), and revolves around the fierce dispute against each other. (In multiplayer though, its inkling against inkling.) Inklings and octarians (especially the elite soldiers called “octolings”) are based, respectively, on squids and octopuses (Fig. 1), two of the most awesome kinds of animals out there.

Splatoon - Fig 1

Figure 1. Top left: an inkling girl in human form (original model from the game). Top right: Loligo vulgaris, an example of a squid species (photo by Hans Hillewaert, 2005; image modified from Wikimedia Commons). Bottom left: an elite octarian soldier, a.k.a. octoling (original artwork from the game). Bottom right: Octopus rubescens, an example of an octopus species (photo by Taollan82, 2007; image modified from Wikimedia Commons).

These animals are mollusks, and, more specifically, cephalopods. The mollusks are the second largest animal group on Earth (after the arthropods, of course) and includes gastropods (snails and slugs), bivalves (clams, mussels and oysters), cephalopods (we’ll come back to them soon) and the little known scaphopods (tusk shells), monoplacophorans, aplacophorans, polyplacophorans (chitons) and some fossil oddities. For those who remember their biological classification, we can put it like this: the class Cephalopoda belongs to the phylum Mollusca.

Cephalopoda is a group that contains a vast array of marine animals. Besides squids and octopuses, it counts with cuttlefish, nautiloids and the fossil belemnites and ammonoids. Today, cephalopods are found everywhere in the sea, from the polar regions to the tropics and from the surface to depths over 5,000 m. There are over 800 known living species of cephalopods, but the fossil record counts with more than 17,000 species (Boyle & Rodhouse, 2005; Rosenberg, 2014).

The class appeared over 450 million years ago during the late Cambrian, the first period of the Paleozoic era (Boyle & Rodhouse, 2005; Nishiguchi & Mapes, 2008). Cephalopods enjoyed a high amount of diversity during the Paleozoic and Mesozoic eras, with hundreds of species of nautiloids and ammonoids (Monks & Palmer, 2002). Most of these forms, however, did not survive to this day. Ammonoids and belemnites were completely extinguished and today we have just a handful of Nautilus species and the group consisting of cuttlefish, squids and octopuses. This group is a latecomer in cephalopod history: it appeared only during the Mesozoic, although some late Paleozoic fossils have tentatively been classified as squids (Boyle & Rodhouse, 2005; Nishiguchi & Mapes, 2008).

OK, so we can all see that Splatoon is a cephalopod-themed game. But before we can say something about the critters starring in Splatoon, we must first clear some Biology stuff. The foremost of these issues is: there is a long-lived and persistent confusion in popular knowledge, art and fiction, regarding squids and octopuses. People just do not seem to know which is which. Many biologists have bemoaned this and tried to set things right in popular works for quite a while (e.g., Lee, 1883; Salvador & Tomotani, 2014). Since these animals are our main theme here, we feel obliged to explain what, after all, are the differences between a squid and an octopus.


For the uninitiated (i.e., those whose sad childhoods did not include wildlife documentaries and visits to zoos and/or aquariums), squids and octopuses look the same. However, if one pays attention and carefully compare one to the other, many differences start to pop up. The first one is the overall shape of their bodies (Fig. 2): squids usually have a bullet-like shape and a more hydrodynamic body, with a pair of fins on their extremity; octopuses have a globose body, which is capable of some serious shape-changing. These animals’ shapes are linked to their way of life: squids are active swimmers, while octopuses live on the sea floor, hiding under rocks or inside burrows.

Splatoon - Fig 2

Figure 2. Diagrams of a squid (above) and an octopus (below), accompanied by the proper scientific terminology of their body parts. Image reproduced from Salvador & Tomotani (2014).

Nevertheless, there is an even more striking difference (Fig. 2): an octopus has only eight arms, while a squid has eight arms and two tentacles (readily identifiable: they are more slender and usually longer than the arms).

The difference in meaning between “arm” and “tentacle” is crucial, but these words unfortunately are used interchangeably in popular writing. The arms of both squids and octopuses are covered with suction cups (or suckers) on their inner surface. (These suckers are sessile in octopuses, but squids have stalked mobile ones.) The tentacles, present only in squids, are smooth (i.e., without suckers) along almost their entire length; only the tentacle’s tip (called “club”) has suckers (Fig. 2).

Despite the popular confusion of squid/octopus and arm/tentacle, it seems Splatoon’s developers took care to be as accurate as they could with their cartoonish squids. The inklings, when in squid form, have the correct number of arms, with the two tentacles clearly differentiated (Figs. 4, 9). Incredibly, this is also true for an inkling in human form and the game’s official Japanese Twitter took some pains to show it is so (Fig. 3).

Splatoon - Fig 3

Figure 3. Diagram showing the correspondence of arms between an inkling (in human form) and a real squid. The tentacles are numbered as 1 and 2, while the remaining arms are numbered 3 to 10. Image taken from Splatoon’s official Japanese Twitter (https://twitter.com/SplatoonJP).

Finally, the last main difference lies inside their bodies. As anyone can tell, the most obvious feature of mollusks is their shells; just think of a snail or a clam and you immediately picture their shells. However, most living cephalopods do not have shells (nautiluses and the tiny Spirula spirula are the exception).

Squids have only a remnant of a shell called a “pen” (or gladius) that serves as a skeletal support for their bodies. Octopuses, however, have absolutely no shell whatsoever. In all other respects, squids and octopuses are very similar, since they are both cephalopods. So now we will dabble a little in cephalopod anatomy, because we will need some concepts to discuss other features from Splatoon.


Before moving on to other topics, we have one final note on tentacles. The inkling boy’s tentacles, more specifically the club on the tip of each tentacle, are smaller than the inkling girl’s is. The official artwork seems to suggest that this is the case (Fig. 4) and this was confirmed in-game at least for the inklings in human form (it was difficult to compare squid forms due to all the shooting). Moreover, this is clearly seen on the Amiibo figures, where the boy’s tentacle clubs are about half the size of the girl’s (Fig. 5).

Splatoon - Fig 4

Figure 4. Inkling girl (top) and boy (bottom) in both human and squid forms. (Original artwork from the game.) Splatoon - Fig 5

Figure 5. Photos of the Amiibo figures of the inkling girl (left) and boy (right) in human form. Note how the girl’s tentacles are much larger than the boy’s.

Sexual dimorphism (i.e., male and female of the same species looking different) is rather common in cephalopods and the difference usually lies on overall body size: in some species the males are larger while in others the females are the larger ones (Boyle & Rodhouse, 2005). However, differences in the size of tentacles and clubs seem to be rare, known only from cuttlefish (Bello & Piscitelli, 2000). Perhaps this matter was not investigated enough in other species, meaning that differences in tentacle or club size could be more widespread in cephalopods. In any case, Bello & Piscitelli (2000) studied the species Sepia orbignyana, in which the females are larger than the males. They discovered that the female’s tentacle clubs were also proportionately larger (in relation to the body) than the male’s. They discuss that this feature is linked to feeding: cuttlefish use their tentacles to capture prey and, since females are larger and need more food, individuals with larger clubs have an advantage (they are able to capture larger prey) and were thus selected through the species’ evolutionary history.

