Valleys Between: bringing environmental issues to games

Niamh Fitzgerald

Little Lost Fox. Wellington, New Zealand.

Email: niamh (at) littlelostfox (dot) com

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Valleys Between[1] is an environmental puzzle game, where your goal is to grow your world for as long as you can while protecting it from threats that will damage its health.

When we started designing Valleys Between we wanted to explore ways to get people thinking about environmental issues, and while the game has evolved during the game development cycle, the core themes of the game are still there. While we considered real world ecology and nature, we realised early on that to create a fun and engaging game we would need to take inspiration from them without being too literal.

One of our goals is to create a strong bond between the player and the world they’ve created, and one of the ways we do this is by allowing you to literally shape the world with your fingertips. Players only have the ability to swipe up or down to interact with the world, but small actions such as pulling a tree up out of the ground can actually have a big impact. Much like the real world, one action isn’t always enough to solve larger problems but a group of small actions can result in a big change.

The beautiful hexagonal environments of Valleys Between.

Many of the games mechanics are inspired by nature, though in a simplified or abstract way. This allows us to craft gameplay that’s enjoyable and relatable without ever straying too far into something that feels completely at odds with reality (at least in most cases). With that in mind we had two important rules that guided our design:

  1. The game is inspired by nature, so the environmental theme should always be present while never overpowering or distracting the player from the gameplay.
  2. We won’t sacrifice enjoyable gameplay for the sake of keeping something too realistic or similar to how our real world works.

These rules allowed us to find a balance between fun and relatable mechanics that are easy for the player to understand. When designing mechanics we often started from an ecological concept and explored how we could distill it down to base elements to see how they could work well within the game. The best way to illustrate this is to look at the primary mechanics in Valleys Between.

At its core, Valleys Between is about creating a thriving world. The first step to doing this is to create an environment where things can grow, so the first move a player makes is to create water tiles in their new world. Water makes all dirt tiles around it turn into grass, and trees can only be planted on grass. To plant a tree, the player pulls up on a grass tile and essentially plucks a fully-grown tree out of the ground. While this is clearly a few steps removed from reality, it feels close enough, and this familiarity helps create a stronger connection between the nature presented in the game and what the player expects from nature in the real world.

Trees that are next to each other can be combined to make a forest, which grows your world by adding a new row of land. In this way, the base relationship between water and trees are shown as being critical to growing a world. Groups of forests can be further combined to make a house, which introduces humans as part of the ecosystem in Valleys Between. While this is an incredibly simplified representation of nature to a few small mechanics in Valleys Between, it’s part of what makes it feel environmentally rich.

Grow a thriving world and find the balance sustain it.

The game wouldn’t be very fun without something challenging you, so we decided to introduce the two sides of human influence on the environment. The first is a positive influence of creating a house by combining trees which helps your world grow and expand. However, as your world grows, we also introduce a negative influence in the form of factories and other man-made objects. Factories threaten the health of your world and they can spill oil to surrounding tiles if you leave them for too long. While there isn’t necessarily an easy action to fix things these things in our world, we wanted players to want to protect their world from these threats even if they can’t stop them from occurring. We also found in early playtests that people became very attached to the animals that wander their world, and this helped them feel connected to it, so we decided to tie these concepts together and have animals act as the primary protectors of your world. Animals wander throughout your world, and while you can influence their path, you aren’t able to control them directly. You can choose to use them to nurture and enhance a specific area, or use them to convert a factory to something that won’t damage the health of your world. Once you’ve used an animal, they fall asleep for a period of time so the player has to choose when to nurture and when to protect their world.

While these mechanics may seem to be quite a stretch from the real world, we’ve found that by taking inspirations from nature rather than literal representations, we’ve been able to craft an enjoyable game.

Animals are the protectors of your world.


Niamh Fitzgerald is a producer and game designer at indie studio Little Lost Fox, based in Wellington, New Zealand. She organised the New Zealand Game Developer Conference in 2017 and 2018, and likes to combine her love of travel with game development by getting involved in game developer events around New Zealand and internationally.

[1] Released in 2018 by Little Lost Fox. Currently available for iPhone/iPad and coming soon to Android. Learn more at

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Cephalopods of the Multiverse

Mark A. Carnall

Oxford University Museum of Natural History. Oxford, UK.

Email: mark.carnall (at) (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: 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: 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:// -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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Douglas Adams and the world’s largest, fattest and least-able-to-fly parrot

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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The system of life on this planet is so astoundingly complex that it was a long time before man even realized that it was a system at all and that it wasn’t something that was just there.” ―Douglas Adams, 1990

Douglas Noel Adams was born on 11 March 1952 in Cambridge, UK, and grew up to become one of geekdom’s most revered icons. Adams is the author of… Well, that is pretty obvious and I should not have to write this down, but I will nonetheless, just because I won’t be able to sleep well otherwise. So bear with me for a moment – here goes: Adams is the author of the trilogy The Hitchhiker’s Guide to the Galaxy, the self-proclaimed world’s largest trilogy, with five books in total[1].

However, unbeknownst to many of his fans, Adams was also an environmental activist. He spearheaded or participated in several conservation initiatives, such as Save the Rhino International. His history with conservation started in 1985, when the World Wide Fund for Nature (better known as WWF) and British newspaper The Observer partnered up, sending writers to visit endangered species to raise public awareness (BBC, 2014). Adams travelled to Madagascar in search of a lemur species, the aye-aye (Daubentonia madagascariensis). As he put it, “My role, and one for which I was entirely qualified, was to be an extremely ignorant non-zoologist to whom everything that happened would come as a complete surprise” (LCtS: p. 1).

In Madagascar Adams met not only weird lemurs, but also British zoologist Mark Carwardine. They enjoyed the experience and decided to travel the world to see other endangered animals. I mean, Adams and Carwardine travelled the world, not the lemurs; the lemurs stayed in Madagascar as far as anyone can tell. According to Carwardine, “We put a big map of the world on a wall, Douglas stuck a pin in everywhere he fancied going, I stuck a pin in where all the endangered animals were, and we made a journey out of every place that had two pins” (BBC, 2014).

Their travels resulted in Last Chance to See, a BBC radio documentary series that aired in the end of 1989. The companion book (by Adams & Carwardine, 1990, henceforth abbreviated as “LCtS”) was published in the following year[2] (Fig. 1). As a matter of fact, Adams considered this book as his favorite work (Adams, 2005).

Figure 1. Cover art of the American edition of Last Chance to See (Harmony Books, New York, 1991).

Despite Adams’s calling himself an “ignorant non-zoologist”, world-renowned evolutionary biologist Richard Dawkins politely disagreed, writing: “Douglas was not just knowledgeable about science. He didn’t just make jokes about science. He had the mind of a scientist, he mined science deeply and brought to the surface… humour, and a style of wit that was simultaneously literary and scientific, and uniquely his own” (Dawkins, 2009: p. xiii).

Last Chance to See describes Adam’s and Carwardine’s travels around the globe to see nearly-extinct species, such as the Amazonian manatee (Trichechus inunguis) and the northern white rhinoceros (Ceratotherium simum cottoni). As one could expect, nearly all the species are mammals, since most of the public are primarily concerned with cuddly and relatable species. I, however, will focus here on the only bird on their list that got an entire chapter for itself. And I’ll do that for various reasons: (1) I am not very normal, so I am not that fond of smelly mammals; (2) it is a success story and people like success stories; and (3) this is a very funny-looking bird, I promise you.

This bird is called kakapo.


Mark Carwardine first described the kakapo to Douglas Adams as “the world’s largest, fattest and least-able-to-fly parrot” (LCtS: p. 7). His description might seem a little disparaging at first, but it was meant in an affectionate way – you cannot help but smile when you see a kakapo. Besides, Carwardine’s description is actually spot-on (Fig. 2).

According to Adams, “[the] kakapo is a bird out of time. If you look one in its large, round, greeny-brown face, it has a look of serenely innocent incomprehension that makes you want to hug it and tell it that everything will be all right” (LCtS: p. 108).

Figure 2. Sirocco, a male kakapo, looking funny as kakapos usually do. Image extracted from New Zealand Birds Online (; credit: Dylan van Winkel.

The kakapo (or kākāpō, in Māori or Te Reo spelling) is a nocturnal flightless bird and its face resemble that of an owl, with the eyes positioned more to the front. For this reason, it is also known as owl-parrot or night parrot. Kakapos have green feathers, speckled with black and yellow (Fig. 3).

Figure 3. A kakapo looking unusually serious. Image extracted from New Zealand Birds Online (; credit: Colin Miskelly (2011).

Furthermore, kakapos are solitary birds, have a polygynous lek mating system (don’t panic, I’ll explain that later), lack male parental care, and breed in irregular intervals (with gaps of 2 to 7 years; Powlesland et al., 2006). Kakapos are so unique that ornithologists classified the species in its own family: Strigopidae. They are the very first lineage to have branched out of the parrot group (the Order Psittaciformes). Even their closest “relatives”, the kaka and the kea (also from New Zealand), are already considered to be very distinct from kakapos.

Being such an ancient lineage of parrots, researchers consider that it could have split off the rest of the parrot groups when New Zealand got separated from the what is now Australia and Antarctica around 80 million years ago (Gibbs, 2016). All the southern landmasses had been previously joined in the supercontinent Gondwana, which was made up of South America, Africa, India, Antarctica, Australia and Zealandia (Fig. 4) and was by that time finishing its separation.

Figure 4. The supercontinent Gondwana during the Triassic (circa 200 million years ago). Image modified from Wikimedia Commons; credit: LennyWikidata (2008).

