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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Moa v Superman

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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During his heroic career Superman fought several foes. Some of these stories are truly memorable, like The Death of Superman (1992–1993), when he faced Doomsday. But many stories just ended up completely forgotten. Granted, there are some stories that most fans prefer to forget, like the film Batman v Superman: Dawn of Justice (2016), but some are curious or weird enough to eventually deserve a fresh look. The story I’m about to tell you is one of the latter kind.

This one happened during the first years of the so-called Bronze Age of Comics (1970–1985). Comic books from the Bronze Age retained lots of elements and conventions from the preceding Silver Age, but started to introduce stories more in tune with social issues, like racism and drugs. Likewise, comics also began including environmental issues and this is the topic I will focus on here. More specifically, on extinction.


It is the first story on Action Comics no. 425 (July 1973), written by Cary Bates, illustrated by Curt Swan and Frank Giacoia. It is called “The Last Moa on Earth!” and by the title alone, you can see it is about a giant extinct bird.

It’s a Bird… It’s a Plane… It’s Super– no, wait, it is actually a bird this time!

My goal here is to guide you through the story and offer some Biology inputs every now and then, explaining some things and “correcting” the bits the comics got wrong. I do know that writers should be free to invent and I wholeheartedly agree with that – it is science fiction after all! However, there are some sciency bits and pieces that are so simple to get right that there can be no excuse for giving the public wrong information.

The story starts off with hunter Jon Halaway in a New Zealand forest, being attacked by a giant flightless bird. He shoots and kills it, and decides to visit a local scientist (in Hawera, a town on the west coast of the North Island) to confirm his suspicions of the bird’s identity.

Elementary, my dear Halaway.

The scientist tells Halaway that he shot a bird thought to be extinct for 500 years and that there were once thousands of these animals in New Zealand. Both pieces of information are correct. Scientists estimated that there were circa 160,000 moa in New Zealand when Polynesian settlers arrived between 1,200 and 1,300 CE (Holdaway & Jacomb, 2000; Wilmshurst et al., 2010). There were nine species of moa in total and the Polynesians (who later became known as the Māori) had already extinguished them all by the early 1,400’s CE (Tennyson & Martinson, 2007; Perry et al., 2014).

The scientist then says that the bird was the largest of the moa species, Dinornis[1] maximus. While indeed this species was likely the largest[2], it inhabited only the South Island of New Zealand. The species from the North Island, where Halaway was hunting, is called Dinornis novaezealandiae. So the writer got the species wrong, but we cannot truly blame him: tens of moa “species” were described throughout the years, mostly because of the huge difference in size between the sexes of some species confused early researchers. Thus, the classification of moa species was really messed up until genetic studies started to be conducted from the late 1990’s onwards.

The skull of a North Island giant moa, Dinornis novaezealandiae. Source: Museum of New Zealand Te Papa Tongarewa (specimen MNZ S.242); ©Te Papa, all rights reserved.

On a similar note, D. maximus is actually an invalid name; the valid name for the South Island giant moa is D. robustus (Gill et al., 2010). That is because “D. maximus” was a second name given to describe the same species; to avoid confusion, only the first name ever used (D. robustus) is valid in these cases.

Halaway estimated the size of the slain moa at 12 feet (approximately 3.6 m), which is quite reasonable. The largest known specimens would have been 2 meters high at their backs or 3 meters high with their necks held straight up (something that they did not do; Tennyson & Martinson, 2007). Moreover, Halaway’s dead bird was a female, which are typically much larger than males in the two Dinornis species (Bunce et al., 2003; Tennyson & Martinson, 2007).

Box 1. What’s a moa anyway?

The moa belong to a group of birds called “ratites”, which also includes ostriches, emus, cassowaries, kiwi, rheas, and the extinct elephant birds. Recent research has shown that moa are not closely related to the other notable New Zealand ratites, the kiwi. Rather, they are closer to the charismatic South America tinamous[3] (Mitchell et al., 2014; Yonezawa et al., 2017). Since tinamous still retain some ability to fly, the moa’s ancestor was actually a flying bird (Gibbs, 2016).

The elegant crested tinamou, Eudromia elegans. Source: Wikimedia Commons (Evanphoto, 2009).

The loss of flight (alongside attaining a large body size) is a common occurrence on island environments where no mammalian predator is present. Other New Zealand species have also lost this ability; besides the kiwi (the typical example of a flightless bird), there are parrots (kakapo), rails (takahē) and wrens.



Halaway realizes that what he did was plain wrong. As mentioned above, during the Bronze Age comics became conscious of social and environmental problems – and extinction is a major problem, since it is usually our fault. This is important because, even though more than 350 years have elapsed after the last dodo was killed, most people still do not really grasp the idea that a species can disappear forever (Adams & Carwardine, 1900).

The “good” Mr. Halaway than devoted all his energy and resources into finding the slain moa’s egg. He succeeds and notes that the egg was being incubated in a hot spring with “strange fumes”. The egg was really big and appear egg-shaped in one panel and spherical in the other. Moa’s eggs were not spherical and not that large. Nevertheless, they were quite big and the largest known intact eggs are 20 and 25 cm tall (respectively, for the North Island and South Island Dinornis).

Of course the strange chemicals will grant the baby moa superpowers; otherwise this wouldn’t be a comic book.

Halaway finally arrives in Metropolis, where he is interviewed by none other than Clark Kent. On the highway, Halaway tells Clark that he wants to redeem himself of his “unforgivable deed” and hope that scientists will figure a way to use the egg to produce more moa. The repented hunter then faints, just as the baby moa hatches and escapes, throwing the car off-balance and into a river.

Clark takes off his suit and glasses and, after he’s more comfortable in his supersuit, saves Halaway and takes him to a hospital. Now I will cut the whole weird plot short and just say that the moa created an “organic link” (whatever that is) with Halaway via a microorganism, and was draining his energy. Typical crazy comic book stuff, but that’s not the point here. So let’s get back to the baby moa.

These “clawed terrors” were actually fluffy herbivores.


Superman starts searching Metropolis for the runaway moa and eventually finds it flying. Yes, flying – without wings, the comic-book moa flies by “thrashing its feet at super-speed”. In fact, Superman notices that the moa can fly faster than a super-sonic jet.

Also, even though just a few hours had passed since the moa escaped, when Superman found it, the bird had already doubled in size. And these were not the only superpowers granted to the moa by the mysterious fumes.

Yep, you read it right – that moa is flying with its feet.

Box 2. The moa’s archnemesis

The moa were herbivores, browsing on several types of leafy herbs, shrubs and trees (Wood et al., 2008). They were so abundant that it is thought their presence in New Zealand resulted in the evolution of a set of counter-measures in some plant lineages, which have small and hardened leaves, and sometimes also spines (Greenwood & Atkinson, 1977; Cooper et al., 1993; Worthy & Holdaway, 2002). But who ate the moa? Well, they were were so large that one would think they had no natural predators before the hungry Polynesians arrived. But that would be wrong – moa were hunted by giant eagles.

Naturally one would think of this – it is New Zealand after all! Source: The Hobbit: An Unexpected Journey (Warner Bros. Pictures, 2012), screen capture.

They are known as Haast’s eagles, after the naturalist who first described them, Sir Johann von Haast. They are the largest known true raptors, in both size and weight. They could reach a 2.6 m wingspan (somewhat smallish for their bulk) and 16 kg in weight, with females being larger (Brathwaite, 1992; Tennyson & Martinson, 2007). To hunt and eat their massive prey, Haast’s eagles had strong legs and feet, with huge claws. Unfortunately, these amazing birds could not survive after the moa became extinct and likely did not last much longer than 1,400 CE (Tennyson & Martinson, 2007).

The skull of a Haast’s eagle, Aquila moorei. Source: Museum of New Zealand Te Papa Tongarewa (specimen MNZ S. 22473); ©Te Papa, all rights reserved.


The moa also gained the ability to use its feathers as projectiles that could even pierce an elephant’s hide (according to Superman). Needless to say, birds cannot do that unless they are also Pokémon. Finally, the moa could instantly regrow lost limbs, a feat that few heroes (and absolutely no birds) can achieve.

Giant Moa uses Feather Barrage. It’s not very effective…
Holy regeneration, Batman!

After some more fighting, Superman understands that the bird just wants to go back home – to that place with the fumes and the lonely pink flower. Superman realizes that the flower is a “Quixa blossom”, as he calls it, and says it is a rare plant found only in northwest New Zealand.

Since my knowledge of plants is fairly limited, I asked a New Zealand botanist for help with this one. I was told that there is no flower with that name in the country and actually nothing that even remotely looks like it.

The “Quixa blossom” is actually the least believable thing in this whole story.

In any event, Superman finds the moa’s home and takes it back there, thus stopping the energy draining effect and saving Halaway. Superman then proclaims the area a “moa preserve” and sets up a fence around it. A thoughtful move, but one that completely overlooks the fact that the supermoa could fly.


The story ends with Halaway saying that “the world owns the moa another chance for survival”. Unfortunately, reality is not so kind: our species has wiped the moa off the face of the Earth and there is no second chance.

Overall, if you ignore the superpowers and the “organic link” stuff, this Superman story is actually a nice portrayal of an extinct species and its tragic fate on the hands of humankind. If nothing else, I hope it has inspired a reader somewhere to become a scientist or to fight to preserve other endangered animals.


Adams, D. & Carwardine, M. (1990) Last Chance to See. William Heinemann, London.

Brathwaite, D.H. (1992) Notes on the weight, flying ability, habitat, and prey of Haast’s Eagle (Harpagornis moorei). Notornis 39: 239–247.

Bunce, M.; Worthy, T.H.; Ford, T.; Hoppitt, W.; Willerslev, E.; et al. (2003) Extreme reversed sexual size dimorphism in the extinct New Zealand moa Dinornis. Nature 425: 172–175.

Cooper, A.; Atkinson, I.A.E.; Lee, W.G.; Worthy, T.H. (1993) Evolution of the moa and their effect on the New Zealand flora. Trends in Ecology & Evolution 8: 433–437.

Mitchell, K.J.; Llamas, B.; Soubrier, J.; Rawlence, N.J.; Worthy, T.H.; et al. (2014) Ancient DNA reveals elephant birds and kiwi are sister taxa and clarifies ratite bird evolution. Science 344: 898–900.

Gibbs, G. (2016) Ghosts of Gondwana: The History of Life in New Zealand. Fully Revised Edition. Potton & Burton, Nelson.

Gill, B.J.; Bell, B.D.; Chambers, G.K.; Medway, D.G.; Palma, R.L.; et al. (2010) Checklist of the Birds of New Zealand, Norfolk and Macquairie Islands, and the Ross Dependency, Antarctica. Te Papa Press, Wellington.

Greenwood, R.M. & Atkinson, I.A.E. (1977) Evolution of divaricating plants in New Zealand in relation to moa browsing. Proceedings of the New Zealand Ecological Society 24: 21–33.

Holdaway, R.N. & Jacomb, C. (2000) Rapid extinction of the moas (Aves: Dinornithiformis): model, test, and implications. Science 287: 2250–2254.

Perry, G.L.W.; Wheeler, A.B.; Wood, J.R.; Wilmshurst, J.M. (2014) A high-precision chronology for the rapid extinction of New Zealand moa (Aves, Dinornithiformes). Quaternary Science Reviews 105: 126–135.