Now, moving on with the anatomy. The mouth of a cephalopod is located in the middle of the circle formed by the arms. It contains a pair of powerful chitinous mandibles, which together are called a “beak” due to their resemblance to a parrot’s beak (Fig. 6).

Splatoon - Fig 6

Figure 6. A (dead) specimen of the squid species Loligo sanpaulensis. The top image show the whole body of the animal. The bottom left image shows the mouth region on the center of the ring of arms and tentacles. The bottom right inset shows a close-up of the mouth, with the beak barely visible on its center. The last two insets on the very bottom right show the beak (removed from the specimen) in frontal and lateral views.

Inside the mouth lies the radula, a rasping tongue-shaped structure equipped with many rows of small chitinous teeth (Fig. 7). The animals use the radula to “scratch” their food and pluck small portions of it. (Note that the radula is a feature common to all mollusks, with the notable exception of the bivalves, which are filter feeders and have lost the radula in the course of evolution.) In Splatoon, the designers’ care is shown here once more: inklings in human form have pointed teeth to emulate the sharp beaks of squids.

Splatoon - Fig 7

Figure 7. The radula (removed from the specimen of Fig. 6), shown straightened out. Notice the neat rows of pointy little “teeth”.

Cephalopods breathe through gills located inside the mantle cavity (Fig. 8). The water enters this cavity through apertures on the mantle edge close to the head and brings dissolved oxygen to the gills. The water is then expelled from the cavity by a structure called “funnel”. The funnel is also capable of expelling a powerful gush of water, which is responsible for the fast jet propulsion movement of cephalopods.

Splatoon - Fig 8

Figure 8. The arrows indicate the way in which water passes through the squid’s body, entering the mantle cavity (shown in transparency), passing by the gills and exiting via funnel (shown in two positions, for forward and backwards movement). Image reproduced from Richard E. Young, 2000, The Tree of Life Web Project (Creative Commons Attribution Non Commercial License 3.0).

If inklings can leap very high from a pond of ink when in squid form, jet propulsion is the reason why. This might better be called “ink-jet-propulsion”, though (Fig. 9).

As a matter of fact, the funnel has an important role in the most characteristic aspect of cephalopod biology: inking. And this is what we will turn to now.

Splatoon - Fig 9

Figure 9. Inkling, in squid form, jumping propelled by its ink-jet-propulsion. (Original model from the game.)


Inside the mantle cavity there is an organ known as “ink sac”, which, as the name implies, is a reservoir of ink. The animals can expel this ink through the funnel and they do this in two very ninja-like manners: as clouds or as pseudomorphs (Derby, 2007). Ink clouds are pretty straightforward, functioning as a smoke screen to allow the cephalopod to escape (Fig. 10); although some recent observations show that they can also be used to confuse and sneak attack prey (Sato et al., 2016). Pseudomorphs (meaning “false-shapes”) are more curious things. They are made of ink and mucus and appear as a well-defined and stable form (maintaining its physical integrity for a while after it is released). These pseudomorphs can be almost as large as the animal releasing them. It is thought they serve as a decoy, a fake double of the cephalopod which will distract the predator and allow it to escape unharmed. Finally, cephalopods’ ink might also contain organic compounds that act either as toxins to deter or damage enemies or as signals to warn conspecifics (members of the same species) of any danger (Derby, 2007).

Splatoon - Fig 10

Figure 10. Ink cloud created by a Humboldt squid, Dosidicus gigas. Image reproduced from Bush & Robinson (2007).

In Splatoon, the main purpose of the ink is, well, to ink stuff. Inklings shoot ink through guns (not funnels), hoping to beat their opponents senseless (“splat” them, as the game puts it) and to ink the largest portion of the battlefield to achieve victory (Fig. 11).

Splatoon - Fig 11

Figure 11. A chaotically inked Splatoon battlefield. (Screenshot from the game.)

Inkling’s ink is the same color as their body (Fig. 12). True cephalopods’ ink is always dark (due to its main constituent, melanin), of course, but we can all agree that different colors of ink was a fair gameplay necessity. And, speaking of body color, cephalopods are the most colorful animals out there (sorry, birds).

Splatoon - Fig 12

Figure 12. An inkling’s ink is always the same color as its body. (Original models from the game.)


Cephalopods are so colorful because they can actually change their body color and also their color patterns. They have specialized cells in their skin called cromatophores, which enable them to instantly change color to camouflage themselves (either to evade predators or to ambush prey; Fig. 13), to communicate with conspecifics, or to ward off predators (Hanlon & Messenger, 1996; Hanlon, 2007; Mäthger et al., 2012). Some scientists even argue that cephalopods can produce waves of changing color patterns to mesmerize prey and make them easier to catch (e.g., Mauris, 1989; Mather & Mather, 2006), but this remains scarcely proved.

Doing justice to cephalopod coloration, inklings come in many colors: turquoise, lime green, purple, pink, orange and blue. And they change colors basically for each battle.

Splatoon - Fig 13

Figure 13. Photo sequence showing an octopus de-camouflaging itself. Image reproduced from Hanlon (2007).

We mentioned above that cephalopods can communicate with each other by changing their color pattern (Shashar et al., 1996; Mäthger et al., 2009). This kind of communication can only work if the animals using them possess a high degree of intelligence. And indeed they do. Cephalopods can solve puzzles, cause all sort of embarrassments for caretakers in aquariums and zoos, and even use tools (Mather, 2008; Finn et al., 2009). Their nervous system is the most complex among invertebrates and, actually, their brain-to-body-mass ratio falls between those of endothermic (birds and mammals) and ectothermic (all the others) vertebrates (Nixon & Young, 2003).

In Splatoon, the inklings and octarians are clearly intelligent enough to build cities and weapons. One of the Sunken Scrolls (a kind of game collectible that tells more about the backstory) reveals that the two races evolved when rising sea levels (yes, that’s Global Warming) wiped out humans and allowed sea creatures to take over (Splatoon Wiki, 2016).

Box 1. Ink and slime

Inklings can only move freely on ink of the same color as theirs; they get stuck in other colors of ink (i.e., those produced by their opponents). Cephalopods, of course, have no such restriction of movement, but another kind of mollusk does. Land snails and slugs produce a mucous slime in order to move; they actually “glide” on top of it. Was this an intentional decision by the game developers, based on actual knowledge of mollusks? Or was this merely a gameplay choice that resulted in a nice molluscan coincidence?

Splatoon - Box 1

A land snail leaving a silvery slime trail on its wake. Photo by “snail ho”, 2007; image modified from Wikimedia Commons.


Now, after such a long immersion in cephalopod biology, it’s past time we try to identify which species exactly (if any) served as the basis for Splatoon’s inklings. (Unfortunately, octarians have a way too generic octopus design to allow some proper identification.)