This break up left Zealandia with no mammals and a bird “paradise” island started to take shape. It is considered that the kakapo followed the trend of oceanic island bird lineages (where nasty mammals are not present) to evolve larger and flightless forms (Powlesland et al., 2006). For instance, that happened with the lineages of the dodo, moa, and elephant bird.


I cannot overstate how weird kakapos are for a parrot – or for a bird, actually. Adams considered the kakapo the strangest and most intriguing of all the creatures he saw during his travels with Carwardine (LCtS: p. 105). So I’ll illustrate that by highlighting some aspects of its biology that are of broader interest or peculiar weirdness. If you, however, are looking for a complete guide to the species’ biology, do take a look at the work of Powlesland et al. (2006).

We already covered that kakapos are nocturnal and flightless, and thus have good hearing and sense of smell, alongside massive legs and feet to walk around and climb trees. Yes, they do not fly, but do climb trees to feed. Evolution works in mysterious ways, it seems. Elliot (2017) wrote: “They often leap from trees and flap their wings, but at best manage a controlled plummet.” I prefer, however, the way Douglas Adams put it: “it seems that not only has the kakapo forgotten how to fly, but it has forgotten that it has forgotten how to fly. Apparently a seriously worried kakapo will sometimes run up a tree and jump out of it, whereupon it flies like a brick and lands in a graceless heap on the ground” (LCtS: p. 109)[3].

It seems kakapos are not able to follow the suggestion of the Hitchhiker’s Guide: “There is an art, it says, or rather, a knack to flying. The knack lies in learning how to throw yourself at the ground and miss. (…) Clearly, it is this second part, the missing, which presents the difficulties” (Adams, 1982). Kakapos just constantly fail to miss the ground.

Overall, kakapos are quite large birds, weighing around 2 kg, but males may weigh up to 4 kg and be 40% larger than females (Eason et al., 2006; Elliot, 2017). Their life span is unknown, but is estimated at 60 to 90 years (Department of Conservation, 2018a, 2018b).

Kakapos are vegetarian and eat almost every possible parts of plants. In fact, they only breed in years with a good abundance of fruit (Cockrem, 2006; Elliot, 2017). In their current habitat, kakapo reproduction is tied with that of the rimu (Dacrydium cupressinum), an evergreen coniferous tree of the podocarp family (Fig. 5). These plants bloom together every 2 to 4 years (sometimes it takes more); the kakapos must wait for the rimu because they depend on its “fruits” (Fig. 6) to feed the chicks (Cockrem, 2006; Ballance, 2010).

Figure 5. A rimu tree is really tall for a flightless bird to climb. Image retrieved from Wikimedia Commons; credit: Kahuroa (2008).
Figure 6. A ripe rimu “fruit”, or better put, a seed sitting on a fleshy cup. Image retrieved from Wikimedia Commons; credit: Department of Conservation (2002).

Unlike any other parrot, kakapos are lek breeders. This behavior is common for other groups of birds and even other animals, though. It consists in males gathering relatively close to each other and starting a competition to show off to females. Birds can do this mainly by song or dance (or both), but might also include somersaults and flying maneuvers. Each female will chose the best performer (in their opinion at least) and successful males typically mate with more than one female during a single season.

Male kakapos sing to attract females. Or rather, they do something akin to “Pink Floyd studio out-takes” (LCtS: p. 111). The most common type of call produced by kakapos is called booming. This is a low-frequency (<100 Hz) resonant call, which can be heard up to 5 km away (Merton et al., 1984; Higgins, 1999). To produce this sound, male kakapos fill up internal air sacs; they can inflate until they look like a fluffy watermelon (Figs. 7, 8). Adams described the sound as a heartbeat, a powerful throb you felt before actually hearing it; and this gave the title to the kakapo’s own chapter in LCtS: “Heartbeats in the Night”.

Figure 7. A male kakapo booming – and looking like a watermelon. Image extracted from New Zealand Birds Online (; credit: Department of Conservation (image ref 10027966, photo by Ralph Powlesland).
Figure 8. How to camouflage as a watermelon in four easy steps. OK, now serious caption: postures of a male kakapo booming. Figure reproduced from Merton et al. (1984: fig. 4). The original caption reads: “(1) Normal stance; (2) Alert static pose between booming sequences; (3) Commencement of booming: inflation of thorax while giving preliminary ‘grunts’; (4) Maximum thoracic inflation during loud booming.”

Booming also serves to indicate the male’s overall location to the female. Once they are close by, males can produce a sharp metallic “ching” call to enable females to pinpoint their exact location (Powlesland et al., 2006). A good place to hear kakapo booming and chinging is New Zealand Birds Online (http://nzbirdsonline.

The female nests on the ground, either on a spot covered by dense vegetation or in natural cavities (Elliot, 2017). Kakapos usually lay 2 to 4 eggs and the female raise the chicks alone (Fig. 9; Cockrem, 2006; Powlesland et al., 2006). Young birds leave the nest within 2 to 3 months, but remain close to their mother’s home range until they are 6.5 to 8.5 months old (Farrimond et al, 2006; Powlesland et al., 2006).

So how do we summarize kakapos? Adams gives us a nice idea: “The kakapo (…) pursues its own eccentricities rather industriously and modestly. If you ask anybody who has worked with kakapos to describe them, they tend to use words like ‘innocent’ and ‘solemn’, even when it’s leaping helplessly out of a tree. This I find immensely appealing” (LCtS: p. 121).

Figure 9. Alice, a female kakapo, on her nest with her two chicks (circa 45 days old). Image extracted from New Zealand Birds Online (; credit: Department of Conservation (image ref 10048384, photo by Don Merton, 2002).

Box 1. Kakapo names

Since there are so few kakapo left and the whole population is managed, each bird has its own name. When Adams and Carwardine visited Codfish Island, they met a kakapo named Ralph. Later on, Adams himself got to name a kakapo Jane, after his then-girlfriend (Balance, 2010). You can check this amazing infographic (by DeMartini et al.) with all the names and family trees of known kakapos: Kakapo/Dashboard1?:embed=y&:display_count=yes&:toolbar=no&:showVizHome=no.

Presently, the most famous kakapo is Sirocco, who became a YouTube star after he tried to mate with Carwardine’s head during the filming of the Last Chance to See TV series (Carwardine, 2010). Today, Sirocco is 21 years old and is the official “spokesbird” for conservation in New Zealand (Department of Conservation, 2018b), a title given to him by then Prime Minister John Key.


Kakapos were present in New Zealand long before humans arrived there: some subfossil bones have been dated from 2500 years ago (Wood, 2006). They were very common and lived throughout both the North and South Islands (Tipa, 2006), with few natural enemies. They were successful in their pre-human environment, but that was soon to change.

Polynesian settlers arrived in Aotearoa[4] between 1200 and 1300 CE (Wilmshurst et al., 2010) and became known as the Māori. As typical of all humans, they brought domestic/pest species with them: dogs and rats.

As many island species, kakapos were only concerned with their known immediate predators; these mostly harmless birds were thus unprepared for a wave of invaders. Kakapos have the strategy of staying perfectly still when facing danger, which works fine against predators that rely on sight. However, this had little effect against dogs, which hunt by scent. The parrots were hunted for food and ornamentation (for instance, the Māori used the feathers in cloaks; Tipa, 2006) and the population declined. Polynesian rats also played a major role, preying upon defenseless kakapo eggs and chicks.

European settlers arrived on the 19th century and, as one might expect, colonization (and new mammalian predators, such as cats and mustelids) accelerated the species’ decline. The Europeans also brought naturalists, who collected specimens for study at museums (Fig. 10). British zoologist George Robert Gray officially named the kakapo Strigops habroptilus[5] in 1845. Later naturalists (some already born in New Zealand) went further, observing live parrots in the wild and studying their natural history.

Figure 10. Museum drawer full of preserved kakapo specimens, from the collection of the Museum of New Zealand Te Papa Tongarewa. Photo by the author (©Te Papa, all rights reserved).

Already in the 1890’s, naturalists became aware that the species was heading towards extinction, so the first efforts in conservation (transferring animals to islands in Fiordland; Fig. 11) were undertaken (Hill & Hill, 1987). They failed and eventually the species fade out from the thoughts of New Zealanders, being considered extinct or nearly so (Ballance, 2010).


That lasted until the work of Williams (1956), which summarized all knowledge about the kakapo and brought it back to the spotlight. With this renewed interest, expeditions were formed to find the species in the southernmost reaches of New Zealand.

A serious take on conservation efforts started again in the 1970’s, when a population of around 200 kakapos was found on Stewart Island (Fig. 11; Powlesland et al., 2006). A new process of translocation and monitoring then began. During the 1980s and 1990s, the animals were all moved to predator-free islands: Codfish, Maud and Little Barrier (Fig. 11; Elliot, 2017). When Adams and Carwardine visited Codfish Island in 1992, there were only around 40 kakapos left (Ballance, 2010; Carwardine, 2010).

Figure 11. Map of New Zealand showing the locations mentioned on the text. Image modified from Wikimedia Commons; credit: NordNordWest (2009).

However, things started to look brighter after a review in the management of the species (Elliot et al., 2001). A strong and focused policy and full support of the government were essential during the decades since (Jansen, 2006). The kakapo population started to recover and can now be considered one of the greatest successes among global conservation programs – and a good example of how our species can, in fact, clean up after its own mess.