Tennyson, A. & Martinson, P. (2007) Extinct Birds of New Zealand. Te Papa Press, Wellington.

Wilmshurst, J.M.; Hunt, T.L.; Lipo, C.P.; Anderson, A.J. (2011) High-precision radiocarbon dating shows recent and rapid initial human colonization of East Polynesia. PNAS 108(5): 1815–1820.

Worthy, T.H. & Holdaway, R.N. (2002) The Lost World of the Moa: Prehistoric Life of New Zealand. Canterbury University, Christchurch.

Wood, J.R.; Rawlence, N.J.; Rogers, G.M.; Austin, J.J.; Worthy, T.H.; Cooper, A. (2008) Coprolite deposits reveal the diet and ecology of the extinct New Zealand megaherbivore moa (Aves, Dinornithiformes). Quaternary Science Reviews 27: 2593–2602.

Yonezawa, T.; Segawa, T.; Mori, H.; Campos, P.F.; Hongoh, Y.; et al. (2017) Phylogenomics and morphology of extinct paleognaths reveal the origin and evolution of the ratites. Current Biology 27: 68–77. 


I am very grateful to Dr. Carlos Lehnebach for the help with flower, to Alan Tennyson for helping me to correct some mistakes on moa/eagle biology, and to Museum of New Zealand Te Papa Tongarewa for allowing the usage of the photographs herein.


Dr. Rodrigo Salvador is a paleontologist/ zoologist who studies mollusks, but just happens to have a soft spot for giant flightless birds. He is a diehard DC Comics fan, but to be honest, he never really liked Superman. Instead, he prefers to read the stories of the caped crusader and his extensive Gotham “family”.

[1] Dinornis means “terrible bird”, just like dinosaur means “terrible lizard”.

[2] The largest tibia (a leg bone) ever found belongs to this species, being 1 m long (Tennyson & Martinson, 2007).

[3] Tinamous are not typically included in the ratites group, rather being historically considered a separate (basal) lineage and grouped together with ratites in the more inclusive “palaeognaths” group. However, the work of Mitchell and collaborators (2014) have placed the tinamous well inside the ratites.

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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.

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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


The ichthyological diversity of Pokémon

Augusto B. Mendes1, Felipe V. Guimarães2, Clara B. P. Eirado-Silva1 & Edson P. Silva1

1Universidade Federal Fluminense, Niterói, RJ, Brazil.

2Universidade do Estado do Rio de Janeiro, São Gonçalo, RJ, Brazil.

Emails: augustobarrosmendes (at) yahoo (dot) com (dot) br; felipevieiragui (at) gmail (dot) com; clara.eirado (at) gmail (dot) com; gbmedson (at) vm (dot) uff (dot) br

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Pokémon, or Pocket Monsters, was originally created for videogames, becoming a worldwide fever among kids and teenagers in the end of the 1990’s and early 2000’s. Currently, it is still a success, with numerous games, a TV series, comic books, movies, a Trading Card Game, toys and collectibles. Through its core products and vibrant merchandising, Pokémon took over the world, influencing pop culture wherever it landed. Despite losing some steam in the early 2010’s, Pokémon is now back to its previous uproar with the release of Pokémon GO, an augmented reality (AR) game for smartphones. This game launched in 2016, with almost 21 million users downloading it in the very first week in the United States alone (Dorward et al., 2017). Thus, Pokémon is indubitably an icon in pop culture (Schlesinger, 1999a; Tobin, 2004).

The origin of Pokémon goes back to two role-playing video games (created by Satoshi Tajiri and released by Nintendo for the Game Boy; Kent, 2001): Pokémon Green and Pokémon Red, released in Japan in 1996. In the West, the Green version never saw the light of day, but the Red and Blue versions were released in 1998, selling together more than 10 million copies. Also in 1998, the Yellow version of the game was released, which has as its most distinct feature the possibility of having Pikachu (the most famous Pokémon) walking side by side with the player in the game. Pokémon Green, Red, Blue and Yellow are the so-called “first generation” of games in the franchise. Today, the Pokémon series is in its seventh generation, with 29 main games released, besides several spin-offs. The TV series, on the other hand, is in its sixth season, with more than 900 episodes.

The games and TV series take place in regions inhabited by many Pokémon and humans. The mission of the protagonist is to win competitions (“Pokémon battles”) against gym leaders who are spread across different cities and regions. For each victory, the protagonist receives a gym badge; with eight badges, he/she is allowed to enter the Pokémon League to try and become the Champion.

For each generation, new Pokémon (and an entire new region) are introduced. In this way, the creatures have a homeland, although most can appear in other regions as well (Schlesinger, 1999b; Whitehill et al., 2016). The seven main regions are: Kanto, Johto, Hoenn, Sinnoh, Unova, Kalos and Alola.

In every region, there are numbered routes that connect cities and landmarks and in which the protagonist travels, finding the monsters in their natural habitats and interacting with other characters. These routes comprise a great range of environments, such as forests, caves, deserts, mountains, fields, seas, beaches, underwater places, mangroves, rivers and marshes, which usually display a huge diversity of Pokémon.

In addition to winning the Pokémon League, the protagonist must complete the Pokédex, a digital encyclopedia of Pokémon. In other words, the trainer must catch all the Pokémon that live in that region, registering each capture in the Pokédex. Each Pokémon has a registry number and an entry text in the Pokédex. Pokémon are usually found in nature, and may be captured with a device called “Pokéball”. Pokéballs are small enough to fit in a pocket, hence the name “Pocket Monsters” (Whitehill et al., 2016).


In the world depicted in the games, there are 801 Pokémon, belonging to one or two of the following 18 types: Normal, Fire, Fighting, Water, Flying, Grass, Poison, Electric, Ground, Psychic, Rock, Ice, Bug, Dragon, Ghost, Dark, Steel and Fairy (Bulbapedia, 2017). Almost all Pokémon are based on animal species, some of them are based on plants or mythological creatures, and a few are based on objects. Curiously, all Pokémon are oviparous, which means they all lay eggs (their development happens inside of an egg and outside of their mother’s body); of course, in the real natural world, this is a reproductive strategy of animals such as fishes, amphibians, reptiles, birds and many kinds of invertebrates (Blackburn, 1999). Moreover, Pokémon might “evolve”, usually meaning they undergo some cosmetic changes, become larger and gain new powers.

In the present work, the Pokémon world was approached by analogies with the real natural world, establishing parallels with actual animals.

A remarkable group of animals represented in Pokémon is the fishes. Fishes are the largest group of vertebrates, with more than 32,000 species inhabiting marine and freshwater environments, a number that roughly corresponds to half of all described vertebrates (Nelson et al., 2016). Showing ample morphological and behavioral variety and living in most of the aquatic ecosystems of the planet, fishes are well represented in the Pokémon world, therefore offering a great opportunity for establishing parallels between the two worlds. The creators of the games not only used the morphology of real animals as a source of inspiration for the monsters, but also their ecology and behavior.

Based on these obvious connections between real fishes and Pokémon, the aim of this work is to describe the ichthyological diversity found in Pokémon based on taxonomic criteria of the classification of real fishes. Ultimately, our goal is to offer useful material for both teaching and the popularization of science.

Table 1. Taxonomic classification of the fish Pokémon. Abbreviations: Ch = Chondrichthyes; Gn = Gnathostomata; Pe = Petromyzontomorphi; Pt = Petromyzontida; Os = Osteichthyes. All images obtained from The Official Pokémon Website (2016).


The first step of our research was a search in the Pokédex (The Official Pokémon Website, 2016) for Pokémon which were related to fishes. The criterion used was the Pokémon’s morphology (resemblance to real fishes). Afterwards, the “fish Pokémon” were classified to the lowest taxonomic level (preferably species, but when not possible, genus, family or even order).

This classification of the Pokémon allowed the comparison of biological data (such as ecological, ethological, morphological traits) from Bulbapedia (2017) with the current knowledge on real fishes (e.g., Nelson et al., 2016). Bulbapedia is a digital community-driven encyclopedia created in 2004 and is the most complete source regarding the pocket monsters.

The final step was a search in online scientific databases (Fishbase, Froese & Pauly, 2016; and Catalog of Fishes, Eschmeyer et al., 2016) in order to obtain the current and precise taxonomy and additional information on habitats, ecology etc. of the fish species.

In the present work, the taxonomic classification used was that proposed by Nelson et al. (2016), who consider the superclasses Petromyzontomorphi (which includes the class Petromyzontida, that is, the lampreys) and Gnathostomata (the jawed vertebrates). Gnathostomata, in turn, includes the classes Chondrichthyes (cartilaginous fishes) and Osteichthyes (bony fishes). Along with this classification, we used the classification proposed by the database ITIS (Integrated Taxonomic Information System, 2016) for comparison at all taxonomic levels. Following identification, the “fish Pokémon” were described regarding their taxonomic and ecological diversity.


As a result of our search, 34 fish Pokémon were identified (circa 4% of the total 801 Pokémon; Table 1) and allocated in two superclasses, three classes, eighteen orders, twenty families and twenty-two genera. Eighteen of the 34 fish Pokémon (circa 53%) could be identified to the species level (Table 2). The features of the real fishes which probably inspired the creation of the Pokémon and other relevant information are described below for each species. To enrich the comparisons, images of the Pokémon (obtained from the Pokédex of The Official Pokémon Website; http://www.pokemon.com) and of the real fishes (illustrations by one of us, C.B.P. Eirado-Silva) follow the descriptions.

Table 2. Taxonomic diversity of the fish Pokémon.

Horsea and Seadra

Species: Hippocampus sp.; Common name: seahorse.

The Pokémon Horsea and Seadra (Fig. 1), which debuted in the first generation of the franchise, were based on seahorses. The long snout, ending in a toothless mouth (Foster & Vincent, 2004), the prehensile, curved tail (Rosa et al., 2006) and the salient abdomen are features of the real fishes present in these Pokémon. Seahorses belong to the genus Hippocampus, presently composed of 54 species (Nelson et al., 2016). The males have a pouch in their bellies where up to 1,000 eggs are deposited by the females. In this pouch, the eggs are fertilized and incubated for a period ranging from 9 to 45 days (Foster & Vincent, 2004). Due to overfishing for medicinal and ornamental purposes, as well habitat destruction, about 33 species of seahorses are considered threatened (Rosa et al., 2007, Castro et al., 2008; Kasapoglu & Duzgunes, 2014).

Figure 1. Horsea, Seadra and Hippocampus sp.

Goldeen and Seaking

Species: Carassius auratus; Common name: goldfish.

Goldeen and Seaking (Fig. 2) were based on the goldfish. This species is one of the most common ornamental fishes worldwide (Soares et al., 2000; Moreira et al., 2011) and it is widely used in studies of physiology and reproduction due to its docile behavior and easy acclimatization to artificial conditions (Bittencourt et al., 2012; Braga et al., 2016). The resemblance between the goldfish and the Pokémon include morphological features, such as the orange/reddish color and the long merged fins, and the name “Goldeen”. The name Seaking, on the other hand, may be a reference to another common name of the species, “kinguio”, from the Japanese “kin-yu” (Ortega-Salas & Reyes-Bustamante, 2006).

Figure 2. Goldeen, Seaking and Carassius auratus.


Species: Cyprinus carpio; Common name: common carp.