The game’s developers have not stated which squid species (or number of species) they used as basis for the inklings. However, gamers on the Internet have referred to the Humboldt squid (Splatoon Wiki, 2016) or to the Japanese flying squid (on some forums). The Humboldt squid (Fig. 14), Dosidicus gigas, in the first place, is not a Japanese species (we assume Japanese game developers basically only use Japanese stuff for their games; see the majority of critters in Pokémon, for instance). The Humboldt squid is found on the Pacific coast of the Americas (Zeidberg & Robinson, 2007). As such, the Japanese flying squid (Fig. 14), Todarodes pacificus, would seem a good choice.

Splatoon - Fig 14

Figure 14. Top left: Dosidicus gigas, the Humboldt squid (image modified from Wikimedia Commons). Top right: Todarodes pacificus, the Japanese flying squid (image modified from Wikimedia Commons). Bottom left: Thysanoteuthis rhombus, the diamond squid (image modified from Wikimedia Commons). Bottom right: inkling in squid form (original model from the game).

The Japanese flying squid belongs to the same family (Ommastrephidae) as the Humboldt squid. The squids in this family, as their common names imply, use jet propulsion to “fly” above the sea surface, covering a few tens of meters in each jump. This behavior is thought to be related to predator avoidance or to save energy as the squids migrate across vast ranges. It could be safely assumed that the jet-propelled jumps of the inklings in Splatoon (Fig. 9) were based on squids from this family. As a matter of fact, other flying squids of the family Ommastrephidae can be found in Japanese waters, such as the “neon flying squid”, Ommastrephes bartramii.

Ommastrephidae species also have a shape similar to the inkling’s squid form (Fig. 14), with large fins forming a triangle on the tip of the body, a tubular section of the mantle leading from it to the head and small arms. As such, it is safe to assume that this family of squids served as inspiration for both behavior and design of the inklings. However, the tubular section in the inkling is where the eyes are located and could thus be interpreted as the whole head. As such, there is another possible species that might have influenced the inkling’s design: the diamond squid (Fig. 14), Thysanoteuthis rhombus (family Thysanoteuthidae). As its name implies, the fins give the animal a diamond shape and extend all the way to its head (as the inkling’s head seem to be immediately below the fins). Moreover, it has short and strong arms and tentacles, like the inklings. This species can be found worldwide, but is an important catch for the Japanese fishing industry (Bower & Miyahara, 2005) and it is to this topic that we will turn next.


It is very common for gaming developers to come up with jokes for April Fools’ and last year even Nintendo joined in. A post on Splatoon’s official Tumblr page read: “So… this whole time I thought ‘Splatoon’ was going to be the name of the game, but it’s not. Splatoon is actually a hot new snack that’s coming out in May! You gotta be squiddin’ me! Now you can have your squid and eat it too! Unless you’re a squid, then… maybe don’t because that’s weird and kinda creepy. Maybe just eat a quesadilla.” A photo of the supposed snack’s package (Fig. 15) accompanied the post.

Splatoon - Fig 15

Figure 15. The April Fools’ Splatoon snack. Image taken from Splatoon’s official Tumblr (http://splatoonus.tumblr.com/).

While it may be a cute April Fools’ joke (the Japanese seem to love these strange snacks), it brings up a very serious question: Japan’s destructive fishing practices. As remarked above, the diamond squid is a very important species for the Japanese fishing industries, but the Ommastrephidae flying squids also consist in large portions of their catch (FAO, 2015a). As a matter of fact, all of these species are heavily exploited and the species are in steady decline (Bower & Ichii, 2005; Bower & Miyahara, 2005; FAO, 2015a).

The loss of a single squid species may not seem much to most people. But when a species go extinct, its absence may lead the whole ecosystem to a disastrous collapse. In the long run, the only thing we’ll be eating from a barren sea will be jellyfish burgers. Japan’s fishing industry is one of the most overexploiting in the world (Clover, 2004: Roberts, 2007; FAO, 2015b); besides, Japan is involved in a huge controversy regarding its whaling practice.

Nobody is expecting people to just stop fishing; the sea is an amazing resource to feed the increasing number of humans on the planet. Moreover, seafood is a staple of Japanese cuisine. Even so, the fisheries could do better with some planning, to avoid species and ecosystems collapses. Some restrictions, accompanied by proper regulation, should be put in place. For us consumers, Clover (2004) offers advice on how to make a difference, buying only lawfully catch seafood (identified by seals of quality or similar markings) and always inquiring where said seafood comes from (to avoid buying something from threatened areas).

Box 2. Mario’s Famous Squid

The inklings’ squid form is also reminiscent of another famous Nintendo squid: Blooper. Everyone should be familiar with this recurrent antagonist from the Mario series, which is present in almost every underwater stage across several Mario titles (Cavallari, 2015). Despite Blooper’s simple design, it can be safely recognized as a squid (Cavallari, 2015) and it is curiously very similar to Splatoon’s inkling. However, according to the game’s developers, they did not use Blooper as a base for the inkling’s squid form design (Splatoon Wiki, 2016).

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Inkling in squid form (left) and Blooper (right). Artwork by CutyAries (http://cutyaries.deviantart.com/); image is a courtesy of the artist.


Typically, invertebrates are not a priority in conservation efforts, since they are usually faced with lots of antipathy. As such, the role of flagship species for raising people’s awareness about biodiversity loss and the dire need of conservation measures is often reserved for mammals and birds (and the eventual tiny cute frog). Cephalopods usually do not get much attention in conservation efforts, but they do have some charismatic species that can serve such purpose (and should definitely be used). Among the squids, the semi-mythic giant squid Architeuthis sp., the animal who originated the legend of the Kraken, is the most obvious choice (Guerra et al., 2011; Salvador & Tomotani, 2014). Among the octopuses, everyone should remember Paul, the “clairvoyant” cephalopod who predicted the results of the 2010 FIFA World Cup, and the little octopus Opisthoteuthis sp. that made the news everywhere this year just for being so damn adorable (Fig. 16).

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Figure 16. Opisthoteuthis sp. Screenshot taken from the YouTube video by SciFry (15/Jun/2015): https://www.youtube.com/watch?v=wv7DfebpL7E.

There is no doubt that the media and works of fiction can help in environmental education and in raising ecological awareness. (It can work against it too, like when the movie Jaws started a shark-killing frenzy.) We believe that games can and should have a much more prominent role in these efforts. This is especially true for a game like Splatoon, which is more children-friendly. After all, we have to teach this ecological awareness to children (adults are already too narrow-minded to listen).

For instance, the game Never Alone (released November 2014) has its story and setting based on Inuit folklore and culture. As the player progresses in the game and encounters different things, he/she unlocks a series of short documentaries which explain facets of the Inuit culture, their arctic environment and arctic animals. This shows that games can both entertain and actually teach something of value.