The last report, from June 2017, counted a total of 154 birds (Elliot, 2017), a number exceeding previous population simulations (Elliot, 2005). Recovering the kakapo from the brink of extinction was a feat, but more challenges remain. Presently, the species is considered as “critically endangered” according to the IUCN’s Red List (BirdLife International, 2016). Although this seems better, it is good to remember that this is just one step away from the “extinct in the wild” status in this classification scheme (which the kakapo held during two issues of the Red List in the mid-1990s). Presently, kakapos only survives on offshore islands and there is still lot of work to be done until we have a viable, and self-sustaining population that does not need human management.

Maybe just panic a little bit…

The kakapo is not the only endangered species in the New Zealand – everyone has heard about kiwis, at least. So what about all the other threatened species, birds and otherwise, in the country? Jansen (2006: 190) ominously wrote: “While extinction of kakapo is now less likely than 10 years ago, the future of the 600+ New Zealand species listed as acutely and chronically threatened (…) and that presently do not receive any management is by no means secure.” So yes, there is still a lot of work to be done.

But why should we care if some species go extinct? Why should we strive so much to save them? Carwardine (LCtS: p. 205) gave what Dawkins (2009) considered to be the typical explanations for business-minded humans: (1) we mess with the environment, everything go haywire, and that ultimately affects our survival, and (2) living beings have their uses as food, drugs, etc. However, Carwardine then presented his preferred explanation, one more typical of scientists and that we say to each other over coffee: we try to save them because they are cool. Or, as Carwardine put it: “There is one last reason for caring, and I believe no other is necessary. It is certainly the reason why so many people have devoted their lives to protecting the likes of rhinos, parakeets, kakapos and dolphins. And it is simply this: the world would be a poorer, darker, lonelier place without them” (LCtS: p. 206).

“Up until that point it hadn’t really clicked with man that an animal could just cease to exist. It was as if we hadn’t realised that if we kill something, it simply won’t be there anymore. Ever. As a result of the extinction of the dodo we are sadder and wiser.” ―Douglas Adams, 1990


Adams, D. (1982) Life, the Universe and Everything. Pan Books, London.

Adams, D. (2005) The Salmon of Doubt: Hitchhiking the Galaxy One Last Time. William Heinemann, London.

Adams, D. & Carwardine, M. (1990) Last Chance to See. William Heinemann, London. [Edition used here: 2009, by Arrow Books, London.]

Ballance, A. (2010) Kakapo: Rescued from the Brink of Extinction. Craig Potton, Nelson.

BBC. (2014) Background. Last Chance to See. Available from: chancetosee/sites/about/last_chance_to_see.shtml [access date: 25 Sep 2018].

BirdLife International. (2016) Strigops habroptila.  The IUCN Red List of Threatened Species 2016. Available from: IUCN.UK.2016-3.RLTS.T22685245A93065234.en [access date: 25 Sep 2018].

Carwardine, M. (2010) Foreword. In: Ballance, A. Kakapo: Rescued from the Brink of Extinction. Craig Potton, Nelson. Pp. 9–10.

Cockrem, J.F. (2006) The timing of breeding in the kakapo (Strigops habroptilus). Notornis 53(1): 153–159.

Colfer, E. (2009) And Another Thing… Penguin Books, London.

Dawkins, R. (2009) Foreword to new edition of Last Chance to See by Douglas Adams and Mark Carwardine. In: Adams, D. & Carwardine, M. Last Chance to See. Arrow Books, London. Pp. xi–xvi.

Department of Conservation (DOC). (2018a) Kākāpō. Available from: https://www.doc.govt. nz/nature/native-animals/birds/birds-a-z/kaka po/ [access date: 26 Sep 2018].

Department of Conservation (DOC). (2018b) Sirocco the kākāpō conservation superstar. Available from: sirocco [access date: 27 Sep 2018].

Eason, D.K.; Elliott, G.P.; Merton, D.V.; Jansen, P.W.; Harper, G.A.; Moorhouse, R.J. (2006) Breeding biology of kakapo (Strigops habroptilus) on offshore island sanctuaries, 1990–2002. Notornis 53(1): 27–36.

Elliott, G.P. (2006) A simulation of the future of kakapo. Notornis 53(1): 164–172.

Elliott, G.P. (2017) Kakapo. In: Miskelly, C.M. (Ed.) New Zealand Birds Online. Available from: [access date: 26 Sep 2018].

Elliott, G.P.; Jansen, P.W.; Merton, D.M. (2001) Intensive management of a critically endangered species: the kakapo. Biological Conservation 99: 121–133.

Farrimond, M.; Elliott, G.P.; Clout, M.N. (2006) Growth and fledging of kakapo. Notornis 53: 112–115.

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

Jansen, P.W. (2006) Kakapo recovery: the basis of decision-making. Notornis 53: 184–190.

Higgins, P.J. (1999) Handbook of Australian, New Zealand and Antarctic Birds. Vol. 4: Parrots to Dollarbird. Oxford University Press, Melbourne.

Hill, S. & Hill, J. (1987) Richard Henry of Resolution Island: a Biography. John McIndoe, Dunedin.

Merton, D.V.; Morris, R.D.; Atkinson, I.A.E. (1984) Lek behaviour in a parrot: the Kakapo Strigops habroptilus of New Zealand. Ibis 126: 277–283.

Powlesland, R.G.; Cockrem, J.F.; Merton, D.V. (2006) A parrot apart: the natural history of the kakapo (Strigops habroptilus) and the context of its conservation management. Notornis 53: 3–26.

Tipa, R. (2006) Kakapo in Maori lore. Notornis 53: 193–194.

Williams, G.R. (1956) The kakapo (Strigops habroptilus, Gray): a review and re-appraisal of a near-extinct species. Notornis 7: 29–56.

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.

Wood, J.R. (2006) Subfossil kakapo (Strigops habroptilus) remains from near Gibraltar Rock, Cromwell Gorge, Central Otago, New Zealand. Notornis 53: 191–193. 


I am very grateful to Colin Miskelly, Dylan van Winkel, the Department of Conservation, and the Museum of New Zealand Te Papa Tongarewa for allowing the usage of their photographs herein. 


Dr. Rodrigo Salvador is a biologist specializing in the classification and evolution of land snails. Yes, you might say, that has nothing to do with kakapos. But it so happens that the universe conspires to keep him entangled with bird work. As a scientist, he learned with Douglas Adams that knowing the right question is sometimes more important than knowing the answer.

[1] Or six, if you count And Another Thing… by Eoin Colfer (2009).

[2] Later, in 1992, a CD-ROM set was published, with photos and audio of Douglas Adams reading the book. In 2009, BBC released a TV series of Last Chance to See, in which British comedian Stephen Fry took the place of the late Adams.

[3] However, he soon changed the tone to blame flying birds instead: “There is something gripping about the idea that this creature has actually given up doing something that virtually every human being has yearned to do since the very first of us looked upwards. I think I find other birds rather irritating for the cocky ease with which they flit through the air as if it was nothing” (LCtS: p. 120).

[4] The Māori name for New Zealand.

[5] Strigops means “owl-faced”, while habroptilus means “soft feather”.

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The entomological diversity of Pokémon

Rebecca N. Kittel

Museum Wiesbaden, Hessisches Landesmuseum für Kunst und Natur, Wiesbaden, Germany.

Email: rebecca.n.kittel (at) gmail (dot) com.

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Pocket Monsters or as they are better known, Pokémon, are playable monsters which first appeared in the 1990’s as a video game in Japan, but soon expanded worldwide. They are still very successful with numerous games, a TV series, comic books, movies, toys and collectibles, additionally to the trading card game and video games. Most recently the release of Pokémon GO, an augmented reality game for smartphones, meant that Pokémon became as popular as never before. The game launched in 2016 and almost 21 million users downloaded it in the very first week in the United States alone (Dorwald et al., 2017).

The games and TV series take place in regions inhabited by humans and Pokémon. Each Pokémon lives in a specific environment (forests, caves, deserts, mountains, fields, seas, beaches, mangroves, rivers, and marshes). The humans try to catch Pokémons with Pokéballs, a device that fits even the largest Pokémon but that is still small enough to be placed into a pocket, hence the name Pocket Monster (Whitehill et al., 2016). After Pokémon have been caught, they are put to fight against each other, just like in the real world, in which humans (unfortunately) let cockerels, crickets, or dogs fight (Marrow, 1995; Jacobs, 2011; Gibson, 2005). The origin of Pokémon goes back to the role-playing game created by Satoshi Tajiri and released by Nintendo for the Game Boy (Kent, 2001). Tajiri was not only a game developer, but like many Japanese adults, grew up catching insects as a child. He wanted to design a game so that every child in Japan could play and let their critters fight, even if they lived in areas which are too densely populated to find insects in the wild. This resulted in the 151 Pokémon in the first versions of the game (“first generation”), with each version adding more Pokémon.

Today, there are 807 Pokémon (seventh generation). Almost all are based on real organisms (mostly animals, but many plants as well), while some depict mythological creatures or objects (e.g., stones, keys). Each Pokémon belongs 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, 2018). All Pokémon in the game are oviparous, which means they all lay eggs; probably because the creator was fond of insects or just for practical reasons.

Certain Pokémon also evolve; however, this kind of evolution is not the same as the biological concept of evolution. In Pokémon evolution is largely synonymous to metamorphosis, such as when a caterpillar turns into a butterfly. As this is the core concept of the game, almost all Pokémon evolve, not only the insects, but also mammals, rocks, and mythological creatures. Usually, they evolve with a complete or incomplete metamorphosis: either they just grow larger, or their look differs significantly between the adult and the young stages.