Possibly the most famous fish Pokémon, Magikarp (Fig. 3) was based on a common carp, a species present in Europe, Africa and Asia, widely used in pisciculture due to its extremely easy acclimatization to many freshwater environments and the high nutritional value of its meat (Stoyanova et al., 2015; Mahboob et al., 2016; Voigt et al., 2016). In some regions of the planet, such as Brazil, the common carp is considered an invasive species, as it was inadvertently released in the wild and poses a threat to the native aquatic fauna (Smith et al., 2013; Contreras-MacBeath et al., 2014).

Figure 3. Magikarp and Cyprinus carpio.

The shared traits between the Pokémon and the real fish are many: the rounded mouth, the lips, the strong orange color and the presence of barbels (“whiskers”) (Nelson et al., 2016). In China, the common carp is praised as an animal linked to honor and strength, due of its ability to swim against the current; an ancient legend tells about carps that swim upstream, entering through a portal and transforming into dragons (Roberts, 2004). In Pokémon, Magikarp evolves into Gyarados, which resembles a typical Chinese dragon.

Chinchou and Lanturn

Species: Himantolophus sp.; Common name: footballfish.

Chinchou and Lanturn (Fig. 4) were based on fishes of the genus Himantolophus, a group of deep-sea fishes found in almost all oceans living in depths up to 1,800 meters (Klepadlo et al., 2003; Kharin, 2006). These fishes are known as footballfishes, a reference to the shape of their bodies. Fishes of this genus have a special modification on their dorsal fin that displays bioluminescence (the ability to produce light through biological means; Pietsch, 2003), which is used to lure and capture prey (Quigley, 2014). Bioluminescence was the main inspiration for these Pokémon, which have luminous appendages and the Water and Electric types. The sexual dimorphism (difference between males and females) is extreme in these fishes: whilst females reach up to 47 cm of standard-length (that is, body length excluding the caudal fin), males do not even reach 4 cm (Jónsson & Pálsson, 1999; Arronte & Pietsch, 2007).

Figure 4. Chinchou, Lanturn and Himantolophus sp.


Species: Diodon sp.; Common name: porcupinefish.

Qwilfish (Fig. 5) was based on porcupinefishes, more likely those of the genus Diodon, which present coloring and spines most similar to this Pokémon. Besides the distinctive hard, sharp spines (Fujita et al., 1997), porcupinefishes have the ability to inflate as a strategy to drive off predators (Raymundo & Chiappa, 2000). As another form of defense, these fishes possess a powerful bacterial toxin in their skin and organs (Lucano-Ramírez et al., 2011; Ravi et al., 2016). Accordingly, Qwilfish has both Water and Poison types.

Figure 5. Qwilfish and Diodon sp.


Species: Remora sp.; Common names: remora, suckerfish.

Remoraid was based on a remora (Fig. 6), a fish with a suction disc on its head that allows its adhesion to other animals such as turtles, whales, dolphins, sharks and manta rays (Fertl & Landry, 1999; Silva & Sazima, 2003; Friedman et al., 2013; Nelson et al., 2016). This feature allows the establishment of a commensalisc or mutualisc relationship of transportation, feeding and protection between the adherent species and its “ride” (Williams et al., 2003; Sazima & Grossman, 2006). The similarities also include the name of the Pokémon and the ecological relationship they have with other fish Pokémon: in the same way remoras keep ecological relationships with rays, Remoraid does so with Mantyke and Mantine (Pokémon based on manta rays; see below).

Figure 6. Remoraid and Remora sp.

Mantyke and Mantine

Species: Manta birostris; Common name: manta ray.

The Pokémon Mantyke and its evolved form Mantine (Fig. 7) were probably based on manta rays of the species Manta birostris, which inhabits tropical oceans (Duffy & Abbot, 2003; Dewar et al., 2008) and can reach more than 6 meters of wingspan, being the largest species of ray in existence (Homma et al., 1999; Ari & Correia, 2008; Marshall et al., 2008; Luiz et al., 2009; Nelson et al., 2016). The similarities between the Pokémon and the real fish are: the body shape, the color pattern, the large and distinctive wingspan and even the names.

Figure 7. Mantine, Mantyke and Manta birostris.

Kingdra and Skrelp

Species: Phyllopteryx taeniolatus; Common name: common seadragon.

Kingdra and Skrelp (Fig. 8) were based on the common seadragon. The resemblances between these Pokémon and the real fish species include the leaf-shaped fins that help the animals to camouflage themselves in the kelp “forests” they inhabit (Sanchez-Camara et al., 2006; Rossteuscher et al., 2008; Sanchez-Camara et al., 2011), and the long snout. Also, the secondary type of Kingdra is Dragon. Although both are based on the common seadragon, Kingdra and Skrelp are not in the same “evolutionary line” in the game.

Common seadragons, as the seahorses mentioned above, are of a particular interest to conservationists, because many species are vulnerable due to overfishing, accidental capture and habitat destruction (Foster & Vincent, 2004; Martin-Smith & Vincent, 2006).

Figure 8. Kingdra, Skrelp and Phyllopteryx taeniolatus.


Species: Pygocentrus sp.; Common name: red piranha.

Piranhas of the genus Pygocentrus possibly were the inspiration for the creation of Carvanha (Fig. 9), a Pokémon of voracious and dangerous habits. The main feature shared by the real fish and the Pokémon is the color pattern: bluish in the dorsal and lateral areas, and reddish in the ventral area (Piorski et al., 2005; Luz et al., 2015).

It is worthwhile pointing out that, despite what is shown in movies and other media, piranhas do not immediately devour their prey; instead, they tear off small pieces, bit by bit, such as scales and fins (Trindade & Jucá-Chagas, 2008; Vital et al., 2011; Ferreira et al., 2014).

Figure 9. Carvanha and Pygocentrus sp.


Order: Carcharhiniformes; Common name: shark.

Sharpedo (Fig. 10), according to its morphological traits (elongated fins), was possibly based on sharks of the order Carcharhiniformes, the largest group of sharks, with 216 species in 8 families and 48 genera. Fishes in this order are common in all oceans, in both coastal and oceanic regions, and from the surface to great depths (Gomes et al., 2010). Several species of Carcharhiniformes are in the IUCN’s (International Union for Conservation of Nature) endangered species list (a.k.a. “Red List”) due to overfishing, as their fins possess high commercial value (Cunningham-Day, 2001).

Figure 10. Sharpedo and a carcharhiniform shark.


Species: Misgurnus sp.; Common name: pond loach.

Barboach (Fig. 11) is likely based on fishes of the genus Misgurnus, natively found in East Asia (Nobile et al., 2017) but introduced in several countries (Gomes et al., 2011). These animals, like M. anguillicaudatus Cantor, 1842, are used as ornamental fishes and in folk medicine (Woo Jun et al., 2010; Urquhart & Koetsier, 2014). The shared similarities between the Pokémon and the pond loach include morphological features, such as the elongated body, oval fins and the presence of barbels (Nelson et al., 2016). The resemblance also extends itself to behavior, such as the habit of burying in the mud (Zhou et al., 2009; Kitagawa et al., 2011) and using the barbels to feel the surroundings (Gao et al., 2014). The secondary type of Barboach, Ground, alongside the ability to feel vibrations in the substrate, seem to be a reference to the behavior of the real fishes.

Figure 11. Barboach and Misgurnus sp.


Species: Silurus sp.; Common name: catfish.

Whiscash (Fig. 12) was based on the Japanese mythological creature Namazu, a gigantic catfish that inhabits the underground realm and is capable of creating earthquakes (Ashkenazi, 2003). Namazu also names the Pokémon in the Japanese language (“Namazun”). In Japan, fishes of the genus Silurus are usually associated with this mythological creature and even the common name of these fishes in that country is “namazu” (Yuma et al., 1998; Malek et al., 2004). In addition, the physical traits of the Silurus catfishes also present in Whiscash are the long barbels (or “whiskers”, hence the name Whiscash) and the robust body (Kobayakawa, 1989; Kiyohara & Kitoh, 1994). In addition to the Water type, Whiscash is also Ground type, which is related to Namazu’s fantastic ability of creating earthquakes.

Figure 12. Whiscash and Silurus sp.


Species: Micropterus salmoides; Common name: largemouth bass.

The Pokémon Feebas (Fig. 13), a relatively weak fish (as its name implies), was possibly based on a largemouth bass, a freshwater fish native to North America (Hossain et al., 2013). The species was introduced in many countries and is often considered a threat to the native fauna (Welcomme, 1992; Hickley et al., 1994; Godinho et al., 1997; García-Berthou, 2002). Similarities between Feebas and the largemouth bass include the large, wide mouth and the brownish coloration, with darker areas (Brown et al., 2009).

Figure 13. Feebas and Micropterus salmoides.


Species: Regalecus sp.; Common name: oarfish.

Often praised as the most beautiful Pokémon of all (Bulbapedia, 2017), Milotic (Fig. 14) certainly lives up to its title. Their long reddish eyebrows were based on the first elongated rays of the dorsal fin of Regalecus species (Nelson et al., 2016), which also share the reddish color of the dorsal fin (Carrasco-Águila et al., 2014). Other similarities between the oarfish and the Pokémon are the elongated body (some oarfishes can grow larger than 3.5 m) and the spots scattered on the body (Chavez et al., 1985; Balart et al., 1999; Dulčić et al., 2009; Ruiz & Gosztonyi, 2010).

Figure 14. Milotic and Regalecus sp.


Species: Monognathus sp.; Common name: onejaw.

Probably based on fishes of the genus Monognathus, which have a large mandible and a long dorsal fin (Nelson et al., 2016), Huntail (Fig. 15) is one of the possible evolutionary results of the mollusk Pokémon Clamperl (the other possibility is Gorebyss; see below). According to Raju (1974), fishes of the genus Monognathus live in great depths and have a continuous dorsal fin that ends in an urostyle (“uro” comes from the Greek language and means “tail”, an element also present in the Pokémon’s name).

Figure 15. Huntail and Monognathus sp.


Family: Nemichthyidae; Common name: snipe eel.

The serpentine body and the thin beak-shaped jaw of Gorebyss (Fig. 16) are features of fishes belonging to the family Nemichthyidae (Nielsen & Smith, 1978). These fishes inhabit tropical and temperate oceans and can be found in depths up to 4,000 meters, in the so-called “abyssal zone” (Cruz-Mena & Anglo, 2016). The Pokémon’s name may be a reference to such habitat.

Figure 16. Gorebyss and a nemichthyid fish.


Species: Latimeria sp.; Common name: coelacanth.

Relicanth (Fig. 17) was based on the coelacanth. The brown coloration, the lighter patches on the body (Benno et al., 2006) and the presence of paired lobed fins (Zardoya & Meyer, 1997) are traits of both the real fish and the Pokémon. It was believed that coelacanths went extinct in the Late Cretaceous, but they were rediscovered in 1938 in the depths off the coast of South Africa (Nikaido et al., 2011). Therefore, the only two living species L. chalumnae Smith, 1939 and L. menadoensis Pouyaud et al., 1999 are known as “living fossils” (Zardoya & Meyer, 1997). Probably for this reason, Relicanth belongs to the Water and Rock types (the “fossil Pokémon” are all Rock-type).

Figure 17. Relicanth and Latimeria sp.