A different (and more direct) kind of approach was taken by Rovio Entertainment Ltd., house of the famous Angry Birds franchise. The company joined forces with BirdLife International to fight against the extinction of the wondrous South Pacific birds. Unfortunately, their crowdfunding campaign reached only about half of its US$ 150,000 target in donations (Save the Birds of the Pacific, 2015). Just for comparison, games overhyped by the media (e.g., Shenmue) were meanwhile gathering over 6 million dollars on Kickstarter; so gamers do have money to spend.

Could Splatoon be used for educating players and raising environmental awareness? Yes, it could. Could Nintendo do the same as Rovio did and join some cephalopod conservation effort? Definitely. Will this ever happen? Likely not; as explained above, Japan is too busy eating all squids in the Pacific Ocean.


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The material analyzed for this study (i.e., the inkling Amiibo figures) is deposited in the private collection of one of the authors (R.B.S.), next to a Kirby and an Elite Stealth Elf.

The squid specimen from Figs. 6 and 7 is deposited in the scientific collection of the Museu de Zoologia da Universidade de São Paulo (São Paulo, Brazil) under the record number MZSP 86430. No mollusks were harmed during this work.

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Shells and bytes: mollusks in the 16-bit era

Daniel C. Cavallari

Museu de Zoologia, Universidade de São Paulo; São Paulo, Brazil.

Email: dccavallari (at) gmail (dot) com

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Mollusks are one of the most diverse groups of organisms known to science. Humanity has described more than 80,000 species of snails and slugs (gastropods), clams, oysters and scallops (bivalves), squids and octopuses (cephalopods), tusk shells (scaphopods) and their less-known kin (e.g., polyplacophorans and aplacophorans), and many more are waiting to be discovered (Chapman, 2009). These soft-bodied and most often shell-bearing animals present a wide variety of shapes, colors and behaviors that has fascinated mankind for ages. Used as currency, tools, jewelry, medicine, and extensively collected and depicted in many different ways, mollusks are strongly tied to human history. It is not surprising that they were, and still are an important part of different cultures everywhere (Simone, 2003; Sturm et al., 2006).

Naturally, in our everyday lives, some of us portray mollusks and their shells in paintings and decoration. They are present in ordinary objects, such as stamps and coins (Robertson, 2011; Todd, 2011). In modern culture, more specifically in the history of video games, the presence of mollusks is a remarkable one. In the late 80s, the so-called fourth generation of consoles (the 16-bit era) began, and with it, many vividly colored games came to be (Kent, 2001). Higher resolution allowed for digital artists to depict characters and draw backgrounds with richer detail. Mollusks showing up in 16-bit era games were naturally part of underwater scenery art, but some were given fairly important roles as memorable characters, plot devices, fierce antagonists and fearsome bosses. Herein we number some of these appearances and take the opportunity to discuss the real-world inspirations that probably originated these 16-bit representations. For practical reasons, only games that had an international or at least an English-language release are included in the following list.


Several platforms existed in the 16-bit era, but the clash between its two greatest consoles was what we could call its “main event” (Fig. 1). The Sega Genesis (also known as the Mega Drive) was launched in Japan in 1988. Its greatest worldwide competitor, the Super Nintendo Entertainment System (short SNES, known as Super Famicom in Japan), was released two years later, in 1990 (Kent, 2001). Games developed for the SNES and Mega Drive platforms marked a generation. Some were the starting point for very successful franchises, and many of them depicted mollusks.

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Figure 1. Clash of titans. Super Nintendo Entertainment System (top) versus Sega Genesis (bottom). Photos by Evan Amos, extracted and modified from Wikimedia Commons.


One of the most famous RPG franchises of gaming history, Final Fantasy includes several 16-bit titles such as Final Fantasy IV (1991), Final Fantasy V (1992) and Final Fantasy VI (1994), among others. These games have since been adapted for more recent platforms such as PlayStation, GBA and mobile. With a multitude of elaborately designed enemies (based on the beautiful artwork by Yoshitaka Amano), Final Fantasy games for the SNES include many mollusk-themed foes. Some of them received special attention from the developers, participating in some of the most memorable moments in the entire franchise (Fig. 2). The enemy names listed below are exactly as they appeared in the SNES, and may not coincide with more recent versions.

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Figure 2. The very first boss in Final Fantasy VI, Whelk (left) ambushes two Imperial soldiers and female protagonist Terra (right). And yes, do not attack the shell: it may have great scientific value! (Screenshot from the actual game.)

Final Fantasy IV (FFIV) mollusk enemies are not exactly remarkable in terms of the game’s plot, but have curious design nonetheless. FangShel and its recolor EvilShel (Figs. 3A and 3B, respectively) are apparently based on bivalves in the family Pectinidae, commonly known as scallops (Fig. 5C). Mindflayer (Fig. 3C) is a Lovecraftian monster of sorts with a head somewhat resembling a squid (a teuthid cephalopod); mindflayers are also recurrent enemies in the Dungeons & Dragons role playing game franchise, and have a similar design. Octomam (Fig. 3D) is a huge colorful octopus-like creature with a crazy smile. It is one of the game’s bosses. Some species of blue-ringed octopus, such as Hapalochlaena lunulata (Fig. 5E), have a similar pointy mantle (the actual head of octopuses roughly corresponds to the region near the eyes; Jereb et al., 2005), and a comparable color pattern.

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Figure 3. Mollusk foes from Final Fantasy IV and V (sprites from the mobile versions of both games). A. FangShel. B. EvilShel. C. Mindflayer. D. Octomam. E. Aquathone. F. Ammona. G. Sucker. H. Luneta. I. Mindflayer (FFV). J. Octoraken. K. MooglEater. L. RockGarter. M. Slug.

Fortunately, Final Fantasy V’s (FFV) roster of enemies had more mollusk representatives. Aquathone (Fig. 3E) and its recolor Ammona (Fig. 3F) are apparently based on reconstructions of ammonites (ancient extinct cephalopods; Fig. 5A), judging by its shell sculpture, cephalic hood (the triangular plate over its head, right above the eyes) and the numerous thin tentacles.  Sucker (Fig. 3G) and its recolor Luneta (Fig. 3H) are apparently based on squids. Both have very clear tentacles with enlarged tips, and curiously vitreous bodies, through which their inner organs can be seen. A translucent body is a characteristic of some squid groups, like glass squids (family Cranchiidae, Fig. 5D). There is yet another Mindflayer (Fig. 3I), though its head resembles squids with smaller fins, such as some species in the family Cirroteuthidae. The last cephalopod representatives are Octoraken (Fig. 3J) and its recolor MooglEater (Fig. 3K), which are surprisingly realistic octopuses. Finally, RockGarter (Fig. 3L) and Slug (Fig. 3M) are different colored versions of a gastropod-based design. They are apparently simple representations of terrestrial gastropods pertaining to the limacoid clade, which is a grouping of air-breathing land snails and slugs that share a common ancestor. Coincidentally, there is a species of keelback slug (family Limacidae), Bielzia coerulans, in which juvenile specimens have a brown colored body while adults are completely blue.