Insects are the largest group of organisms on earth (Zhang, 2011). There are more than one million described species of insects, of a total of 1.8 million known organisms (Zhang, 2011). They occupy all terrestrial environments (forests, fields, under the soil surface, and in the air) and freshwater; some are even found in the ocean. Additionally, they show a wide range of morphological and behavioral adaptations. This biodiversity is not reflected in the Pokémon world. In the present Generation VII, only 77 of the 807 Pokémon are “Bug type”: about 9.5% of all Pokémon. The aim of this work is to describe the entomological diversity of Pokémon based on taxonomic criteria of the classification of real insects.


The Pokédex was the source of primary information on Pokémon (Pokémon Website, 2018). The criteria to identify insects are either based on the type (Bug type) or morphology (resembles a real insect). Afterwards, the insect Pokémon were classified to the lowest possible taxonomic level (family, genus, or species) according to their real world counterparts. This classification of the Pokémon allowed the comparison of their biological data (such as ecological or morphological traits; Bulbapedia, 2018) with the current knowledge of real insects. The information of the biology of real insects is largely based on Borror et al. (1981).


Not all Bug types are insects; many of them represent other arthropods, like spiders, while some are from other invertebrate groups (Table 1). Also, five insect Pokémon do not belong to the Bug type (e.g., Trapinch (#328) is a Ground type; Table 2). In total, insects represent only 62 of the 807 Pokémon. In comparison, the vertebrate groups are overly well-represented by birds (61), mammals (232), reptiles (57), amphibians (23), and fishes (39) (Table 3).

Eleven insect orders are represented in the Pokémon world, namely Blattodea (with 1 Pokémon), Coleoptera (11), Diptera (3), Hemiptera (7), Hymenoptera (6), Lepidoptera (22), Mantodea (4), Neuroptera (3), Odonata (2), Orthoptera (2), Phasmatodea (1). They are listed below in systematic order.

Table 1. List of the 20 Pokémon that are Bug type, but are not insects. Mostly, they belong to other groups within the phylum Arthropoda.
Table 2. Taxonomic classification of the insect Pokémon (Arthropoda: Hexapoda: Insecta). All images are official artwork from Pokémon games (obtained from Bulbapedia, 2018). An asterisk (*) denotes Pokémon that are not Bug type.
Table 3. Comparison between the diversity of Pokémon “species” and their respective representatives in the natural world (Zhang, 2011).

Order: Odonata

Families: Libellulidae and Aeshnidae

Genera: Erythrodiplax and Anax

Yanma (#193) evolves to Yanmega (#469).

Yanma is a large, red dragonfly Pokémon. Like all dragonflies and damselflies, it lives near the water and hunts other insects for food. Yanma is territorial and prefers wooded and swampy areas. Based on its appearance, it belongs to the dragonfly family Libellulidae, and further to the genus Erythrodiplax Brauer, 1868.

Yanmega on the other hand is a large, dark green Pokémon. It is actually a different real-world species. Not only the colors are different, but also the morphology, like the appendages on the tip of the tail. Based on this, it belongs to the dragonfly family Aeshnidae, and to the genus Anax Leach, 1815. One could argue that it is based on Meganeura Martynov, 1932, a very large (wingspan up to 70 cm) but extinct dragonfly genus from the Carboniferous Period. However, the size alone should not be the indicator to classify the species, as many insectoid species are larger in the Pokémon world compared to the real world.

Order: Mantodea

Family: Mantidae

Scyther (#123) evolves to Scizor (#212, incl. Mega-Scizor).

Scyther is a bipedal, insectoid Pokémon. It is green with cream joints between its three body segments, one pair of wings and two large, white scythes as forearms. Scyther camouflages itself by its green color. Based on its appearance, it is classified as a praying mantis (or possible a mantidfly).

Scizor is also a bipedal, insectoid Pokémon. It is primarily red with grey, retractable forewings. Scizor’s arms end in large, round pincers. It appears to be based on a praying mantis, maybe with some references to flying red ants and wasp-mimicking mantidflies.

Although Scizor evolves from Scyther, they are very different and would actually be two different real-world species. Not only are the colors different, but also the morphology: the arms end in either scythes or pincers; Scyther has one pair of wings, Scizor has two.

Fomantis (#753) evolves to Lurantis (#754).

Fomantis is a plant-like and, at the same time, an insect-like Pokémon. Its main body is pink, with green hair, green tufts on the head, and green leaves as a collar. Fomantis is somewhat bipedal and is likely based on the orchid mantis Hymenopus coronatus Olivier, 1792 (Fig. 1), which is known for being able to mimic the orchid flower, along with the orchid itself.

Figure 1. Adult male of Hymenopus coronatus. Credit: Sander van der Wel (2010), Wikimedia Commons.

Lurantis is also plant- and insect-like. It is pink, white, and green. Lurantis looks and smells like a flower, to attract and then attack foes (and prey). It also disguises itself as a Bug Pokémon for self-defense. Lurantis is likely based on the orchid mantis as well as the orchid flower itself, as it is impossible to say where the flower ends and the insect starts. Orchid mantises mimic parts of a flower, by making their legs look like flower petals. Well camouflaged, they can wait for their prey, which will visit the flower for nectar.

Order: Blattodea

Pheromosa (#795).

Pheromosa is a bipedal anthropomorphic Pokémon. It has a rather slender build and is mostly white. Pheromosa originates from the Ultra Desert dimension in Ultra Space. Pheromosa is based on generic cockroaches just after they have molted (Fig. 2); during this stage, the animals are pale and vulnerable until their exoskeleton hardens and darken.

Figure 2. A freshly-molted cockroach (family Blattidae), leaving its exuvia behind. Credit: Donald Hobern (2010), Wikimedia Commons.

Order: Orthoptera

Family: Gryllidae

Kricketot (#401) evolves to Kricketune (#402).

Kricketot is a bipedal, bug-like Pokémon. It has a red body with some black and white markings. By shaking its head and rubbing its antennae together, it can create a sound that it uses to communicate. Based on its appearance, it is a cricket.

Kricketune is also a bipedal Pokémon with an insectoid appearance, also primarily red with some black and tan colored markings. It can produce sound by rubbing its arms on the abdomen. Kricketune appears to be based on crickets due to their sound-producing ability, but it somewhat resembles a violin beetle.

Both Kricketot and Kricketune are depicted with only 4 limbs, whereas insects are largely defined by having exactly six legs.

Order: Hemiptera

Families: Gerridae and Fulgoridae

Surskit (#283) evolves to Masquerain (#284).

Surskit is a blue insectoid Pokémon with some pink markings. It produces some sort of syrup, which is exuded as a defense mechanism or to attract prey. This Pokémon can also secrete oil from the tips of its feet, which enables it to walk on water as though skating. Surskit usually inhabits ponds, rivers, and similar wetlands, where it feeds on microscopic, aquatic organisms. This Pokémon is based on water striders. However, a water strider does not ooze syrup and neither does it need oil to walk on water; it can walk on water due to the natural surface tension.

Masquerain is a light blue Pokémon with two pairs of wings. On either side of its head is a large antenna that resembles an angry eye. These eyespots are used by many real-life moths and lantern-flies to confuse and intimidate would-be predators. Masquerain is in fact based on a lantern-fly.

Both “species”, water striders and lantern-flies, are only distantly related, belonging to two different families within the “true bugs” (Hemiptera).

Family: Cicadidae

Nincada (#290) evolves to Ninjask (#291) and then to Shedinja (#292).

Nincada is a small, whitish, insectoid Pokémon. The claws are used to carve the roots of tree and absorb water and nutrients. Nincada builds underground nests by the roots of trees. It is based on a cicada nymph, which lives underneath the soil surface. However, a cicada nymph usually does not have fully developed wings. Instead, they have short wing stubs which eventually will become fully functional wings – as usual amongst hemimetabolous insects.

Ninjask is a small, cicada-like Pokémon with two pairs of wings. Its body is mostly black with some yellow and grey markings. Ninjask is a very fast Pokémon and it can seem invisible due to its high speed. It is based on an adult cicada, with the colors somewhat resembling Neotibicen dorsatus (Say, 1825) (Fig. 3).

Shedinja is a brown and grey insectoid Pokémon. A hole between its wings reveals that its body is completely hollow and dark, as it possesses no internal organs. It is based on the shed husk (exuvia) that cicadas and other hemimetabolous insects leave behind when they molt.

Figure 3. Adult female of Neotibicen dorsatus, the bush cicada. Credit: Yakkam255 (2015), Wikimedia Commons.

Paras (#046) evolves to Parasect (#047).

Paras is an orange insectoid Pokémon with an ovoid body. On the top it has two little red and yellow mushrooms known as tōchūkasō. The mushrooms can be removed at any time, and grow from spores that are doused on this Pokémon’s back at its birth by the mushroom on its mother’s back. Tōchūkasō is an endoparasitoid that replaces the host tissue and can affect the behavior of its insect host. The base insect is based on a cicada nymph. The real-world tōchūkasō live on hepialid caterpillars in Tibet. However, there are many more species of entomopathogenic fungi in the world, most notable the genus Cordyceps (L.) Fr. (1818).

Parasect is an orange, insectoid Pokémon that has been completely overtaken by the tōchūkasō mushroom. The adult insect has been drained of nutrients and is now under the control of the fully-grown tōchūkasō. Parasect can thrive in dank forests with a suitable amount of humidity for growing fungi. The base insect is a deformed version of what is probably a cicada nymph, the parasitic mushroom having caused a form of neoteny, when the adults look like a juvenile form.

Order: Neuroptera

Family: Myrmeleontidae

Trapinch (#328) evolves to Vibrava (#329) and then to Flygon (#330).