Species: Helostoma temminckii; Common name: kissing gourami.

The silver-pinkish coloration, the peculiar mouth formed by strong lips and the habit of “kissing” other individuals of their species (which is actually a form of aggression!) are features of the kissing gourami (Sterba 1983; Sousa & Severi 2000; Sulaiman & Daud, 2002; Ferry et al., 2012) that are also seen in Luvdisc (Fig. 18). Helostoma temminckii is native to Thailand, Indonesia, Java, Borneo, Sumatra and the Malay Peninsula (Axelrod et al., 1971), but due to its use an ornamental fish and the irresponsible handling by fishkeepers, it has been introduced in other parts of the world (Magalhães, 2007).

Figure 18. Luvdisc and Helostoma temminckii.

Finneon and Lumineon

Species: Pantodon buchholzi; Common name: freshwater butterflyfish.

Finneon and Lumineon (Fig. 19) were probably based on the freshwater butterflyfish. Finneon has a caudal fin in the shape of a butterfly and Lumineon, like Pantodon buchholzi, has large pectoral fins (Nelson et al., 2016) resembling the wings of a butterfly (hence the popular name of the species). Butterflyfishes are found in West African lakes (Greenwood & Thompson, 1960); their backs are olive-colored while their ventral side is silver, with black spots scattered throughout the body; their fins are pink with some purplish spots (Lévêque & Paugy, 1984). Both Pokémon have color patterns that resemble the freshwater butterflyfish.

Figure 19. Finneon, Lumineon and Pantodon buchholzi.


Family: Serrasalmidae; Common name: piranha.

The two forms of the Pokémon Basculin (Fig. 20) seem to have been inspired on fishes from the Serrasalmidae family, such as piranhas. Basculin, like these fishes, has a tall body and conical teeth (Baumgartner et al., 2012). Piranhas are predators with strong jaws that inhabit some South American rivers. Curiously, they are commonly caught by local subsistence fishing (Freeman et al., 2007).

Figure 20. Basculin’s two forms and a serrasalmid fish.


Species: Mola mola; Common name: sunfish.

The very name of this Pokémon is evidence that it was inspired on Mola mola, the sunfish (Fig. 21). Moreover, Alomomola, just like the sunfish, has a circular body with no caudal fin (Pope et al., 2010). The sunfish is the largest and heaviest bony fish in the world, weighting more than 1,500 kg (Freesman & Noakes, 2002; Sims et al., 2009). They inhabit the Atlantic and Pacific Oceans, feeding mainly on zooplankton (Cartamil & Lowe, 2004; Potter & Howell, 2010).

Figure 21. Alomomola and Mola mola.

Tynamo, Eelektrik and Eelektross

Species: Petromyzon marinus; Common name: sea lamprey.

The evolutionary line Tynamo, Eelektrik and Eelektross (Fig. 22) was probably inspired by the life cycle of the sea lamprey, Petromyzon marinus: Tynamo represents a larval stage, Eelektrik a juvenile, and Eelektross an adult. As a larva, the sea lamprey inhabits freshwater environments and, after going through metamorphosis, the juvenile migrates to the ocean, where they start to develop hematophagous (“blood-sucking”) feeding habits (Youson, 1980; Silva et al., 2013). Eelektrik and Eelektross, like the sea lamprey, have a serpentine body and a circular suction cup-mouth with conical teeth. In addition, the yellow circles on the side of these Pokémon resemble the gill slits of lampreys (which are of circular shape) or the marbled spots of P. marinus (Igoe et al., 2004).

It is worth mentioning that Eelektrik and Elektross also seem to possess name and characteristics (Electric type and serpentine body with yellow spots) inspired by the electric eel (Electrophorus electricus Linnaeus, 1766), a fish capable of generating an electrical potential up to 600 volts, making it the greatest producer of bioelectricity in the animal kingdom (Catania, 2014). However, a remarkable characteristic of Eelektrik and Eelektross is the jawless mouth structure of the superclass Petromyzontomorphi species. The electric eel has a jaw and thus belongs to the superclass Gnathostomata (jawed vertebrates) (Gotter et al., 1998).

Figure 22. Tynamo, Eelektrik, Eelektross and P. marinus.


Order: Pleuronectiformes; Common name: flatfish.

Flattened and predominantly brown in color, Stunfisk (Fig. 23) appears to have been based on fishes of the order Pleuronectiformes. Popularly known as flatfishes, these animals have both eyes on the same side of the head and stay most of their lives buried and camouflaged on sandy and muddy substrates of almost every ocean, feeding on fishes and benthic invertebrates (Sakamoto, 1984; Kramer, 1991; Gibb, 1997). It is likely that the primary type of Stunfisk, Ground, is based on the close relationship between pleuronectiform fishes and the substrate they live in. Species of this group are very valuable for the fishing industry (Cooper & Chapleau, 1998).

Figure 23. Stunfisk and a pleuronectiform fish.


Species: Phycodurus eques; Common name: leafy seadragon.

Dragalge (Fig. 24), a Pokémon belonging to the Poison and Dragon types, was based on a leafy seadragon. This species is found in Australia and it is named after its appearance: this fish has appendages throughout its body that resemble leaves (Larson et al., 2014). This feature, also present in the Pokémon, allows the leafy seadragon to camouflage itself among algae (Wilson & Rouse, 2010). Dragalge is the evolved form of Skrelp, a Pokémon based on a common seadragon (see above).

Figure 24. Dragalge and Phycodurus eques.


Species: Sardinops sagax; Common name: Pacific sardine.

Wishiwashi (Fig. 25) was probably based on the Pacific sardine, a pelagic fish with high commercial value and quite abundant along the California and Humboldt Currents (Coleman, 1984; Gutierrez-Estrada et al., 2009; Demer et al., 2012; Zwolinski et al., 2012). The lateral circles of the Pokémon are a reference to the dark spots present on the lateral areas of the real fish (Paul et al., 2001). Furthermore, Wishiwashi has the ability to form a large school, just as sardines do (Emmett et al., 2005; Zwolinski et al., 2007).

Figure 25. Wishiwashi and Sardinops sagax.

Another parallel is the geographic location: the Pokémon belongs to Alola, a fictional region based on Hawaii, and S. sagax is one of the most common sardines in the Eastern Pacific Ocean. From the mid-1920’s to the mid-1940’s, for example, S. sagax supported one of the largest fisheries in the world. The stock collapsed in the late 1940’s, but in the 1990’s it started to recover (McFarlane et al., 2005).


Species: Rhinecanthus rectangulus; Common name: reef triggerfish.

Bruxish (Fig. 26) was probably inspired by the species Rhinecanthus rectangulus, the reef triggerfish of the Hawaiian reefs and other tropical regions (Kuiter & Debelius, 2006; Dornburg et al., 2008). Bruxish has powerful jaws, just like the reef triggerfishes that prey upon a wide variety of invertebrates, such as hard-shelled gastropods, bivalves, echinoderms and crustaceans (Wainwright & Friel, 2000; Froese & Pauly, 2016).

Figure 26. Bruxish and Rhinecanthus rectangulus.

Besides the strong jaw, the overall body shape and the flashy coloring, another parallel can be seen: this Pokémon is an inhabitant of the Alola region (the Pokémon version of Hawaii) and R. rectangulus is actually the state symbol fish of the Hawaiian archipelago (Kelly & Kelly, 1997).


The majority of the identified Pokémon (85.29%) is, expectedly, Water-type. A large portion of them (29.41%) was introduced for the first time in the third generation of the franchise, in the Hoenn region.

Figure 27. Representativeness of fish classes in Pokémon.

Only three fish Pokémon were classified in the superclass Petromyzontomorphi (8.82%): the lamprey-like Tynamo, Eelektrik and Eelektross, all of them belonging to the same evolutionary line. In the superclass Gnathostomata, the class Osteichthyes is represented by the highest number of Pokémon: 28 in total (82.35%, Fig. 27). Inside this class, the most representative groups were the order Syngnathiformes (14.71%, Fig. 28), family Syngnathidae (15.63%, Fig. 29) and the genus Petromyzon (10.00%, Fig. 30).

Figure 28. Representativeness of fish orders in Pokémon.

Most of the real fishes on which the Pokémon were based (55.88%, Fig. 31) live in marine environments, followed by freshwater (continental water environments, 32.35%) and finally, brackish water (estuarine environments, 11.76%).

The “fish” species found in the Pokémon world consists of a considerable portion of the ichthyological diversity in our world. According to Nelson et al. (2016), the Osteichthyes class corresponds to 96.1% of all vertebrate fish species (30,508 species), followed by the Condrichthyes with 3.76% (1,197 species) and the Petromyzontida with just 0.14% (46 species). In Pokémon, the proportions of taxa (taxonomic group) that inspired the creatures follow a roughly similar distribution: within the 26 taxa in which the evolutionary families of the Pokémon were based, 23 are Osteichthyes class (88.46%), two are Condrichthyes (7.7%) and one is Petromyzontida (3.84%). If the games follow a pattern of introducing more fish Pokémon over time, it is expected that these proportions will gradually become more equivalent as each new generation of the franchise is released.

Figure 29. Representativeness of fish families in Pokémon.


Our analysis shows that fish Pokémon are very diverse creatures, both taxonomic and ecologically, despite being a small group within the Pokémon universe (with 801 “species”).

The fish Pokémon are represented by several orders, families and genera of real fishes and, as previously stated, this is actually a relevant sampling of the ichthyological diversity of our planet. The marine Pokémon described here are inhabit from abyssal zones to coastal regions, including reefs. The creative process of the fish monsters in the game must have included a fair share of research on real animals.

Figure 30. Representativeness of fish genera in Pokémon.

The Hoenn region, which has the largest playable surface and includes areas with “too much water”, is also the region with the highest number of fish Pokémon. Furthermore, the majority of these Pokémon live in the marine environment and belongs to the Osteichthyes class, as is observed for real fishes (Nelson et al., 2016; Eschmeyer et al., 2016). However, it is also important to underline that marine fishes are those with the more attractive colors and shapes and, therefore, higher popular appeal, which is vital for a game based in charismatic monsters (Darwall et al., 2011; McClenachan, 2012; Dulvy et al., 2014).

Figure 31. Environments inhabited by the fish Pokémon.

In the present work, the analogy between fish Pokémon and real species allowed a descriptive study of the “Pokéfauna” in a similar manner in which actual faunal surveys are presented. These surveys are an important tool for understanding the structure of communities and to evaluate the conservation status of natural environments (Buckup et al., 2014). It is noteworthy that the association of the monsters with real fishes was only possible because Pokémon have several morphological, ecological and ethological traits that were based on real species.

Pokémon is a successful franchise and many of its staple monsters are already part of the popular imaginary. The creation of the pocket monsters was not done in a random manner; they were mostly inspired by real organisms, particularly animals, and often have specific biological traits taken from their source of inspiration. Thus, analogies between Pokémon and our natural world, such as the ones performed here, open a range of possibilities for science outreach.


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Augusto Mendes began his journey as a Pokémon trainer in his childhood, when his parents gave him a green Game Boy Color with Pokémon Red for Christmas. Currently, he is a master’s degree student in the Program of Marine Biology and Coastal Environments of UFF, where he works with zooarchaeology of fishes and education.