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Figure 4. Mollusk enemies in Final Fantasy VI (sprites from the mobile version of the game). A. Nautiloid. B. Cephaler. C. Ultros. D. Presenter.

Though the mollusks in Final Fantasy VI (FFVI) were not as numerous as in the preceding titles, some were given far more important roles. In fact, the very first boss in the game is none other than Whelk (Fig. 2), a scary giant gastropod. It surely looks like a terrestrial snail (despite the odd position of the eyes) but not much else can be said on the real world animal that inspired its design. The Presenter (Fig. 4D) is a recolored version of Whelk, with a different head-foot (it’s actually holding a treasure chest), and not much more can be said about it as well. Other minor enemies worth noting include Nautiloid (Fig. 4A) and its recolor Cephaler (Fig. 4B), which as the names may indicate are probably based on cephalopods. The shell color pattern of both resemble actual species of nautiloid cephalopods, such as Nautilus pompilius (Fig. 5B), though the arms with suckers do not.

Finally, the most (in)famous of all these mollusk antagonists, and perhaps the most well-known mollusk in the Final Fantasy franchise, Ultros (Fig. 4C) is a purple octopus-themed creature with an odd “smile” and rolling eyes. It is undoubtedly a comic relief character, with a total of four (hilarious) appearances in the game. The most important encounter involves the cherished Opera House scene. Ultros denominates itself an octopus, though it lacks the characteristic arm suckers of most real-life cephalopods. It can spit ink, however, as it has an ink-related attack that can render player characters blind. Since FFVI, Ultros is a recurrent character/enemy in the franchise.

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Figure 5. Plausible real-life inspirations for 16-bit Final Fantasy molluks. A. A digital reconstruction of the ammonoid Asteroceras obtusum by Nobu Tamura. B. A live specimen of Nautilus pompilius. C. A live individual of the scallop Argopecten irradians. D. Photo of a glass squid (Cranchiidae) juvenile by Uwe Kils. E. A photo of the blue-ringed octopus Hapalochlaena lunulata, by Jens Petersen. (All images were extracted and modified from Wikimedia Commons; images are public domain unless otherwise stated.)


Super Mario is a world famous game series that actually predates the SNES, coming from the 8-bit generation of consoles. Nevertheless, the world’s most famous plumber evolved amazingly well into the 16-bit era, with several successful titles such as the remakes of Super Mario Bros. 2 and 3, Super Mario World, and Super Mario RPG: Legend of the Seven Stars, in the early to mid-1990s.

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Figure 6. Blooper the squid welcomes Mario to the underwater world in the SNES version of Super Mario Bros. (Screenshot from the actual game.)

A humble yet persistent mollusk antagonist in the series, Blooper is present in almost every underwater stage in several Mario titles. It is pretty much a squid-like creature (Fig. 6), and its main offensive strategy (as is the case in most of the earliest Mario’s enemies) consists of causing damage upon contact. A very basic maneuver, of course, but a pretty dangerous one in confined spaces crowded with other enemies. Blooper is loosely based on real world squids. It possesses enough features to be recognized as a cephalopod (the class of mollusks that includes octopuses, squids and their allies) and more specifically a Teuthida (the order that includes squids and their allies). It has arms and tentacles with suckers, and fins on both sides of the body (not exactly a head), conferring it a pointy silhouette, but not much else can be inferred from its relatively simple and stylized design.


Breath of Fire (BoF) is a roleplaying game series that had its first installment launched in 1993 for the SNES. The sequel, Breath of Fire II, came a year later. Both games are focused on the adventures of Ryu, a dragon-descendant warrior on a continuous quest to uncover the history of his clan. As is the case in most RPGs, both BoF games had quite varied enemy lists. A few of these opponents were apparently based on mollusks.

The earliest mollusk-themed appearance in the first game is Tentacle (Fig. 7A), which would be followed much later by its recolored version Nautulis (Fig. 7B). They are probably based on reconstructions of ammonites (Fig. 5A), very similarly to some Final Fantasy monsters (see above). Squid and Octo (Figs. 7C and 7D, respectively) are cephalopod twins, and two game bosses no less. Judging by the shape of the mantle and fins, both were based on squids, but their real-world affinities beyond that are uncertain.

Breath of Fire II had a few more mollusk adversaries, starting with Amonica (Fig. 7E) and Cuttlecb (Fig. 7F). They look like an odd crossover between a cephalopod (tentacles and arms, very large eyes and an evident, uncovered siphon) and a marine gastropod belonging in the superfamily Trochoidea, judging by the rounded pyramidal outline of their shells. Roadslug (Fig. 7G) and R-Slug (Fig. 7H) are heavily stylized representations of terrestrial gastropods in the limacoid clade (see the Final Fantasy section above). Finally, the only mollusk boss in the game is Babaruku (Fig. 7I), the evil high priest of St. Eva in its Lovecraftian might. It looks like a mindflayer of sorts, a nasty human-octopus hybrid, but not much else can be said on its possible real-life counterpart.

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Figure 7. Molluscan enemies from Breath of Fire and BoF II (sprites from the actual games). A. Tentacle. B. Nautulis. C. Squid. D. Octo. E. Amonica. F. Cuttlecb. G. Roadslug. H. R-Slug. I. Babaruku.


First released in Japan in 1994, the game had a sinister plot involving rival demon lords fighting over ancient magical gems of power. This, as well as the fantastic organ music, creepy characters and enemies and beautifully-drawn gloomy scenarios gave the game one of the darkest tones of its generation.

Amidst this demonic war for absolute supremacy, an underwater stage boss – named Holothurion (Fig. 8) – actually posed a serious threat as a horrendous giant sea snail. His arsenal of menacing moves included creating overwhelming water currents that tossed the player character around (even toward deadly shell spikes or its huge uninviting mouth), and spitting chunks of an apparently toxic ink-like substance. It could also retract its body into the shell as a defensive maneuver – a very “snaily” move indeed.

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Figure 8. Top: Demon’s Crest protagonist Firebrand (in his green water-dwelling form, to the left) battles Holothurion, the giant demonic snail (right). (Screenshot from the actual game.) Bottom: Original artwork of Holothurion from the game.

Despite the hyperbolic boss-like size, Holothurion looks to be an amalgam of marine and perhaps terrestrial snail species. It has a bulky, orange-colored shell, with spines on the body whorl’s (the largest shell whorl) shoulder and on the upper portion of each whorl. This outline and color is reminiscent of some iconic strombid species like the queen conch, Lobatus gigas or the West Indian fighting conch, Strombus pugilis (Fig. 9D). The shell’s aperture (opening) has an unusual outline, and a clear, long siphonal canal is apparent. The series of holes observed on the shell’s dorsum are found in a few gastropods (Fig. 9C), such as abalones (family Haliotidae; Poutiers, 1998).