Trapinch is an orange, insectoid Pokémon. This Pokémon lives in arid deserts, where it builds its nest in a bowl-shaped pit dug in sand. It sits in its nest and waits for prey to stumble inside. Once inside, the prey cannot climb back out. It is based on the larval stage of the antlion, which lives in conical sandy pits before maturing into winged adults.

Vibrava is a dragonfly-like Pokémon. Vibrava’s wings are not fully developed, so it is unable to fly very far. However, it is able to create vibrations and ultrasonic waves with its wings, causing its prey to faint. Vibrava is a saprotroph – it spits stomach acid to melt its prey before consumption. Vibrava is based on the adult stage of an antlion. Adult antlions and dragonflies look from a distance quite similar and are therefore often mistaken for each other.

Flygon is a desert-dwelling insectoid dragon with a green body and one pair of wings. Its wings make a “singing” sound when they are flapped. It uses this unique ability to attract prey, stranding them before it attacks. It is based on the winged, adult stage of the antlion.

Order: Coleoptera

Family: Lucanidae

Pinsir (#127, incl. Mega-Pinsir).

Pinsir is a bipedal beetle-like Pokémon with a brown body and a large pair of grey, spiky pincers on top of its head. Pinsir is based on a stag beetle.

Grubbin (#736) evolves to Charjabug (#737) and then to Vikavolt (#738).

Grubbin is a small insectoid Pokémon. It has a white body with three nubs on either side resembling simple legs. Grubbin typically lives underground. It uses its jaw as a weapon, a tool for burrowing, and for extracting sap from trees. Grubbin appears to be based on a larval beetle, also known as “grubs”.

Charjabug is a small cubic Pokémon resembling an insect-like battery. Its body consists of three square segments with two brown stubs on each side. It generates and stores electricity in its body by digesting food. This energy is stored in an electric sac. Charjabug appears to be based on a cocooned bug and a battery. It may also be based on the denkimushi (Monema flavescens Walker, 1855), a caterpillar in Japan that, when touched, can give a sting that is said to feel like an electric shock (Fig. 4).

Vikavolt is a beetle-like Pokémon with a large pair of mandibles. It produces electricity with an organ in its abdomen, and fires powerful electric beams from its huge jaws. Vikavolt appears to be based on a stag beetle. Its straight, scissor-like mandibles resemble those of Lucanus hayashii Nagai, 2000.

Figure 4. Larva of Monema flavescens. Credit: Pan et al. (2013), Wikimedia Commons.

Family: Coccinellidae

Ledyba (#165) evolves to Ledian (#166).

Ledyba is a red ladybird-like Pokémon with five black spots on its back. Female Ledyba have shorter antennae than male Ledyba. Ledyba is a very social Pokémon, e.g. in the winter they gather together to keep each other warm. Ledyba is probably based on the five-point ladybird Coccinella quinquepunctata Linnaeus, 1758 due to its color and/or on the harlequin ladybird Harmonia axyridis (Pallas, 1773), which clusters together in the winter.

Ledian is a large red bipedal ladybird-like Pokémon. Female Ledians’ antennae are shorter than the males’. Ledian sleeps in forests during daytime inside a big leaf.

Family: Scarabaeidae

Heracross (#214, incl. Mega-Heracross).

Heracross is a bipedal beetle-like Pokémon with a blue exoskeleton. The prolonged horn on its forehead ends in a cross-shaped (males) or heart-shape (females) structure. Heracross is most likely based on the Japanese rhinoceros beetle Allomyrina dichotoma Linneaus, 1771 (Fig. 5).

Figure 5. Adult male of Allomyrina dichotoma. Credit: Lsadonkey (2016), Wikimedia Commons.

Family: Lampyridae

Volbeat (#313) and Illumise (#314).

Volbeat is a bipedal firefly-like Pokémon. Its body is black with some blue, yellow, and red portions. It has a spherical yellow tail, which glows to communicate and draws geometric patterns in the sky while in a swarm. This is a male only Pokémon “species”; Illumise is its female counterpart. Volbeat lives in forests near clean ponds and is attracted by the sweet aroma given off by Illumise. It is based on a firefly like its counterpart Illumise. Its appearance may be based on a greaser, a subculture from the 1950’s.

Illumise is a bipedal firefly-like Pokémon. It is black and blue with some yellow markings. This is a female only Pokémon “species”; Volbeat is its male counterpart. It is a nocturnal Pokémon that lives in forests.  Illumise does not seem to share its coloring with any particular species. Illumise may be based on flappers, a 1920’s women’s style. Its mating behavior only slightly resembles the behavior of real-world fireflies, in which females use light signals to attract mates.

Family: Elateridae

Karrablast (#588) evolves to Escavalier (#589).

Karrablast is a round bipedal Pokémon with a yellow and blue body. When it senses danger, it spews an acidic liquid from its mouth. It targets another Pokémon, Shelmet, so it can evolve. It resides in forests and fields, and it often hides in trees or grass if threatened. Karrablast may be based on a Japanese snail-eating beetle due to its preference for attacking Shelmet, a snail-like Pokémon.

Escavalier is an insectoid Pokémon wearing a knight’s helmet. Its tough armor protects its entire body. It flies around at high speed, jabbing foes with its lances. Escavalier is probably based on the Drilus Olivier, 1790 genus, with references to a jousting knight. Drilus larvae are known for eating snails and stealing their shells, explaining why it attacks Shelmet and takes its shell to evolve into Karrablast.

Order: Hymenoptera

Family: Tenthredinidae

Weedle (#013) evolves to Kakuna (#014) and then to Beedrill (#015, incl. Mega-Bedrill).

Weedle is a small larval Pokémon with a body ranging in color from yellow to reddish-brown. It has a conical venomous stinger on its head and a barbed one on its tail to fend off enemies. Weedle can be found in forests and usually hides in grass, bushes, and under the leaves it eats. Weedle appears to be based on the larva of a wasp or hornet, although these real-world larvae usually don’t have defense strategies. The only larvae which feed directly off leaves are those of sawflies.

Kakuna is a yellow cocoon-like Pokémon. Kakuna remains virtually immobile and waits for its “evolution” to happen, often hanging from tree branches by long strands of silk. Although Kakuna is the pupa stage of a Hymenoptera, it showcases a silky cocoon, a feature usually found in Lepidoptera and only some Hymenoptera, like sawflies.

Beedrill is a bipedal, wasp-like Pokémon. Its forelegs are tipped with long, conical stingers. It stands on its other two legs, which are long, segmented, and insectoid in shape. Beedrill has two pairs of rounded, veined wings, and another stinger on its yellow-and-black striped abdomen. By its color pattern, Beedrill looks like a vespid wasp, but due to the previous stages of this Pokémon species, it must be based on Tenthredo scrophulariae Linneaus, 1758, the figwort sawfly.

Family: Apidae

Combee (#415) evolves to Vespiquen (#416, female).

Combee is a small insectoid Pokémon that resembles three social bees inside three hexagonal pieces of honeycomb stuck together; the top two have wings. Female Combee have a red spot on the forehead. Male Combee are not known to evolve into or from any other Pokémon. The sex ratio of Combee is 87.5% male and 12.5% female. Combee can fly with its two wings as long as the top two bees coordinate their flapping. They gather honey, sleep, or protect the queen. Combee is based on a mix of bees and their larvae living in honeycombs. (Bees arrange their honeycombs in a vertical manner, whereas wasps arrange them horizontally.)

In the hive of the real-world honey bee (Apis mellifera Linneaus, 1758), there is usually one queen bee and up to 40.000 female workers. So, the sex ratio of Combee does not reflect the ratio of female (workers) and male (drones) honey bees, but of the reproductive bees, the drones and the fertile queens. The larger number of drones is needed, since each queen will often mate with 10–15 males before she starts a new hive. Usually, drones can make up to 5% of the bees in a hive.

Vespiquen is a bipedal bee-like Pokémon with a yellow and black striped abdomen resembling an elegant ballroom gown. Underneath the expansive abdomen are honeycomb-like cells that serve as a nest for baby Combee. Vespiquen is a female-only Pokémon “species”. Vespiquen is the queen of a Combee hive, controlling it and protecting it, as well as giving birth to young Combee. The horizontal honeycombs hints that this “species” is a wasp rather than a bee.

Family: Formicidae

Durant (#632).

Durant is an ant-like Pokémon with a grey body and six black legs. It is territorial, lives in colonies and digs underground mazes. Durant grows steel armor to protect itself from predators. Durant is based on an ant, possibly the Argentine ant (Linepithema humile Mayr, 1868), due to the jaw and their invasive behavior.

Order: Lepidoptera

Family: Papilionidae

Caterpie (#010) evolves to Metapod (#011) and then to Butterfree (#012).

Caterpie is a green caterpillar-like Pokémon. It has yellow ring-shaped markings down the sides of its body and bright red “antenna” (osmeterium) on its head, which releases a foul odor to repel predators. The appearance of Caterpie helps to startle predators; Caterpie is probably based on Papilio xuthus Linnaeus, 1767, the Asian swallowtail (Fig. 6). The osmeterium is a unique feature of swallowtails. Caterpie will shed its skin many times before finally cocooning itself in thick silk. Its primary diet are plants.

Metapod is a green chrysalis Pokémon. Its crescent shape is based upon a Swallowtail chrysalis with a large nose-like protrusion and side protrusions resembling a Polydamas Swallowtail or Pipevine Swallowtail chrysalis (genus Battus Scopoli, 1777).