Felipe Guimarães is in love with Pokémon (since he first watched the TV series) and the natural world. He graduated in Biology from the UERJ, where he worked with taxonomy and ecology of fishes. He also works with popularization of science and environmental education.

Clara Eirado-Silva, when she was eight years old, told her parents she would study sharks. She has always been passionate about art too and draw since her childhood. Currently, she holds a “Junior Science” scholarship, working on fishing ecology with emphasis on reproductive biology. In her free time, she draws her much loved fishes.

Although Pokémon is not exactly Dr. Edson Silva’s cup of tea, he watched all movies with his daughter, who’s crazy about the little monsters. As fate would have it, his work on population genetics of marine organisms attracted a master’s student (A.B.M.) who’s an equally crazy pokéfan. May Arceus not spare him from the monsters!

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Barbara M. Tomotani1 & Rodrigo B. Salvador2

1 Netherlands Institute of Ecology; Wageningen, The Netherlands. Rijksuniversiteit Groningen; Groningen, The Netherlands. Email: babi.mt (at) gmail (dot) com

2 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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The video game Overwatch is Blizzard Entertainment’s new hit, released on May 2016 for Microsoft Windows, PlayStation 4 and Xbox One. In the game, the so-called “heroes” spend most of their time trying to kill each other to secure a payload. While the morals of these self-proclaimed heroes are rather open to debate, one of them at least has some redeeming personality traits. The hero Bastion is a nature-loving animal-friend robot. Actually, the single animal to appear in the whole game (besides the hominids, of course) is Bastion’s pet bird, called Ganymede (Fig. 1).


Figure 1. Left: Bastion with Ganymede (official artwork from the game). Image extracted from Overwatch Wiki. Right: Ganymede (official artwork from the game). Image extracted from “Bastion Reference Kit” (official Overwatch website).

Ganymede’s design is an original creation of Blizzard’s artists, although it resembles in shape and size a northern cardinal, Cardinalis cardinalis (Linnaeus, 1758), a common species in Canada and the USA. Cardinals usually have a red plumage (Fig. 2A), but there are rare naturally occurring yellow mutants, called xanthochroic cardinals (Fig. 2B). Ganymede also has a white area around its eyes, a trait not seen in cardinals, but well-known from species of the genus Zosterops (commonly known as “white-eyes”; Fig. 2C), which live in tropical Africa, Southeast Asia and Australasia.

bastion-fig-2Figure 2. Left: A male northern cardinal, Cardinalis cardinalis. Photo by Stephen Wolfe (2011); image extracted and modified from Wikimedia Commons. Center: A xanthochroic northern cardinal. Photo by Jim McCormac (2013), extracted from “Ohio Birds and Biodiversity”. Right: A Japanese white-eye, Zosterops japonicas (Temminck & Schlegel, 1847). Photo by Laitche (2016); image extracted and modified from Wikimedia Commons.

Despite being based in an American species, Ganymede seems to be native to European forests. The bird appears on its home forest in the animated short The Last Bastion (from August 2016), which takes place in the outskirts of Stuttgart, Germany. There is no bird here in Stuttgart that looks like Ganymede (one of us lives here, by the way). Actually, in the whole European bird fauna, only the golden oriole, Oriolus oriolus (Linnaeus, 1758), comes close to it, with its yellow color and dark horizontal stripe across the eyes (Fig. 3). However, its slenderer body shape, thinner beak and lack of crest are all very different from Ganymede.


Figure 3. A male golden oriole (Oriolus oriolus). Photo by Pawel Ryszawa (2008); image extracted from Wikimedia Commons.

Moreover, in the Eichenwalde stage (which, in the game’s lore, is located nearby Stuttgart), there is a painting resting above the hunting lodge’s fireplace (Fig. 4). This painting shows four local bird species; one of them is the “Ganymede species”, while the others seem to be actual species: the Eurasian blue tit (Cyanistes caeruleus (Linnaeus, 1758)) and two titmice. The latter are American species and seem to represent the tufted titmouse (Baeolophus bicolor (Linnaeus, 1766)), even though one of them is more bluish in color.


Figure 4. Fireplace of the hunting lodge in the Eichenwalde stage, with close-up of the painting. Screenshots from the game.

As we pointed out before, Ganymede’s design is an original creation and does not represent an actual species, although some of its features might be traced to the cardinal. Despite the problems with Ganymede’s identification, Bastion’s bird friend can also appear in the guise of actual real-life bird species. To do so, the player must simply equip a “skin” for Bastion (”skin” is basically the gaming jargon for “outfit”). By changing Bastion’s “skin”, Ganymede’s appearance may also change.

The common and rare skins (alongside the legendary Overgrown skin) do not change Ganymede’s appearance, but the epic and legendary skins do. Here we identify all the bird species that most resembles Ganymede’s look and tell a little bit about their biology.


Let’s start with the “proper” red northern cardinal, Cardinalis cardinalis (Fig. 5A). This species belongs to the family Cardinalidae and is also commonly known as redbird, being easy to identify due to its color, black “mask” and crest. Ganymede appears as a male cardinal (females are light brown). These birds eat mainly seeds, grains and fruits, but feed their young with insects. They are found from Belize and Guatemala, through Mexico and eastern USA, all the way to Canada. The species was introduced by humans in other American states, like California and Hawaii. Cardinals are common in residential areas and visit bird feeders. They were prized as pets due to their bright plumage and song, but thankfully now have full legal protection.

Next, we have Ganymede appearing as a blue jay, Cyanocitta cristata (Linnaeus, 1758) (Fig. 5B). This species belongs to the family Corvidae (ravens, crows, jays and magpies) and has a distinct color pattern. As a matter of fact, the color pattern of Ganymede’s wings is a little bit simplified when compared to the actual bird’s complicated gradation of colors. Blue jays can be found in central and eastern USA and Canada; they eat nuts, grains and small invertebrates. These birds are typically monogamous, pairing for life; genders are similar in plumage and size. Blue jays are very intelligent, with complex social systems.


Figure 5. Bastion’s skins, accompanied by a close-up of Ganymede and a photo of the actual bird species in which he was based. Bastion’s skins are screenshots from the game; the images were extracted from Overwatch Wiki. A. Bastion’s Omnic Crisis skin. Northern cardinal, Cardinalis cardinalis (photo by Stephen Wolfe, 2011; image extracted and modified from Wikimedia Commons). B. Bastion’s Defense Matrix skin. Blue jay, Cyanocitta cristata (photo by Mdf, 2005; image extracted and modified from Wikimedia Commons).

Bastion’s two “wooden” skins are fittingly accompanied by a Ganymede looking like two species of woodpeckers (family Picidae): the red-naped sapsucker, Sphyrapicus nuchalis Baird, 1858 (Fig. 6A; although it is also reminiscent of the pileated woodpecker, Dryocopus pileatus (Linnaeus, 1758), and the downy woodpecker, Dryobates pubescens (Linnaeus, 1766)) and the Arizona woodpecker, Leuconotopicus arizonae (Hargitt, 1886) (Fig. 6B). The sapsucker, as its name implies, drills hole in trees to feed on the plant’s sap, also eating insects that are attracted to the sap. These birds can be found throughout the Great Basin region and the Rocky Mountains, in North America. The Arizona woodpecker has a more restricted range, occurring in the southern parts of Arizona (obviously) and New Mexico, USA, and in western Mexico. This species feed mainly on insects (especially beetle larvae), but may also eat fruits and acorns. Similar to the case of the blue jay above, the color pattern on Ganymede’s head, chest and wings are very simplified in relation to the real animals. Also, there is some divergence in color: while the male Arizona woodpecker has a red crest, Ganymede has a yellow one, which makes him more similar to female woodpeckers.


Figure 6. Bastion’s skins, accompanied by a close-up of Ganymede and a photo of the actual bird species in which he was based. Bastion’s skins are screenshots from the game; the images were extracted from Overwatch Wiki. A. Bastion’s Antique skin. Red-naped sapsucker, Sphyrapicus nuchalis (photo by Glenn Bartley, 2011; extracted from Glenn Bartley Nature Photography, used with permission). B. Bastion’s Woodbot skin. Female (left) and male (right) Arizona woodpecker, Leuconotopicus arizonae (photos respectively by Alan Wilson, 2007, and Nature’s Pics Online, 2007; images extracted and modified from Wikimedia Commons).

The last two of Bastion’s skins are based on steampunk designs. Therefore, they needed a more city-dwelling bird to accompany him. Ganymede thus appears as a rock pigeon, Columba livia Gmelin, 1789 (family Columbidae), the common pigeon we have in large cities. The Gearbot skin has a common rock pigeon (Fig. 7A), while the Steambot skin is accompanied by an albinistic pigeon (Fig. 7B). We judge it is an albinistic (instead of a leucistic; see Box 1 below) bird, because the beak also does not have the usual black pigmentation (it is pinkish yellow). We could not check if the same is true for Ganymede’s legs, though, as we have yet to unlock this very expensive skin in the game.


Figure 7. Bastion’s skins, accompanied by a close-up of Ganymede and a photo of the actual bird species in which he was based. Bastion’s skins are screenshots from the game; the images were extracted from Overwatch Wiki. A. Bastion’s Gearbot skin. Rock pigeon, Columba livia (photo by Diego Delso, 2012; image extracted and modified from Wikimedia Commons). B. Bastion’s Steambot skin. Albinistic rock pigeon, Columba livia (photo by Maria Corcacas; image extracted from Project FeederWatch, a partner organization of the Cornell Lab of Ornithology and Bird Studies Canada, used with permission).

Unsurprisingly, all the birds above are American (Blizzard’s headquarters is in California). As explained above, the depictions are not completely true-to-life, but simplified in some instances. This is to be expected, we guess, since the game’s developers would not need focusing too much on a scientifically accurate depiction of a bird. They would rather be more worried about making all the shooting fun. Nevertheless, it seems the team at Blizzard clearly put a lot of effort in making Ganymede, as not only his appearance but also his movements in the game are all very realistic (the model for Ganymede in the animated short The Last Bastion was done based on the pet parrot of a Blizzard employee). The two pigeon “skins” for Ganymede even change his body shape to make him look like a pigeon.

Box 1. Albinism and leucism

Both albinism and leucism are genetic variations, meaning they are conditions defined by the genes the animal inherits from its parents. Albino animals show a complete (or partial) absence of the pigment called melanin in their skin, hair, feathers, scales, cuticles and irises. Melanin is responsible for brown and black colors. Thus, albinos are very light-skinned, with white hairs and red eyes (the lack of pigment in the eyes means that the light is reflected by the blood vessels). This failure to produce melanin is usually caused by the absence or malformation of an enzyme involved in its production. Common albino animals include white lab rats and mice and rabbits. People with albinism are also rather common.

In leucism, however, there is only partial loss of pigmentation. This means paler hairs (or feathers, etc.), often “creamy” in color, but with no changes to the eyes. It is also different from albinism in another regard: leucism is a reduction in several types of pigment, not only melanin. Leucistic peacocks are very commonly bred in captivity and leucistic lions are a fan-favorite in zoos.

On the opposite side of albinism, there is a condition called melanism. The over-deposition of the black pigment melanin in hairs (or feathers, etc.) results in very dark animals, like the black jaguar.