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Figure 9. Possible real-world inspirations for Holothurion. A. The sea slug Glaucus atlanticus (photo by Sylke Rohrlach). B. Three specimens of the sea slug Chromodoris willani (photo by Juuyoh Tanaka). C. Dorsal view of an empty shell of the abalone Haliotis varia. D. Dorsal view of an empty shell of Strombus pugilis (both shell photos by H. Zell). (All photos extracted and modified from Wikimedia Commons.)

The demonic snail’s body, or rather its soft parts, is a matter of further discussion. The blue coloration is not itself rare, but it is more common in non-shelled species such as nudibranchs (sea slugs). Besides being possibly similarly blue, the sea slugs Glaucus atlanticus (Fig. 9A) and Chromodoris willani (Fig. 9B) both may have darker blue lateral stripes with an adjacent, lighter stripe just below it. The posterior portion of Holothurion’s foot has a few “spiky” appendages. They could be part of a very bizarre operculum (a structure that closes the shell aperture when the animal retracts into the shell), if they weren’t actually mobile structures that react to the snail’s mood. This may be another similarity with some nudibranchs, which have comparable posterior appendages (gills; as in C. willani), some of which can be contractile or retract into a cavity (Fahey & Gosliner, 2004). The demonic snail’s head looks like that of most shelled marine gastropods, with a single pair of cephalic tentacles. They are probably eye-bearing, though there are no discernible eyes on the in-game sprite. This somehow contradicts the official artwork (Fig. 8), which depicts two pairs of cephalic tentacles most similar to a land snail, and the upper pair appears to bear eyes at the tips. In any case, after such an in-depth analysis, I can only conclude that such a freakish molluscan gestalt could only be brought into existence by demonic forces.


The Donkey Kong Country series featured 3 games in the SNES platform. It is a very successful franchise that began as a very dynamic platformer. The game protagonists and antagonists are animal-based: simian protagonists (the Kongs) and their animal buddies face off against an evil crocodile gang and their allies in richly-designed, nature-themed scenarios.

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Figure 10. A common underwater enemy, Clambo (at the bottom) throws pearl bullets at the Kongs. (Screenshot from the actual game.)

The first game of the series was released in 1994. It is the only one among its 16-bit counterparts to include mollusks. Clambo (Fig. 10) is a pearl-spitting bivalve, a stationary opponent waiting to hit any simian swimmers with vicious underwater pearl bullets. Well, in fact and contrary to what the name indicates, Clambo might be an oyster (family Ostreidae), and not an actual clam. Judging by the thick shell with strong undulated sculpture, it could be a zigzag oyster (genus Lopha). Alternatively, it could indeed be a clam, a giant clam in the genus Tridacna (Fig. 12A). Tridacnids have a somewhat similar shell outline and are known to produce pearls, though not of the nacreous (iridescent) type (WJC, 2013). Giving the name some credit, the second option seems more plausible.

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Figure 11. Touching the cephalopod Croctopus (bottom left) could prove deadly. (Screenshot from the actual game.)

Croctopus (Fig. 11), a swirling and rather deadly cephalopod, is another recurring mollusk enemy and perhaps the most fearsome (and sometimes annoying) one in the game. Its offensive abilities are not the fanciest of the franchise, merely consisting of inflicting damage to the player characters upon contact. It is, however, invincible and often very fast-moving, and will invariably damage the player characters upon touch despite the circumstances (even while riding your favorite swordfish buddy, for instance). Surely, it was a foe to avoid. Croctopus’ design is loosely based on Indo-Pacific blue-ringed octopus species belonging to the genus Hapalochlaena (Fig. 12B). Blue-ringed octopuses are venomous animals, and their venom is nothing short of a cocktail of deadly substances. Accidents involving humans are often fatal, so any resemblance between Croctopus’ offensive power and real life is -not- a coincidence (Sheumack et al., 1978; McMichael, 2013).

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Figure 12. Possible real-life inspirations for Clambo and Croctopus, respectively. A. A live specimen of Tridacna (public domain image). B. A blue-ringed octopus, Hapalochlaena maculosa (photo by Sylke Rohrlach, extracted and modified from Wikimedia Commons).

Another enemy worth mentioning is Squidge, a fast-moving, translucent creature. Despite the name, it does not have much to do with squids, being more of a jellyfish (a cnidarian) with an odd pair of angry eyes. It moves around by agitating its gelatinous body in an umbrella-like fashion, and its tentacle-like posterior appendages are more akin to cnidarian oral arms, lacking the typical suckers of cephalopod arms and tentacles.


The first Mega Man X game was released in Japan in 1993 and in North America a year later. Set in a dystopian future, the game’s plot comes down to an Aasimovian conflict between evil robots (mavericks) aimed at destroying their human creators and the ones that wish to defend them. Bosses in the X series were mostly animal-themed. Their design was based on lizards, mammals, fish, insects, birds and, of course, mollusks.

Launch Octopus (Figs. 13, 15A) was the underwater stage boss in the first game. Featuring six torpedo-firing arms (which adds to eight members if you count the legs), and the power to produce powerful whirlpools, this octopus-themed enemy could give the player a good headache. It had more weaknesses than a normal Mega Man boss would however; it was vulnerable to two of X’s weapons, one of which (boomerang cutter) could sever his awesome arms, greatly crippling his combat ability. In any case, assigning this robotic foe to a single cephalopod species or genus is no easy task. There is a small hint to a possible biological correlation, though: his arms have a single row of suckers, or rather, devices that look like suckers. Uniserial suckers are characteristic of some octopod families, like Eledonidae, Amphitretidae and Megaleledonidae (Strugnell et al., 2014). Amphitretids have vitreous bodies, and Megaleledonids inhabit arctic environments. Both facts cannot be applied to Launch Octopus, leaving Eledonidae as the most plausible option.

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Figure 13. Mega Man X (left) versus Launch Octopus (right). (Screenshot from the actual game.)

Snails may not be the quickest animals out there. One might assume they would not make for astounding enemies in a quick-paced shooter/platformer like Mega Man X2. Well, Crystal Snail (Figs. 14, 15C), a snail-themed maverick, proves this theory wrong. Possessing an invulnerable shell with rocket-like propellants, it can fly around the screen, and even change its trajectory in midair. It can also spit a gooey mucous substance that crystalizes upon contact, imprisoning Mega Man in a shiny crystal coffin. To top it off, Crystal Snail has an amazing time-slowing special ability, which turns the world into a sluggish hell while it moves freely. On a first glance, Crystal Snail’s design is based on a terrestrial gastropod, bearing two pairs of cephalic (head) tentacles and a characteristic shell. The shell itself is bulky, and seems to be somewhat planispiral (coiled in a single horizontal plane). There are several species with planispiral shells, but most are not that bulky or rounded. It seems more likely that the maverick’s design was based on a more common species such as the garden snail (Cornu aspersum), and the shell’s odd coiling axis is merely an artistic interpretation.

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Figure 14. Mega Man X (right) versus Crystal Snail (left). (Screenshot from the actual game.)