Butterfree is a butterfly Pokémon with a purple body and large, white wings, somewhat resembling a black-veined white Aporia crataegi (Linneaus, 1758). Although it is supposed to be a butterfly, it lacks the proboscis, which is typical of Lepidoptera, and presents teeth instead. Additionally, the body does not consist of the typical three segments of insects. Therefore, each stage seems to be based on a different species.

Figure 6. Larva of Papilio xuthus, with everted orange osmeterium. Credit: Alpsdake (2011), Wikimedia Commons.

Families: Geometridae and Arctiidae

Venonat (#048) evolves to Venomoth (#049).

Venonat has a round body covered in purple fur, which can release poison. It feeds on small insects, the only Lepidoptera caterpillar which is known to feed on prey instead of leaves belong the genus Eupethecia Grote, 1882 (Geometridae). However, Venonat does not resemble a caterpillar in general body shape or numbers of legs.

Venomoth is a moth-like Pokémon with a light purple body and interestingly two small mandibles. It has two pairs of wings, which are covered in dust-like, purple scales, although the color varies depending on their toxic capability. Dark scales are poisonous, while lighter scales can cause paralysis. These scales are released when Venomoth flutters its wings. The general appearance resembles species belonging to the Actiidae.

There is no cocoon stage for this species it is doubtful whether both stages were based on the same real-life species.

Family: Riodinidae

Scatterbug (#664) evolves to Spewpa (#665) and then to Vivillon (#666).

Scatterbug is a small caterpillar Pokémon with a grey body. If threatened by a bird Pokémon, it can spew a powder that paralyzes on contact. Similarly, the large white butterfly Pieris brassicae (Linneaus, 1758) is known to throw up a fluid of semi-digested cabbage, which contains compounds that smell and taste unpleasant to predators, such as birds.

Spewpa is a small insectoid Pokémon with a grey body covered by white furry material. In order to defend itself, Spewpa will bristle its “fur” to threaten predators or spray powder at them. Spewpa is based on a generic pupa of a moth or butterfly, probably a silkworm cocoon.

Vivillon is a butterfly-like Pokémon with wings that come in a large variety of patterns, depending in which climate it lives or rather, in which real-world region the player is. There is a total of 20 patterns known. It would be interesting to know whether they evolved due to allopatric speciation or if it is a case of mimicry.

Family: Psychidae

Pineco (#204) evolves to Forretress (#205).

Pineco is a pine cone-like Pokémon without visible limbs. It is based on a bagworm, the caterpillar stage of psychid Lepidoptera. Bagworms cover themselves with a case (the bag) made of surrounding material. This Pokémon uses tree bark and thus resembles a pine cone.

Forretress is a large spherical Pokémon, also without any visible limbs. It lives in forests, attaching itself immovably to tree trunks. Forretrees is also based on a bagworm.

Different bagworm species are adapted to their environment, to the plants they eat, and to the materials available for producing their case. Therefore, Pineco and Forretress are actually based on two different species, as they both are caterpillars. There is no adult stage for this Pokémon.

Burmy (#412) evolves to Wormadam (#413, female) or Mothim (#414, male).

Burmy is a small pupa-shaped Pokémon with a black body and six stubby legs. It is based on a bagworm pupa, which will metamorphose into a winged moth if male, or wingless moth if female. Burmy can change its “cloak” (case) depending on the environment it last battled.

Wormadam is a black bagworm-like Pokémon with a cloak of leaves, sand, or building insulation. Its cloak depends on Burmy’s cloak when it evolved, and so does it type (Grass, Ground or Steel). It is a female-only “species”, with Mothim as its male counterpart. Female psychid moth either don’t have wings at all or have only small wing stubs that don’t develop fully.

Mothim is a moth-like Pokémon with two pairs of legs and two pairs of wings, one larger than the other. Mothim is a nomadic nocturnal Pokémon, searching for honey and nectar. Instead of gathering honey on its own, it raids the hives of Combee. It is a male-only “species”, with Wormadam as its female counterpart.

Family: Nymphalidae

Wurmple (#265) evolves to Silcoon (#266) and then to Beautifly (#267).

Wurmple is a small caterpillar-like Pokémon with a mostly red body and many spikes on the top of its body. It can spit a white silk that turns gooey when exposed to air. Spikes or hairy appendages are common amongst nymphalid caterpillars. Also, it has five pairs of legs, whereas insects are known to have only three pairs of legs. However, many lepidopteran caterpillars have additionally “prolegs” (small fleshy stub-like structures) to help them move.

Silcoon is a cocoon-like Pokémon which is completely covered by white silk. Silcoon also uses the silk to attach itself to tree branches. Nymphalid cocoons are usually not woolly or hairy, but smooth.

Beautifly is a butterfly-like Pokémon with two pairs of wings. Beautifly has a long and curled black proboscis that it uses to drain body fluids from its prey. In the real world, Lepidoptera usually drink the nectar of flowers. One of the few exceptions are the species of the genus Calyptra Ochsenheimer, 1816, which pierce skin of animals and drink blood.

Family: Saturniidae

Wurmple (#265) evolves to Cascoon (#268) and then to Dustox (#269).

The caterpillar stage of this species is morphologically identical to the caterpillar stage of the “species” above: Wurmple. It appears that Wurmple can evolve in two forms: due to mimicry, sympatric speciation or are there morphological or biological characters, which have not been notices yet?

Cascoon is a round cocoon-like Pokémon covered in purple silk. Saturniid cocoons are usually covered in silk.

Dustox is a moth-like Pokémon. It has a purple body, two pairs of tattered green wings, and – just like Beautifly – two pairs of legs. Dustox is nocturnal and is instinctively drawn to light. Clearly, this is a moth. Some of the markings on its wings resemble typical markings of noctuid moths, but the big “fake eye” is typical of saturniids.

Larvesta (#636) evolves to Volcarona (#367).

Larvesta is a fuzzy caterpillar-like Pokémon. It has five red horns on the sides of its head, which it can use to spit fire as a defensive tactic to deter predators. Larvesta is based on a saturniid caterpillar.

Volcarona is a large moth-like Pokémon with four small feet and three pairs of wings. It releases fiery scales from its wings. Just like Larvesta, Volcarona is based on a saturniid moth, likely the Atlas moth Attacus atlas (Linneaus, 1758).

Order: Diptera

Family: Bombyliidae

Cutiefly (#742) evolves to Ribombee (#743).

Cutiefly is a tiny Pokémon with large wings. Cutiefly appears to be based on the bee fly, specifically the species Anastoechus nitidulus (Fabricius, 1794) (Fig. 7).

Ribombee is a tiny insectoid Pokémon with a large head, slightly smaller body, and thin arms and legs. It is covered in fluffy yellow hair. Two wings nearly as large as its body sprout from its back. The wings are clear with three brown loop designs near the base. Its four thin limbs have bulbous hands or feet. Ribombee uses its fluffy hair to hold the pollen it collects from flowers. It is based on a bee fly.

Figure 7. Adult of Anastoechus nitidulus. Credit: karakotokako (2007), image retrieved from https://

Family: Culicidae

Buzzwole (#794).

Buzzwole is a bipedal anthropomorphic Pokémon. It has four legs and two pairs of orange translucent wings. It uses its proboscis to stab and then drink “energy” off its enemies/prey. Buzzwole originates from the Ultra Desert dimension in Ultra Space. It is based on a mosquito and may specifically derive inspiration from Aedes albopictus (Skuse, 1894), which is an invasive species worldwide.

Mixed Orders: Lepidoptera and Phasmatodea

Families: Tortricidae, Hesperiidae, and Phylliidae

Sewaddle (#540) evolves to Swadloon (#541) and then to Leavanny (#542).

Sewaddle is a caterpillar-like Pokémon with a green body with three pairs of legs. It makes leafy “clothes” using chewed-up leaves and a thread-like substance it produces from its mouth. The leafy hood helps Sewaddle to hide from enemies. Sewaddle appears to be based on the caterpillar of the silver-spotted skipper Epargyreus clarus (Cramer, 1775), which produce silk and fold leaves over themselves for shelter (Fig. 8).

Swadloon is a round yellow Pokémon inside of a cloak of leaves. It lives on the forest ground and feeds on fallen leaves. Swadloon appears to be based on the chrysalis of Epargyreus clarus. Epargyreus clarus fold leaves over themselves for shelter as they age and, when cocooning, eventually use silk to stick the leaves together and form its chrysalis.

Leavanny is a bipedal, insectoid Pokémon with a yellow and green body with leaf-like limbs. It lives in forests and uses its cutters and sticky silk it produces to create leafy “clothing”. It also warms its eggs with fermenting fallen leaves. Leavanny has the features of several insects. Primarily it appears to be a bipedal leaf-insect (Phylliidae). Its general body structure is also similar to that of Choeradodis Serville, 1831 mantises, which also have laterally expanded thoraxes and abdomens.

Figure 8. Larva of Epargyreus clarus. Credit: Seth Ausubel (2013), image retrieved from


Only 11 insect orders (out of 30) are represented in the Pokémon world. Possible more, as differentiation of insect Pokémon and non-insect Pokémon are sometimes difficult. The main reason is, that many insect Pokémon are not depicted as a typical insect with its segmented body, the six legs, and two pairs of wings[1]. Many are depicted as bipedal (e.g., #401 Kricketot) or even in an anthropomorphic way (e.g., #795 Pheromosa). Also, insectoid Pokémon typically have only four limbs (instead of six). Many insectoid Pokémon also have fewer wings than insects (except for #637 Volcarona, which has more). Therefore, the definition of what is an insect Pokémon is debatable.