Nevertheless, despite all the care in making Ganymede, there are some major inconsistencies (besides the whole “American-bird-in-German-forest” issue discussed above). Until Gamescom (in August 2016, when the animated short The Last Bastion was premiered), we supposed that Ganymede was a male. This was based on: (1) the name, which is a male one (originally from Greek mythology); (2) it is crested and colorful, which is common of male birds, while females often have a plainer look; and (3) it sings a lot, which is also a typical male activity in birds (usually used for defending territory or courtship).However, in the aforementioned animated short, Ganymede is building a nest, which is typical female behavior. It is very rare for male birds to do the nest building (this is only seen, for instance, in species of weavers and megapodes). Moreover, Bastion’s Overgrown skin, which relates to the short, has a nest with eggs place on the robot’s shoulder (Fig. 8). Needless to say, only females can lay eggs. Moreover, the incubation and hatching is usually also done by females; male birds only rarely incubate eggs. Of course, the eggs from the Overgrown skin are way too large (Fig. 8) to belong to Ganymede anyway.


Figure 8. Bastion’s Overgrown skin (screenshot from the game). Image extracted from Overwatch Wiki.

Ganymede’s sex is never directly alluded to in the game or official material, although sometimes we could find the pronouns “he” and “his” referring to it on Blizzard’s websites. Curiously, the same is true for Bastion, who is almost always referred to by the pronoun “it”, but sometimes by “he”.


The player can also customize Bastion’s victory pose, which is shown after the match if he/she was part of the winning team. One of Bastion’s poses is called Birdwatching, because, well, he is watching his bird (Fig. 9).

It might sound surprising to some that birdwatching is not only an actual pastime but a very popular one at that. But what exactly is it?


Figure 9. Bastion’s Birdwatching victory pose (screenshot from the game). Image extracted from Overwatch Wiki.

Birdwatching, also called birding, is basically an activity of wildlife observation, where you go out to observe, of course, birds. You can do this, of course, with the naked eye, but it is better done with a good pair of binoculars (or sometimes a telescope). It’s a hobby that actually attracts a huge lot of people (Fig. 10), especially when a rare bird is involved. There are, of course, guides for beginners explaining everything about how to start birding, like Birding for Beginners: A Comprehensive Introduction to the Art of Birdwatching (by S. Buff, 2010), and websites like All About Birds (by the Cornell Lab of Ornithology).


Figure 10. Birdwatchers (often called simply “birders”) at Caerlaverock, UK, watching a rare (in Europe) White-tailed Lapwing, Vanellus leucurus (Lichtenstein, 1823). Photo by MPF (2007); Image extracted and modified from Wikimedia Commons.

After you’ve started your birding campaigns, you will want to know the names of the birds you’re seeing. To identify the bird species, you can use one of the several field guides and handbooks in existence. These books have drawings and/or photos of the birds, with guidelines to identify them. These guides are usually restricted to a single country (or sometimes just part of it, if the country is too large, like Brazil or the USA) or continent (like Europe). It’s very easy to find one at your bookstore or online store, since they are often called “Birds of Somewhere” (Fig. 11). Of course, there are now also websites that act as these guides, such as the RSPB’s Bird Identifier (see References below).


Figure 11. Examples of “bird books”, with the covers of Birds of Australia (by K. Simpson & N. Day, 2010, 8th ed.), Birds of Venezuela (by S.L. Hilty, 2003, 2nd ed.) and Collins Bird Guide (by L. Svensson et al., 2010, 2nd ed.).

Moreover, birdwatching actually involves a lot of hearing, because you will most often hear the bird before seeing it (if you see it at all). Thus, it is also good to know what each species’ vocalization sounds likes. There are several websites to identify birds’ calls, such as the Smithsonian’s Guide (see References below). Of course, both for image and song identification, there are now lots of apps, such as eBird Mobile, BirdsEye, Collins Field Guide and Bird Song Id, among several others. Unfortunately, these websites and apps are still largely restricted to the USA and Europe, while the greatest (and some would say most splendorous) bird diversity is found in Australasia and tropical America.

Birdwatching is all about enjoying nature and having fun, but birders worldwide abide by a “code of conduct” of sorts (see, for example, the code of the American Birding Association). Nowadays, our more ecological-prone society is concerned about the impact that our activities have on the animals and their environment. Thus, birdwatching etiquette usually includes promoting the welfare of birds and their habitats, limiting the birders’ impact (photographing, using playback devices, keeping your distance from nests etc.) and thus mitigating the stress caused to the animals. Basically, have fun, but let the birds live their life – that’s what Bastion does anyway.


American Birding Association. (2016) ABA Code of Ethics. Principles of Birding Ethics. Available from: http://www.aba.org/about/ethics.html (Date of access: 23/Jul/2016).

BirdLife International. (2012) The IUCN Red List of Threatened Species. Available from: http:// www.iucnredlist.org/ (Date of access: 30/Jun/2016).

Cornell Lab of Ornithology, The. (2016a) All About Birds. Available from: https://www.allabout birds.org/ (Date of access: 30/Jun/2016).

Cornell Lab of Ornithology, The. (2016b) Project FeederWatch. Available from: http://feeder watch.org/ (Date of access: 30/Jun/2016).

Gamepedia. (2016) Overwatch. Available from: http://overwatch.gamepedia.com/Overwatch_Wiki (Date of access: 30/Aug/2016).

Glenn Bartley Nature Photography. (2016) Bird Photography from Canada and around the World. Available from: www.glennbartley.com (Date of access: 30/Jun/2016).

McCormac, J. (2016) Ohio Birds and Biodiversity. Available from: http://jimmccormac.blogspot. nl/ (Date of access: 30/Jun/2016).

Mcgraw, K.J.; Hill, G.E.; Parker, R.S. (2003) Carotenoid pigments in a mutant cardinal:  implications for the genetic and enzymatic control mechanisms of carotenoid metabolism in birds. The Condor 105: 587–592.

Overwatch. (2016) Bastion Reference Kit. Available from: https://playoverwatch.com (Date of access: 30/Jun/2016).

Overwatch Wiki. (2016) Overwatch Wiki. Available from: https://blzgdapipro-a.akamaihd.net/med ia/reference/bastion_reference.pdf (Date of access: 26/Jun/2016).

RSBP. (2016) Bird Identifier. Available from: http://www.rspb.org.uk/discoverandenjoynature/discoverandlearn/birdidentifier/ (Date of access: 23/Jul/2016).

Smithsonian’s National Zoo. (2016) Guide to North American Bird Songs and Sounds. Available from: http://nationalzoo.si.edu/scbi/migratory birds/education/nasongkey.pl (Date of access: 23/Jul/2016).


We are very grateful to Anne Marie Johnson (Project FeederWatch) and Glenn Bartley for granting us permission to use their photos here.


Barbara Tomotani is a birdwatcher bird-scientist and was a marked presence at Blizzard’s store booth during Gamescom, hoping to find a Ganymede plush.

Rodrigo Salvador is a biologist, but now is found mostly escorting payloads as either D.Va or Lúcio. He has his fair share of Plays of the Game as Bastion, though.

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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).

Splatoon - Box 2

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).

Splatoon - Fig 16

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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Bush, S.L. & Robison, B.H. (2007) Ink utilization by mesopelagic squid. Marine Biology 152: 485–494.

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

Clover, C. (2004) The End of the Line: How Overfishing Is Changing the World and What We Eat. Ebury Press, London.

Derby, C.D. (2007) Escape by inking and secreting: marine molluscs avoid predators through a rich array of chemicals and mechanisms. The Biological Bulletin 213: 274–289.

Bello, G. & Piscitelli, G. (2000) Effect of sex on tentacular club development and relationships with feeding efficiency and growth in Sepia orbignyana (Cephalopoda, Sepiidae). Ophelia 53: 113–118.

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Finn, J.K.; Tregenza, T.; Norman, M.D. (2009) Defensive tool use in a coconut-carrying octopus. Current Biology 19(23): 1069–1070.

Guerra Á., Gonzáles Á.F., Pascual S., Gawe E.G. (2011) The giant squid Architeuthis: an emblematic invertebrate that can represent concern for the conservation of marine biodiversity. Biological Conservation 144(7): 1989–1997.

Hanlon, R.T. & Messenger, J.B. (1996) Cephalopod Behaviour. Cambridge University Press, Cambridge.

Hanlon, R. (2007) Cephalopod dynamic camouflage. Current Biology 17(11): 400–404.

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Mather, J.A. & Mather, D.L. (2006) Apparent movement in a visual display: the ‘passing cloud’ of Octopus cyanea (Mollusca: Cephalopoda). Journal of Zoology 263: 89–94.

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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.

Check other articles from this volume


The birds of James Bond

Rodrigo B. Salvador1 & Barbara M. Tomotani2

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

2 Netherlands Institute of Ecology; Wageningen, The Netherlands. Rijksuniversiteit Groningen; Groningen, The Netherlands. Email: babi.mt (at) gmail (dot) com

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“The name is Bond, James Bond.”

This particular British Secret Service agent is known worldwide through numerous books, comics, videogames and, of course, films. James Bond was created by Ian Fleming and the series now outlives its creator, continuing to grow on a somewhat constant rate. Fleming’s superspy character was based on many people he met during the time he spent serving in the British Naval Intelligence Division during World War II. In his own words, James Bond “was a compound of all the secret agents and commando types I met during the war”.

But what few know is where the name comes from. Actually, it was not invented by Fleming for the character; instead, it was borrowed from a real person. So who was the original James Bond and how Fleming came to know him and to borrow his name?


James Bond was born in Philadelphia on 4 January 1900. After his mother’s death during his teens, in 1914, he moved with his father to England, going to Cambridge University and receiving his degree in 1922. Back in Philadelphia, after less than three years working for a banking firm, his love of natural history led him to join an expedition of the ANSP (Academy of Natural Sciences of Philadelphia) to the lower Amazon River in Brazil. His father, Francis E. Bond, who led an ANSP expedition (when James was 11) to the Orinoco Delta, perhaps influenced James’ decision, as well as his interest in the natural sciences.

James Bond, in 1974. Photo taken at the ANSP by Jerry Freilich. (Source: Wikimedia Commons.)

After the expedition to the Amazon, James Bond became a true ornithologist (see Box 1 for a glossary) and curator of the ANSP and started to publish many scientific papers on the South American birds. Nevertheless, he soon decided that the focus of his studies would be the Caribbean birds and this became his life’s work. He spent the next decades travelling through the Caribbean islands and studying their avifauna. The main result of his work in the region was the book “Birds of the West Indies” (1936), containing a scientific account (with descriptions, habits, geographic distribution etc.) of all the known species from the islands. The book was renamed “Field Guide of Birds of the West Indies” on its second edition (1947), but reverted to the original name on the third edition (1961). Also, from the third edition onwards, the book featured color plates of the birds (by Don R. Eckelberry) and more simplified descriptions. This made the book more similar to modern field guides, making it a must for scientists and birdwatchers alike. After the final edition (1985), Bond kept the book updated via a series of 27 supplements. He finished revising a sixth edition shortly before his death (on 14 February 1989, after a years-long fight with cancer).


Cover of the first edition of “Birds of the West Indies”, featuring the Jamaican tody (Todus todus).

From all the islands that James Bond visited, perhaps the one that most fascinated him was Jamaica, where he realized that the native avifauna was derived from North America, and not from South America as was previously supposed. This kind of study is part of the discipline known as Biogeography and led Bond, in 1971, to establish a biogeographic boundary between the Lesser Antilles and Tobago. This line separates two zones, the West Indies and South America, each with its own type of avifauna. This later led David Lack to propose, in 1973, the name “Bond’s Line” for this boundary.