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Figure 15. Mega Man X foes and their plausible natural inspirations. A. Launch Octopus (official game art by Capcom). B. The Eledonid cephalopod, Eledone cirrhosa (public domain image from a 1896 monograph by Jatta Giuseppe). C. Crystal Snail (official game art by Capcom). D. The common garden snail, Cornu aspersum (image by Rasbak, extracted and modified from Wikimedia Commons).


Released in Japan and North America in 1995, Chrono Trigger became a commercial success, and received extensive critical acclaim. It was an interesting RPG, with an innovative combat mechanics and an enticing plot revolving around time travel and apocalyptic events.

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Figure 16. The Rainbow shell (center) harbored in Guardia’s treasury. (Screenshot from the actual game.)

An interesting, optional aspect of the game’s plot, the rainbow shell (Figs. 16, 17A) was a huge specimen that remained hidden in a series of unnatural pre-historic caves in 600 A.D, guarded by a plethora of dinosaur enemies and a fearsome Tyrannosaurus-like boss. There it lied, waiting to be discovered by Crono, the game’s silent protagonist, and his friends. Once claimed by the heroes and after a series of plot twists involving the kingdom of Guardia in a future timeline, the rainbow shell became part of the royal treasure. It was, in fact, its most important piece, and could also be used as raw material to fabricate some of the game’s most powerful gear and weapons (such as the Rainbow blade, Crono’s top weapon).

At first glance – and considering the “ancient pre-historic cave” context – the rainbow shell looks like a fossilized ammonoid, an ancient shelled cephalopod. Indeed, some ammonoids are among the largest shelled mollusks that ever lived (Monks & Palmer, 2002). Given its coiling axis and the strong axial sculpture (with thick, sequential ribs), as well as its huge size (it took no less than 8 soldiers to move it out of the cave!), the rainbow shell could have pertained to the family Desmoceratidae, like species in the genus Parapuzosia, which could reach over 3 m in diameter (Teichert & Kummel, 1960). Ammonoid fossils with an iridescent shade (a rainbow-like shine) are not unheard of (Fig. 17B), and could have inspired Akira Toriyama’s artwork and in-game sprite.

Mollusks - Figure17

Figure 17. Iridescent lookalikes. A. The Rainbow shell (official Chrono Trigger art by Akira Toriyama). B. An iridescent ancient ammonite fossil on display at the American Museum of Natural History, New York (image by James P. Fisher III, extracted and modified from Wikimedia Commons).


One of the most famous series in the 16-bit era and greatly cherished by Sega Genesis owners, Sonic the Hedgehog had many successful titles in the early to mid-90s. Sonic games were very agile platformers, with colorful and complex ambiences full of twists and turns, challenging enemies and bosses. Enemies in the series were mostly robots created by the infamous Dr. Ivo “Eggman” Robotnik. The (not so) good doctor based some of his creations on mollusks, of course.

In Sonic the Hedgehog 2, Octus (Fig. 18A) is an octopus-like robot that jumps using its robotic arms, and tries to hit Sonic with energy projectiles. It is obviously based on a cephalopod, but not much else can be inferred. In sonic the Hedgehog 3, Clamer (Fig. 18B) and Snail Blaster (Fig. 18C) were, respectively, bivalve and snail (a terrestrial gastropod) based artillery robots. While they’re not attacking the protagonists, both can hide inside their shells, which offer considerable protection.


Earthworm Jim 2 (1995): A smaller and recurrent enemy in the Level Ate stage, it appears to be based on terrestrial gastropods.

Joe & Mac 2 (1994): Had some minor unnamed mollusk-themed antagonists.

Kirby Super Star (1996):  A minor, white squid-like enemy called Squishy (Figs. 18D, 19) is present throughout the game. It is similar to Super Mario’s Blooper in some ways.

The Legend of Zelda: A Link to the Past (1991): A very common mollusk antagonist called Octorok (Fig. 18E) attacked the protagonist by spiting rocky projectiles. It is a stylized octopus, apparently, and has a Dark World counterpart called Slarok.

Lufia II (1995): A few of the game’s antagonists were mollusks, namely Evil Fish (an octopus, Fig. 18H), Drill Shell (a marine snail with a spiky shell, Fig. 18G) and Ammonite (yet another ammonoid-based monster, Fig. 18I).

Super Ghouls n’ Ghosts (1991): A devilish red-shelled clam named Eyeball Clam (Fig. 18F) attacked the protagonist by spitting eyeballs towards him. It is surely based on real-life clams (probably tridacnid, see Donkey Kong above).

Mollusks - Figure18

Figure 18. Sonic the Hedgehog’s mollusk enemies, and some additional honorable mentions. A. Octus (official art from Sonic the Hedgehog 2). B. Clamer (official art from Sonic the Hedgehog 3). C. Snail Blaster (official art from Sonic the Hedgehog 3). D. Squishy (official art from Kirby’s Dream Land). E. Octorok (official art from The Legend of Zelda: A Link to the Past). F. Eyeball Clam (official art from Super Ghouls n’ Goblins). G. Drill shell (sprite from Lufia II). H. Evil Fish (sprite from Lufia II). I. Ammonite (sprite from Lufia II).

Mollusks - Figure19

Figure 19. Kirby (left) and Squishy (right) have a nice conversation by the sea in Kirby Super Star. (Screenshot of the actual game.)


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Praise Helix!

Rodrigo B. Salvador

Staatliches Museum für Naturkunde Stuttgart; Stuttgart, Germany.

Eberhard Karls Universität Tübingen; Tübingen, Germany.

Email: salvador.rodrigo.b (at) gmail (dot) com

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It is not everyday that I manage to join two of my main interests, mollusks and mythology. So rejoice!, for today is one of those days. I bid you welcome to the Cult of the Helix.

So how was this cult born? Nature-worshipping barbarians coping in a dangerous environment? An old bearded guy receiving revelations in the desert? A bald hermit meditating in the mountains? Well, none of the above. The Cult of the Helix was born in a most unorthodox manner: on the first iteration of Twitch Plays Pokémon. Wait, what?


Twitch Plays Pokémon (henceforth “TPP”) was a crowdsourced event in which everyone could type commands through the website’s chat window and try to finish the game that was being streamed, namely Pokémon Red. It took a little more than two weeks for the players to complete the game and this was more than enough time for the birth of an entirely new religion. But how exactly did that happen?

With thousands of people giving commands at the same time, there was a huge confusion and progress was very slow at the beginning. Then some programmer had the idea of initiating a system (named “Democracy Mode”) in which the game compiled votes every 10 seconds and the command inputted on the game was the one with most votes. People could vote to switch between Democracy and the original mode (hence renamed as “Anarchy Mode”) at any time. Most people preferred Anarchy, because it was supposedly more fun, and turned to Democracy only when it seemed otherwise impossible to advance in the game.