One clue is to look at the types each Pokémon belongs to. However, from the circa 80 Bug-type Pokémon, only about 60 are insects. The others belong to other arthropods groups, like Chelicerata, Crustacea, and Myriapoda. This is not surprising, as often creepy crawlies (basically everything that is small with legs) are all addressed as “bugs”. In fact, only member of the insect order Hemiptera are called “true bugs”.

Interestingly, Prado & Almeida (2017) have included Pokémon on their insect list, which are doubtful: #251 Celebi, #247 Pupitar, and #206 Dunsparce. None of them are considered insects here. Celebi may resemble a bipedal somewhat anthropomorphic insectoid, but nothing of the lifestyle or beyond the vague appearance gives a clue to an insect. Similarly, #247 Pupitar, might look like a pupa of an insect. However, both its “larval” stage (#256 Larvitar) and its final stage (#248 Tyranitar) resemble a dinosaur or some sort of dragon. Only the hint of “pupa” in its name, links Pupitar to an insect. Lastly, #206 Dunsparce was classified as a Hymenoptera by Prado & Almeida (2017). Is may look somewhat like an insect, even showing two pairs of wings (and no legs at all). Dunsparce, however, is based on a mythical “snake-like animal” called Tsuchinoko, also known as “bachi hebi” (or “bee snake”). Finally, Prado & Almeida (2017) have classified #212 Scizor as “unknown”, but here it is treated as a praying mantis (Mantodea). Similarly, those authors have classified #284 Masquerain as a Lepidoptera, but here we treat is as a true bug (Hemiptera).

Lastly, #649 Genesect resembles somewhat an ant covered by steel. However, according to the Pokédex (Pokémon Website, 2018), it is a man-made machine.

Compared to the vertebrates (birds, mammals, reptiles, amphibians, and fishes), many more insects live on earth (66,000 described species to about 1 million, respectively; Zhang, 2011). This ratio is, however, not represented in the Pokémon world (Table 3), most likely due to the fact that the majority of people prefer (cute and cuddly) furry animals over creepy insects, even though butterflies and dragonflies are regarded as beautiful.


Borror, D.J.; DeLong, D.M.; Triplehorn, C.A. (1981) An Introduction to the Study of Insects. Saunders College, Philadelphia.

Bulbapedia (2018) The community driven Pokémon encyclopedia. Available from: http://bulbaped (Date of access: 10/Sep/ 2018).

Dorward, L.J.; Mittermeier, J.C.; Sandbrook, C.; Spooner, F. (2017) Pokémon GO: benefits, costs, and lessons for the conservation movement. Conservation Letters 10(1): 160–165.

Gibson, H. (2005) Detailed Discussion of Dog Fighting. Michigan State University, East Lansing.

Jacobs, A. (2011) Chirps and sheers: China’s srickets slash. The New York Times. Available from: world/asia/chirps-and-cheers-chinas-crickets-clash-and-bets-are-made.html (Date of access: 10/Oct/2018).

Kent, S.L. (2001) The Ultimate History of Video Games. Crown Publishing Group, New York.

Morrow, L. (1995) History they don’t teach you: a tradition of cockfighting. White River Valley Historical Quarterly 35(2): 5–15.

Official Pokémon Website, The. (2018) The Official Pokémon Website. Available from: http://poke  (Date of access: 10/Sep/2018).

Prado, A.W. & Almeida, T.F.A. (2017) Arthropod diversity in Pokémon. Journal of Geek Studies 4(2): 41–52.

Whitehill, S.; Neves, L.; Fang, K.; Silvestri, C. (2016) Pokémon: Visual Companion. Pokémon Company International / Dorling Kindersley, London.

Zhang, Z.-Q. (2011) Animal biodiversity: an outline of higher-level classification and survey of taxonomic richness. Zootaxa 3703: 1–82.


I am grateful to Seth Ausubel (https://www. for kindly granting permission to use his photograph of Epargyreus clarus on this article. I would also like to thank Miles Zhang for valuable comments on an earlier version of the manuscript.


Dr. Rebecca Kittel is an entomologist working on parasitoid wasps. She is interested in all sorts of interactions of insects with human beings, regardless of whether they are real-life insects or purely fictional.

[1] Not all insects have two pairs of wings, though. For instance, the Diptera (flies) have only one, while the Siphonaptera (fleas) have none.

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Zoological Nomenclature of Ice and Fire

Evangelos Vlachos

CONICET & Museo Paleontológico Egidio Feruglio, Trelew, Chubut, Argentina.

Email: evlacho (at) mef (dot) org (dot) ar

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Valar gūrēñis — All men must learn

The diversity of the World of Ice and Fire (Westeros, Essos and the other continents combined) is remarkable. All kinds of species of animals and plants are known, including some mythical creatures. The purpose of this contribution is to provide a system of nomenclature for the most important animal species from the World of Ice and Fire. This new system is based on the High Valyrian language, and aims to provide a set of names that can be applied to the various species of life that survived, or even became extinct, in this world.

The World of Ice and Fire is a fictional world. Although most of the wild and domesticated animals are the same or similar to our own, there several animals that are unique to it. Also, more than one ‘species’ of humans survive in this world, now mostly isolated in remote islands like Ibben and the Sothoryos. The Common Tongue, spoken mainly in the Seven Kingdoms of Westeros, is given to us through the books in English; but this doesn’t mean that it is English. Even if a direwolf is called a direwolf in the books, it probably sounded differently in the Common Tongue.

Back to our world, following the pioneering work of C. Linnaeus in 1758 the need of a stable and universal system of biological nomenclature became necessary. Since then, a set of rules has been created, revised, used and applied to Zoological Nomenclature, forming the so-called International Code of Zoological Nomenclature (ICZN, or simply ‘the Code’). The latest edition was published in 1999, and some parts of the Code have been recently (2012) amended to include names and acts published in electronic-only journals.

I will briefly present the main features of this system of nomenclature for those not entirely familiar with it. The backbone concept of nomenclature is the binomen: each species name is formed by two components, the genus name and the specific name; both are written in italics and the genus name is capitalized (e.g., Homo sapiens). The ICZN offers a graphical summary of the whole process of naming animal taxa[1], which is summarized in Box 1 below. The reader should, of course, consult the Code for further details.

Box 1. Basic steps for naming taxa

  1. The name must be contained in a published work (published sensu the ICZN);
  2. The name must be available (sensu the ICZN);
  3. The name must be properly formed, following the instructions of the ICZN.

Names that do not conform these rules are unavailable names (including the so-called ‘naked names’), and can be made available later for the same or different concept. If these conditions are met, the available names enter the zoological literature. Once part of the literature, the names ‘compete’ for validity, which mainly refers to the so-called ‘Principle of Priority’. Simply put, the oldest available name applied to a taxon is the valid name for this taxon (Art. 23.1, ICZN). The other names are invalid names, including synonyms, homonyms, and dubious names. Of course, in real life things are not so simple, as there are several exemptions from these rules and a multitude of complicated cases; the Code contains numerous articles and examples that try to account for all these situations.

Obviously, the purpose of this article is to propose a set of names for the animals of the World of Ice and Fire, but a curious reader might ask: do those names also become part of the ‘real life’ zoological nomenclature? The answer is no, these names will not form part of the zoological nomenclature for the main following reasons:

  1. As the Journal of Geek Studies is an electronic publication, any name (or nomenclatural act) published in it should conform to the rules of Art. 8.5 (ICZN) for works published/distributed electronically. But it fails to conform to the provisions of the sub-article 8.5.3, which mandates the registration of the work and the names on the Official Register of Zoological Nomenclature (a.k.a. ZooBank).
  2. Even though several of the animals of the World of Ice and Fire are referred to the Common Tongue with similar names and concepts of wild and domesticated animals that exist or existed in our world (e.g., a dog, a horse, a mammoth), those animals are actually purely hypothetical concepts (sensu Art. 1.3.1, ICZN) that exist in the fantasy World of Ice and Fire and the mind of G.R.R. Martin. Thus, they are excluded from the zoological nomenclature.
  3. The names, as published herein, are not formed properly according to the Code. Both words are capitalized, not italicized, with diacritic signs, and are connected by a dash.

Therefore, all the names herein are unavailable names for our ‘real life’ zoological nomenclature. I suppose that a similar need of a system of nomenclature would be eventually necessary in the World of Ice and Fire as well, most probably among its scholars—the Maesters. The study of the natural world has largely been neglected by the great Maesters of the Citadel, in Oldtown. Maester Yandel in his work (Martin et al., 2014) provides some basic information on various animals — in many cases by citing other authors — but without any specific focus on nature. However, one cannot understand and explain the mysteries of the world, unless they are able to explain and describe the life on it. Therefore, and to avoid misunderstandings among Maesters across the continent, this new system of nomenclature would greatly assist in the communication among scholars in the World of Ice and Fire.

I strongly insist that the Maesters of the Citadel should try to promote the study of the natural mysteries of the world. I further propose that the Maester who will complete the study of a significant portion of the natural world should be awarded a wooden link to add to his chain. This link should be made by a weirwood tree and would symbolize that all life on the World is related, and originated from a common root, just like the branches and leaves of a weirwood tree.


In order to differ from the common, vernacular, names of the animals in the Common Tongue of the World of Ice and Fire, their scientific names will be created in the High Valyrian.

The Valyrian languages are a group of languages that were spoken in the past, with High Valyrian being spoken in Valyria and its descendants languages (Astapori and Meereenesse Valyrian) spoken in Astapor and Meereen respectively, as well as a variety of dialects and corruptions of the pure High Valyrian spoken in the Free Cities (Martin et al., 2014). Although several words in High Valyrian were already present in the books of the series The Song of Ice and Fire written by G.R.R. Martin, the language was created by D.J. Peterson for the TV series (Peterson, 2013).