Map of the Caribbean Islands, showing the West Indies avifaunal region, encompassed by Bond’s Line. (Source: Bond, 1993.)

Besides the books, Bond published more than 100 scientific papers and was awarded many medals and honors throughout his career. He is known today as the father or Caribbean ornithology. What he did not expected though, was the other Bond, which appeared in Jamaica of all places, and caused him a certain deal of consternation.


It was only in 1960–1961 that Bond discovered his fictional namesake from Ian Fleming’s novels, after several novels had already been published (the first one, “Cassino Royale”, dates from 1953). This led his wife Mary to write the book “How 007 Got His Name” (published in 1966). In this book, she tells how she jokingly wrote a letter to Fleming saying that he had “brazenly taken the name of a real human being for your rascal!”

Fleming was a British novelist and spent a couple of months every year in his estate (named Goldeneye) on Oracabessa Bay, on the northern coast of Jamaica. He was interested in the Jamaican wildlife and had a growing collection of book on shells, birds, fish and flora. Also, as any keen birdwatcher on the Caribbean, Fleming used the “Field Guide of Birds of the West Indies” (he had the 2nd edition, from 1947) and was thus very familiar with the name James Bond. On his reply to Mary’s letter, he explained that he “was determined that my secret agent should be as anonymous a personality as possible. (…) At this time one of my bibles was, and still is, Birds of the West Indies by James Bond, and it struck me that this name, brief, unromantic and yet very masculine, was just what I needed and so James Bond II was born.” On a later interview, Fleming explained further his choice of name: “I wanted the simplest, dullest, plainest-sounding name I could find, ‘James Bond’ was much better than something more interesting, like ‘Peregrine Carruthers’. Exotic things would happen to and around him, but he would be a neutral figure – an anonymous, blunt instrument wielded by a government department.”

The Goldeneye estate, as of 2011. (Source: Wikimedia Commons.)

On that letter to Mary, Fleming added that in return for using the name he could offer “your James Bond unlimited use of the name Ian Fleming for any purpose he may think fit. Perhaps one day he will discover some particularly horrible species of bird which he would like to christen in an insulting fashion.” This never happened though. Finally, Fleming also invited the Bonds to visit him in Jamaica. This happened in 1964, when the Bonds were there researching and paid a surprise visit to Fleming. This was shortly before the novelist’s death six months later, and luckily, this one-time meeting was captured in video for a future documentary. At first, Fleming was suspicious of Bond’s identity and asked him to identify some birds. Bond, of course, passed the test with flying colors and Fleming had the happiest day of the rest of his life.


Jamaica, despite being a rather small country, has a very diverse avifauna. There are circa 320 bird species living in Jamaica, including migrants. From these, 28 are endemic species, 12 are endangered and 14 are introduced. Some of these species have fascinated James Bond, Ian Fleming and countless other tourists and birdwatchers. Moreover, since Ian Fleming was such a keen birdwatcher, birds sometimes featured in his stories (and later in the films), and a collection of bird trivia can be found in Box 2 further below.

We will now briefly introduce some of the more interesting Jamaican birds and explore a little bit of their natural history and even folklore.

Red-Billed Streamertail (Trochilus polytmus)

The red-billed streamertail, also known as doctor bird or scissortail hummingbird, appears in Fleming’s short story “For Your Eyes Only” (1960). The first lines of the story are: “The most beautiful bird in Jamaica, and some say the most beautiful bird in the world, is the streamer-tail or doctor humming-bird.” It is very hard to crown a “most beautiful” bird, but the red-billed streamertail is indeed remarkable. The feathers on the male’s tail (the “streamers”) are longer than their actual body and make a humming sound during flight. James Bond (the ornithologist) seems to agree; well, partially, at least: his book says that the “adult male is the most spectacular West Indian hummingbird”.

This species is the most abundant and widespread bird in Jamaica and was actually selected as the country’s national bird. Frederic G. Cassidy (1962–2000), who studied the evolution of the English language in Jamaica, says that the name doctor bird comes from the way the animals spear the flowers with their beaks to feed. Still, the term “doctor” also carries a superstitious overtone (as in “witch-doctor”) and Cassidy notes that natives referred to these hummingbirds as “god birds”.

Male (top) and female (bottom) of the Red-Billed Streamertail (Trochilus polytmus). (Source: Wikimedia Commons.)
Male (top) and female (bottom) of the Red-Billed Streamertail (Trochilus polytmus). (Source: Wikimedia Commons.)


Jamaican Tody (Todus todus)

The todies belong to the order Coraciiformes, a group that also includes kingfishers, rollers and bee-eaters. The Jamaican tody was at first believed to be a species of hummingbird. Later, it received the name of robin, due to its small size and round appearance. This early folk name still survives in Jamaica as robin red-breas’, an allusion to the bird’s red colored patch below the beak and a copy of the English name of another bird. Robin redbreast is the old name of the European robin (Erithacus rubecula), a totally unrelated species.

The Jamaican tody is a tiny bird that feeds on insects and fruits, nesting in excavated burrows. James Bond was especially interested in the nesting behavior of birds and studied this topic at length. He chose the Jamaican tody as the cover of the first edition of “Birds of the West Indies” (1936). It has a very small geographic distribution and its population seems to be steadily decreasing in the last decade.

The Jamaican Tody, Todus todus. (Source: Wikimedia Commons.)

Jamaican Poorwill (Siphonorhis americana)

Also known as Jamaican pauraque, this nocturnal bird is a species of nightjar, of the family Caprimulgidae. The family name comes from the Latin caprimulgus (goatsucker) and reflects the absurd folk “lore” that these birds sucked milk from goats.

Very little is known about the Jamaican poorwill – it had been extinct long before Bond’s studies, since 1859. It was driven to extinction by introduced rats and mongooses, alongside the usual human-caused habitat destruction. Since the birds nest on the ground, their eggs are easy prey for these introduced mammals. Nevertheless, there are some recent (1998) records of caprimulgids from the regions of the Milk River and the Hellshire Hills in the country, but they remain unconfirmed. Thus, a very small population of poorwills might still exist in these remote regions. Curiously, Bond had also previously alluded to the possibility of a surviving population of these birds on the semi-arid Hellshire Hills.

The Jamaican poorwill, Siphonorhis americana. (Source: Rothschild, 1907.)

Jamaican Blackbird (Nesopsar nigerrimus)

The Jamaican blackbird (family Icteridae) is the only species in its genus and all of its names are rather misleading. Firstly, it is not an actual blackbird (Turdus merula, family Turdidae), which is a species of thrush. Nevertheless, the family Icteridae is popularly known as “New World blackbirds”, so we can let this one slip. As for the scientific name, the genus name comes from the Greek neso (island) and psar (starling) and, as one might guess, this bird is completely unrelated to true starlings (family Sturnidae). Finally, the specific epithet (see Salvador, 2014, for a crash course in species’ scientific names) means simply “very black”, which might not be so descriptive of a “blackbird” after all.

The Jamaican blackbird, Nesopsar nigerrimus. (Source: Wikimedia Commons.)

Nevertheless, a local Jamaican popular name for this bird is “wild-pine sergeant” and is more accurate than the other names. These birds feed on insects they find in tree bark or bromeliads (locally known as “wild-pines”) and are adapted to climbing trees, similar to woodpeckers. They inhabit the montane forests of Jamaica and are arranged in pairs of birds, each pair occupying a vast territory. The severe deforestation caused by mining, forestry, charcoal production and agriculture has led to an extreme habitat loss incompatible with the blackbirds’ large territories. The species is thus considered endangered, but only some very shy efforts have been made towards its preservation.

Sad Flycatcher (Myiarchus barbirostris)

The sad flycatcher (together with the lesser Antillean pewee, Contopus latirostris) is commonly called little Tom-fool by the Jamaican people, for its habit of refusing to fly away when threatened. This flycatcher species inhabits the forests of Jamaica and, as their name imply, feed on insects. In fact, the genus name comes from the Greek muia (fly) and archos (ruler), while the specific epithet refers to the presence of rictal bristles. These bristles are modified feathers (that look like mammals’ whiskers) projecting from the beak; they not only provide tactile feedback (as whiskers do), but also supposedly protect the birds’ eyes as they consumes their wriggly insect prey.

The sad flycatcher, Myiarchus barbirostris. (Source: Wikimedia Commons.)

To avoid confusion, we must note here that the sad flycatcher is part of the group known as “New World flycatchers” or “tyrant flycatchers” (the family Tyrannidae). The “Old World flycatchers” belong to another family, Muscicapidae, which is only distantly related to the Tyrannidae.

Jamaican Crow (Corvus jamaicensis)

This bird is locally known as “jabbering crow” of “gabbling crow”, for it can produce a variety of jabbering sounds (besides the common “caw” of crows). Their incessant jabbering may also sound like indistinct human languages and, to the British, rather like Welsh people, which led to the birds being nicknamed “Welshmen” in a typical bout of Brit humor.

The Jamaican crows live mainly in the country’s uplands, but may come down to the lowlands during the dry season. They feed mainly on fruit and invertebrates, but may occasionally eat other birds’ eggs and nestlings.

The Jamaican crow, Corvus jamaicensis. (Source: Internet Bird Collection, IBC155934. Courtesy of Ken Simonite.)



Bond’s work with the Caribbean avifauna set the basis for ornithology in the region and most of his insights have been continuously proved accurate. As such, his influence in science shall remain relevant for a long time to come. Well, at least until humans have extinguished all the bird species in the region – unfortunately, birds live only once and Jamaica has already lost three of its endemic species. Meanwhile, the other Bond also remain a relevant figure in popular culture and imagination, with his over-the-top stories, exotic locations, strange villains, Bond girls, fancy suits, weaponized cars and a number of crazy gadgets. James Bond has thus the (somewhat dubious) honor of having his name twice immortalized in History, as a brilliant ornithologist and as a womanizing superspy. (We believe the latter is better remembered than the former though.)

But for those of you thinking that a birder’s life is much duller than a spy’s life, some words from the naturalist and writer Alexander F. Skutch (1904–2004) might change your mind or at the very least make you revisit your beliefs: “our quest of them [birds] takes us to the fairest places; to find them and uncover some of their well-guarded secrets we exert ourselves greatly and live intensely.”


Avibase. (2015) Bird Checklists of the World. Jamaica. Available from: http://avibase.bsc-eoc.org/checklist.jsp?region=jm&list=clements  (Date of access: 02/Apr/2015).

Bond, J. (1993) A Field Guide to the Birds of the West Indies. Fifth edition (Peterson Field Guides). Houghton Mifflin Harcourt, Boston.

Bond, M.F.W.P. (1966) How 007 Got His Name. Collins, London.

Cassidy, F.G. (2006) Jamaica Talk: Three Hundred Years of the English Language in Jamaica. University of the West Indies Press, Kingston.

Clements, J.F.; Schulenberg, T.S.; Iliff, M.J.; Roberson, D.; Fredericks, T.A.; Sullivan, B.L.; Wood, C.L. (2014) The eBird/Clements checklist of birds of the world. Version 6.9. Available from: http://www.birds.cornell.edu/clementschecklist/download/ (Date of access: 02/Apr/2015).