TPP is a very boring way of playing Pokémon and the players soon turned to other stuff in order to make it a little more exciting. They started to interpret whatever was happening in the game in a way that it would make sense from a cosmic point of view. And, as a matter of fact, many bad things were happening in the game – in the Anarchic world of TPP, bad moves and poor strategies were running amok. Not intentionally, mind you, but as a result of the way in which commands were given and computed. This way, items were discarded, pokémons were released and, even worse, eevees turned into flareons.


But let’s return to the Helix. One item in particular could not be discarded; it was the Helix Fossil (the fossilized shell of a ammonite-like pokémon). And, boy, people spent a lot of time in the inventory clicking on the Helix Fossil (and thus receiving in return the message “This isn’t the time to use that”). It did not take long for people to decide that the fossil was a god and that Red, the protagonist, was consulting it as a sort of oracle in order to discover the best way to proceed on his adventure.

Lord Helix. Artwork by Chlorine17 (http://chlorine17.deviantart.com/).

From this point onwards, the mythology of the Helix developed really fast. The Helix Fossil had been previously chosen by the players in spite of the Dome Fossil, which then became the Enemy, or the Devil, if you will. The Helix represented Anarchy Mode, while the Dome represented Democracy. The pidgeot, the most reliable pokémon in battle, became Bird Jesus; flareon became the False Prophet, a servant of the Dome Fossil; and many other pokémons received places in the mythology, accompanied by a lot of fanart on the internet. Long story short, eventually the players revived the fossil (yes, that’s possible in the game in a very Jurassic Park style) and received the pokémon omanyte in return. He was the resurrected god, Lord Helix. And then they went on to beat the game, but that’s not important – let’s take a closer look at the whole religion thing.


The Church of the Helix was born in a very short time span and possibly already have more followers than many of the world’s “true” religions. In a sense, Helixism has itself become a true religion and, more than that, it was created consciously through the consensus of a tribe (here defined as a group of people sharing the same interests and symbols). This is perhaps an example of Durkheim’s totemism. According to him, this is the most fundamental and primitive style of religion. The totem (here, the Helix) is a reflection of the tribe’s consciousness, chosen as a symbol to represent it. Symbols are an important part of any religion and the main pillar of totemism. Symbols are the representation (or perhaps translation) of the abstract principles of a religion in material form and, thus, allow the cult to develop and flourish.

Durkheim’s ideas were much disputed, of course (despite having received certain revival now in the light of research on the evolutionary roots of religious behavior), but the parallel was too strong to be ignored here. For Lévi-Strauss, for instance, the totem is a kind of animal with which a particular tribe identify themselves. In this case, it is not consciously chosen. Therefore, this view does not accommodate so nicely with the TPP’s Helix cult, since it was consciously (albeit somewhat accidently) chosen by its followers, which supposedly don’t identify themselves as an omanyte.

Granted, there are yet further difficulties: to begin with, Helixism was not born “naturally”, like a totemic religion developing in a group of humans some tens of thousands years ago. Rather, it was in a large part built on the common features of Christianity (including its symbology and usual artistic depictions). This, of course, merely reflect the cultural background of most players, but make comparisons with theoretical works more complicate and perhaps even more tenuous.

The Helix mythology. Artwork by Twarda8 (http://twarda8.deviantart.com/).

Of course, this is not a serious foray into the origins of religions in general or the meaning of a peculiar newborn religion. These are just some random thoughts that came to me when I first saw the Helix cult in all its glory. Helixism will probably never be treated seriously by its followers (well, at least I hope so). Still, the Church of the Helix functioned in its own manner as a true religion does, giving an identity to a group, making them stick together and driving them forwards (there was even a petition to make March 1st the National Helix Day in the USA). As such, it is a unique and amazing event and I do hope that somebody will someday seriously study it.


The Helix Fossil and the pokémons you get from it, omanyte and its evolved form omastar, are based on actual mollusks: the ammonites.

Top row: the helix fossil (left) and omastar (right), as they appear in official Pokémon artwork. Bottom row: Asteroceras sp. (left), an actual fossil ammonite shell from the Jurassic of England, and an artistic reconstruction of the animal (right), by N. Tamura (http://ntamura.deviantart.com/).

The ammonites are a completely extinct branch of cephalopod mollusks – besides ammonites, the class Cephalopoda comprises squids, octopuses, cuttlefish, nautiluses and the also extinct belemnites. Ammonites once ruled the seas and diversified in thousands upon thousands of species, but unfortunately, they died together with the dinosaurs in the great extinction event at the end of the Cretaceous. They received their name in ancient Rome, for the fossil shells were compared to the ram’s horns of the Egyptian god Ammon.

By the image above, one can see that both the fossil item and the pokémon are reasonably representative of ammonites (although the pokémon’s shell is positioned like a snail’s shell, not like a cephalopod’s!). But I do have an issue with the name: “helix” comes from the Greek, through Latin, and simply means “spiral”. Up to here, it is a fitting name. However, Helix (notice the italics) is already the name of a genus of land snails, which includes common garden snails and edible snails.

A Helix snail: Helix lucorum. Image taken from: Wikimedia Commons.

Land snails are, of course, gastropods, which is an entirely different class of mollusks altogether and only distantly related to the cephalopods (and thus to ammonites). They could at least have chosen a better name; a good deal of ammonites have names ending in “ceras”, for instance (which means “horn” in Greek). But Pokémon is a complete failure for names – gastrodon is another poorly named molluscan pokémon. But I’ll let this whole name deal slide just this once, since this fossil has spawned the first mythology ever based around a mollusk – and that is truly something to be happy about.

But since they have chosen the name Helix, I have a final comment to make (which may be somewhat disturbing for the faithful), for one must be consistent with his choices. “Helix” is feminine, so we would have a Lady Helix, not a Lord Helix. Unfortunately, pokémons still did not have genders in Pokémon Red (this feature was only introduced in the so called Generation II, i.e., the Gold/Silver games), so we will never know Helix’s gender for sure. In any case, I bet it would have been a surprise for the followers to discover that their god was actually a goddess.

Last but not least, if you have any important questions, feel free to do like Red and consult the Helix Fossil, in this charming website: http://askhelixfossil.com/#313usi.

Praise the Helix!


I’d like to thank the artists from deviantArt.com, who kindly let me use their works here.


Durkheim, E. (1912) The Elementary Forms of the Religious Life. George Allen & Unwin, London.

Leach, E. (2010) The Structural Study of Myth and Totemism. Routledge, London.

Lévi-Strauss, C. (1963) Totemism. Beacon Press, Boston.

Mithen, S. (1999) Symbolism and the supernatural. In: Dunbar, R.; Knight, C.; Power, C. (Eds.) The Evolution of Culture: A Historical and Scientific Overview. Rutgers University Press, New Jersey. Pp. 147–172.

Monks, N. & Palmer, P. (2002) Ammonites. Smithsonian Books, Washington D.C.

Wade, N. (2009) The Faith Instinct: How Religion Evolved and Why It Endures. Penguin Press, London.

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