Chancellor, H. (2005) James Bond: The Man and His World. John Murray, London.

Cruz, A. (1978) Adaptive evolution in the Jamaican Blackbird Nesopsar nigerrimus. Ornis Scandinavia 9(2): 130–137.

IUCN (International Union for Conservation of Nature). (2014) The IUCN Red List of Threatened Species. International Union for Conservation of Nature and Natural Resources. Available from: http://www.iucnredlist.org/ (Date of access: 03/Apr/2015).

Lederer, R. & Burr, C. (2014) Latin for Bird Lovers. Timber Press, New York.

MI6-HQ. (2015) MI6 – The Home of James Bond 007. Available from: http://www.mi6-hq.com/ (Date of access: 02/Apr/2015).

Parker, M. (2015) Goldeneye. Where Bond Was Born: Ian Fleming’s Jamaica. Pegasus Publications, Winnipeg.

Parkes, K. (1989) In Memoriam: James Bond. The Auk 106(4): 718–720.

Rothschild, W. (1907) Extinct Birds. An attempt to unite in one volume a short account of those birds which have become extinct in historical times – that is, within the last six or seven hundred years. To which are added a few which still exist, but are on the verge of extinction. Hutchinson & Co., London.

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

Skutch, A.F. (1977) A Bird Watcher’s Adventures in Tropical America. University of Texas Press, Austin.

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Geeky nature

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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Everybody knows that each species on the planet eventually receives a so-called “scientific name”, a two-piece Latin-like name that serves the purpose of scaring people away from science – even more than they already naturally are. So what good do scientific names do?

Cyanocitta cristata, the blue jay. Image taken from: Wikimedia Commons.

Well, for starters, having an official name assures that every single scientist in the world will refer to a species by its scientific name. This makes it a lot easier to find information about a given species in the vast scientific literature. Just imagine how easier it is to simply search the literature for information on Cyanocitta cristata instead of looking for citations of its popular names: blue jay (in English), arrendajo azul or urraca azul (in Spanish), Blauhäher (in German), geai bleu (in French), ghiandaia azzurra americana (in Italian), gaio azul (in Portuguese) etc.

Species in the genus Panthera are all closely related to each other and, thus, all have similar characteristics. Top row, from left to right: tiger (P. tigris), leopard (P. pardus) and a reconstruction of the fossil Longdan tiger (P. zdanskyi). Bottom row, from left to right: jaguar (P. onca), lion (P. leo) and snow leopard (P. uncia). Image taken from: Wikimedia Commons.

Moreover, by stating that a tiger (Panthera tigris) belongs in the genus Panthera, we are saying that it is more closely related to the other species in the same genus (such as the lion, Panthera leo, and the jaguar, Panthera onca) than to any other member of the cat family (called Felidae), such as the Canadian lynx (Lynx canadensis) or the saber-toothed cat (Smilodon fatalis). These statements are the basis for organizing the tree of life.

Now, let us take a moment to review how scientific names work. They have two parts. The first one is the name of the genus, like Panthera in the example above. The second part is called the “specific epithet”, like tigris for the tiger. Now mind you that the species name is not simply tigris. The word tigris means nothing by itself, unless accompanied by the genus name. As such, the complete name of the tiger species is Panthera tigris.

The specific epithet (the cristata of the blue jay example) is usually not a random word. It may help describing a species, giving an idea of what it is like or where it comes from. Let’s take a look now at some useful specific epithets:

  • Take the snail species called Eoborus rotundus, for instance. The specific epithet implies that this particular snail is rotund or round and this is something that makes it different from other species in the same genus. For instance, the species Eoborus fusiformis is, like the name implies, spindle-shaped. As such, the specific epithet serves to point out a feature that makes the species easy to distinguish (diagnose, in the jargon) from other closely related species.

  • The specific epithet can also reflect the place where the species lives or, at least, where it was first found. For instance, we expect to find a bird named Tangara brasiliensis in Brazil and a slug called Arion lusitanicus in Portugal. Sometimes this fails though: the bird Tangara mexicana is not found in Mexico – perhaps a lack of geographical knowledge of the person who named it.

  • An epithet may also reflect the kind of habitat where the species lives in or its mode of life. The snail Cepaea hortensis received this epithet because it is commonly found in groves and orchards.

The round Eoborus rotundus (left) and the spindle-shaped Eoborus fusiformis (right) are fossil land snails species from the Paleocene/Eocene of Brazil.

Also, there are the not-so-useful names, the ones that are given in honor of someone, commonly a great scientist who usually worked with that group of animals before. For instance, there are loads of species, such as the snail Bulimulus darwini, named after Charles Darwin. Of course, Darwin deserves all the honors possible, but sometimes this habit of naming can become more a matter of ass-kissing than anything else. It is thus common (and useless) to name species after the person who funded the research or even after people who are completely irrelevant to science, such as the zoologist’s wife or children. Therefore, we have lots of women’s proper names, especially in the butterflies. Even worse, almost all birds of paradise are named after European nobility or royalty. It might be cute, be it is useless.

Sometimes, a species is named after a mythological being. This is often also useless, despite being way more awesome, like the owl genus named Athene. Yet, it might also be useful sometimes. For instance, the snail Brasilennea arethusae was named after the nymph Arethusa. This snail was the first fossil land snail found in Brazil and naming it after a forest-dwelling nymph made this very clear (at least to people who know their mythology), in a manner similar to the example of Cepaea hortensis above. Another example is Pseudotorinia phorcysi, a snail that lives in the deep sea, named (by myself and two colleagues) after the Greek deity Phorcys, the god of the hidden dangers of the deep sea.

Halystina umberlee. The photo on the left was taken on a light stereomicroscope. The one on the right was taken using a scanning electron microscope, which reveals much more details about the structures of tiny creatures.

And now, finally, I arrived where I wanted: the geek names. Some species have received names coming from geek culture. As the first example, there is Halystina umberlee. This is also a deep-sea snail named by myself and the same two colleagues, but this time, instead of the Greek god Phorcys of the example above, we used the goddess Umberlee. She is also a goddess of the dangers of the deep sea, but she is a fictitious deity, coming from the so-called Faerûnian pantheon of the Dungeons & Dragons RPG. To my knowledge, I was the first geek to name a species after something D&D-ish. But I’m far from being the first geek in the history of zoological nomenclature.

The goddess Umberlee rising from the waves (taken from the book Faiths & Pantheons by Eric L. Boyd & Erik Mona, 2002, published by Wizards of the Coast).

Back in the 19th century, geek zoologists did not have Tolkien or Star Trek yet, so they named their species after the geeky literature of their time. For instance, the jumping spider Bagheera kiplingi – the genus named after the character and the specific epithet after the writer.

From the middle of the 20th century onwards, geekness became much more pervasive. Just to exemplify, we have the spiders Pimoa cthulhu and Aname aragog, the fossil plant Phoenicopsis rincewindii, the mussel Ladella spocki, the fish Bidenichthys beeblebroxi, the dinosaur Dracorex hogwartsia and a whole lot from the Tolkienverse: the weevil Macrostyphlus gandalf, the fossil mammals Protoselene bombadili and Mimatuta morgoth, the leafhopper Macropsis sauroni etc.

The dinosaur Dracorex hogwartsia, from the late Cretaceous of North America. Its skull really looks like that of a “typical” dragon, but the animal was disappointingly an herbivore. Image taken from: Wikimedia Commons.

Genera (this is the plural of genus!) have also been named after geek culture: the worm Yoda, the slug Smeagol (which has its own precious family, Smeagolidae), the crustacean Godzillius, the snail Cortana (this one is also my fault), the lizard Smaug, the fish Batman (why not an outright bat is something that also baffles me) and the tardigrade (microscopic creatures also known as sea-bears) Beorn, among many others.

One species that deserves a full paragraph here is Han solo. Yes, exactly, I’m talking about the Chinese trilobite. In the official description (from 2005), the author Samuel T. Turvey says that the name comes from to the Han Chinese (by far the most numerous ethnic group in China today) and that the specific epithet solo is because the species is the youngest fossil in the family (meaning the last or sole survivor). Still, Turvey later said that it was all a bet; some friends dared him to name a species after a Star Wars character. But Turvey was rather cowardly in this. He could have stated up front (and proudly) where the name came from. There is no rule in the International Code of Zoological Nomenclature (the code that regulates the names) against this. I have done it myself and lots of geeks before me have been doing it for a long time. The official description of the fossil turtle genus Ninjemys reads: “Ninja, in allusion to that totally rad, fearsome foursome epitomizing shelled success; emys, turtle.” And no editor or reviewer can prevent the name being given. Well, perhaps they could back in 1900-something, where everybody was worried with proper-this and proper-that, but, come on, not in 2005! Dr. Turvey, you have made geekdom both proud and disappointed at the same time. Please get things right from the start next time.

Skull of the fossil teenager ninja turtle Ninjemys oweni, from the Pleistocene of Australia. Those are some pretty badass spikes and it actually looks a little bit like Slasher. Image taken from: Wikimedia Commons.

OK, I grant you that geek names are not very useful, but they sure give a little color to zoological (and sometimes also botanical) nomenclature. Taxonomy (the science of naming and classifying living creatures) is very nice and all, but the scientific papers in the area can be very arid and lifeless. Therefore, I think that it is a very valid endeavor to try to have some fun while doing taxonomy, especially if you are a geek and have a whole pantheon of heroes, gods and monsters to get your inspiration from.


I am very grateful to Ed Greenwood, creator of the Forgotten Realms (and, thus, of Umberlee) for his very kind comments on the new species named in honor of the goddess. Also, many thanks to my co-authors of scientific papers for allowing my geekness to run free when naming species.


If you want to know exactly how species are formally described and get their official names, this is the best guide out there: Winston, J.E. (1999) Describing Species: Practical Taxonomic Procedure for Biologists. Columbia University Press, New York.

A less academic approach to the whole naming process can be found in: Wright, J. (2014) The Naming of the Shrew: A Curious History of Latin Names. Bloomsbury Publishing, London.

For a more philosophical view and musings about the importance of naming species for scientists and non-scientists alike, try this one (you might want to skip chapter 9 though, which is far too exaggerated on its glorification of molecular taxonomy): Yoon, C.K. (2010) Naming Nature: The Clash Between Instinct and Science. W.W. Norton & Company, New York.

If you want a taste of what a real taxonomic paper looks like, try this one (where Halystina umberlee came from): Salvador, R.B.; Cavallari, D.C.; Simone, L.R.L. (2014) Seguenziidae (Gastropoda: Vetigastropoda) from SE Brazil collected by the Marion Dufresne (MD55) expedition. Zootaxa 3878(6): 536–550.

For the ones who like rules and want to take a look at the “laws” presiding over animal names, the International Code of Zoological Nomenclature (ICZN, for the intimate) is the one and only guide: http://iczn.org/iczn/index.jsp.

Last but not least, Mark Isaak has compiled a lot of geeky scientific names on his website: www.curioustaxonomy.net/etym/fiction.html. I must confess that I did not know most of them, since they are insect names (rather removed from my area of study). In any case, it is always good to know that I am not alone – there are many other geek zoologists and paleontologists out there. Just take a look at the sheer amount of Lord of the Rings and Silmarillion names; it’s amazing!

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