For the purpose of establishing the ‘Zoological Nomenclature of Ice and Fire’, the names will be written in High Valyrian, with the use of the letters of the Latin alphabet (High Valyrian was certainly written in its own alphabet). The source of linguistic information is the Dothraki Wiki (2018; information stored therein is copyrighted by the Language Creation Society, HBO, and G.R.R. Martin).

The main objective of this work is to name the main species of animals (e.g., the species of humans) and also provide some names for large groups (e.g., a name for ‘mammals’). The basic information comes from the bestiary of A Wiki of Ice and Fire (2018, and references therein). Parts of this work have been preliminary published in the subreddit r/asoiaf ( by the author, under the alias E_v_a_n (2017, and references therein). Very few names have been proposed by some other redditors and they are not included herein. The terms ‘species’, ‘subspecies’, and ‘genus’ are used in a similar sense as in modern taxonomy and nomenclature for simplicity.

The various names were created based on the following basic rules and recommendations, which are illustrated by examples where necessary. The formation of the majority these rules is based largely on valuable comments of David J. Peterson, whom I deeply thank.

Rule 1. Names for large groups consist of a single word, whereas names for ‘species’ consist of two words. Example: Valar for humans, Sylvie-Valar for the wise humans, which is included in Valar.

Rule 2. The two words comprising the ‘species’ names are hyphenated and each start with a capital letter. We do not know if such kind of punctuation was present in High Valyrian. The purpose of adding the hyphen here is mainly to distinguish these names from original binomina in nomenclature.

Rule 3. Group names are written in small capitals. This rule is only for stylistic purposes.

Rule 4. All original diacritics of High Valyrian must be kept. Besides its stylistic purpose, the application of this rule further distinguishes the names herein from original names in nomenclature.

Rule 5. Formation of group names is done either with nouns in the collective or adjectives with the addition of the derivational affix –enka (meaning ‘like’). Example A: To form the name of the group of humans (‘equivalent’ to a genus name) we could use the word ‘vala’ (1lun; man) in the collective, as Valar. Example B: To form the name of the group of reptile-like animals we could use the word ‘rīza’ (1lun; reptile, lizard) with the addition of the derivational affix –enka (adj. I), as Rīzenka. Note that in this case we need to use only the root of the word ‘rīza’ (rīz–).

Rule 6. Formation of a species name is done with the combination of an adjective and a noun in the collective. Note that adjectives must agree in gender (i.e., lunar, solar, terrestrial, aquatic), case, and number, with the noun they modify; however, as the noun is in the collective, the adjective should be in the singular. Also, the adjective goes before the noun it modifies. Example A: To create the name for the wise humans we could use the combination of the noun ‘Valar’ (1lun; ‘all the men’, in the collective) with the adjective ‘Sylvie’ (adj. III). The singular of this adjective would be ‘Sylvie’ for lunar/solar and ‘Sylvior’ for terrestrial/aquatic (in the singular; see Rule 5 above). As the word ‘Valar’ is of lunar gender, it should be combined with the adjective in the lunar gender as well, as Sylvie-Valar. Example B: To create an adjective from a noun one should use one of the derivational affixes like –enka (adj. I) (see Rule 5). Again, there must be agreement in gender.

Rule 7. To create a name that consists of three components (‘equivalent’ to a subspecies or for other purposes), insert the third component in its proper place according to the desired meaning, again in agreement to Rule 6. Example: For the name of the white walkers, supposedly a further subdivision of the wise humans, we could use the name Sylvie-Valar, inserting in between the adjective ‘Timpa’ (adj. I) in the lunar gender and in singular, as Sylvie-Timpa-Valar. In this arrangement it reads: ‘all the wise white men’. Contrary to our own nomenclature, the position of the components may vary depending on the desired meaning. For example, ‘all the white wise men’ would read as Timpa-Sylvie-Valar. Both versions are equivalent for nomenclatural purposes herein.

Rule 8. To form a name from a toponym, one should add the derivational suffix –sīha, or –īha (depending if the root ends in consonant or vowel), to form an adjective of Class I. It then follows in agreement to Rule 6. Alternatively — and this could be done with other names as well, not only with toponyms — one could use the derivational suffix –ōñe (which means ‘from the’) to form a Class II adjective. Example A: To name the species of humans from Ibben, we could add the suffix –īha, as Ibbenīha-Valar. In this form it reads: ‘all the Ibbenian humans’. Example B: Ibbenōñe-Valar. In this form, it reads: ‘all the humans from Ibben’. This is a quite useful suffix to form many other names as well (see below).

All original information below comes from The Song of Ice and Fire books (Martin, 1996, 2000, 2005, 2011) and The World of Ice and Fire (Martin et al., 2014). For simplicity, I will not add these citations below.

The relationships among the main ‘species’ named herein are depicted across the branches of a weirwood tree (Fig. 1).

Figure 1. The taxonomy of the animals of the World of Ice and Fire, depicted on the branches of a weirwood tree.

The maps presented herein (Figs. 2 and 4) are based on the original map available in Wikimedia Commons (CC-BY-SA 4.0), which was subsequently edited in Adobe Photoshop (removing words) and Adobe Illustrator (tracing) to create the final ‘clean’ version for this article. Silhouettes of animals are re-drawn manually from pictures available online with permission to be modified.

Figure 2. The distribution of known animal species in the World of Ice and Fire, excluding those with cosmopolitan distribution.

Abbreviations: Nouns: numbers denote the declension, followed by the abbreviated gender (aq, aquatic; lun, lunar; sol, solar; ter, terrestrial). Adjectives (adj.): Roman numerals indicate the class.



(all the names; from the noun ‘brōzi’, 5lun, meaning ‘name’)

Dȳñenka, animals.

Etymology. Dȳñenka, from the word ‘dȳñes’ (4sol; animal) and the suffix –enka (adj. I), which means ‘like’; altogether the name means ‘animal-like’.

Remarks. The distribution of the animals of the World of Ice and Fire is shown in Figure 2. Those with a roughly cosmopolitan distribution (e.g., horses) were excluded for simplicity.

Jūlrenka, mammal-like animals.

Etymology. Jūlrenka, from the word ‘jūlor’ (3aq; milk) and the suffix –enka (adj. I).

Uēpys-Nusper, all the ancient cows or aurochs.

Etymology. Uēpys from the adjective ‘uēpa’ (adj. I; old); Nusper from the nominative collective of the noun ‘nuspes’ (4sol; cow).

Remarks. This is the ancestor of the modern-day cows, and was larger, with longer and more robust horns. Although not present in most of Westeros as a result of domestication, their presence is reported beyond the Wall, and are served in feasts in some of the Great Houses of the North.

Lantarōvatsienkys-Ñomber, all the elephants with two big teeth.

Etymology. Lantarōvatsienkys, from the combination of the words ‘lanta’ (adj. I; two), ‘rova’ (adj. I; big), ‘atsio’ (3lun; tooth), and the suffix –enkys, referring to the animals’ large tusks; Ñomber from the noun ‘ñombes’ (4sol; elephant).

Remarks. Native to Essos, quite common in Astapor.

Krubenkys-Ñombītsor, all the dwarf elephants.

Etymology. Krubenkys, from of the word ‘krubo’ (3lun; dwarf) and the suffix –enkys; Ñombītsor from the noun ‘ñombes’ (4sol; elephant) and the diminutive suffix –ītsos (2sol), in the collective.

Remarks. Related to elephants, but never reaching a large size; used as transportation in Volantis.

Timpa-Kēlior, all the white lions or hrakkars.

Etymology. Timpa from the adjective ‘timpa’ (adj. I; white); Kēlior, from the collective of the noun ‘kēlio’ (3lun; lion).

Remarks. A rare species of white lion, native to the Dothraki Sea.

Dothrakōñe-Anner, all the horses of the Dothraki.

Etymology. Dothrakōñe, from the Dothraki, the horselords, and the suffix –ōñe (adj. II); Anner, from the nominative collective of the word ‘anne’ (4lun; horse).

Remarks. Widespread on the entire world, medium of transportation, and used in combat as well. They are especially important for the Dothraki horselords.

Rizmenkys-Annītsor, all the dwarf horses of the sand or sand steeds.

Etymology. Rizmenkys the word ‘rizmon’ (3ter; sand) and the suffix –enkys (adj. I); Annītsor from the word ‘anne’ (4lun; horse) and the diminutive suffix –ītsos (2sol) in the collective.

Remarks. Long neck, narrow head, slim and swift, with red, golden, black or pale fur. Bred in Dorne.

Starkenka-Zoklar, all the wolves of the Starks or direwolves.

Etymology. Starkenka, from the name of House Stark, whose sigil is the direwolf, and the suffix –enka (adj. I); Zoklar from the nominative collective of the word ‘zokla’ (1lun; wolf).

Remarks. An ancient relative of the common wolf, but much more robust and strong. Absent south of the Wall. However, a dead female direwolf was found south of the Wall; Ned Stark’s children and Jon Snow were allowed to keep and raise the pups (Fig. 3).

Figure 3. The first known occurrences of Starkenka-Zoklar south of the Wall, seen here as two pups of a female direwolf. A typical example of Sylvie-Ēlie-Valar (Jon Snow) for scale. Screen capture from Episode #1 (‘Winter is Coming’), Season #1, of Game of Thrones (HBO, 2011–present).

Qohorōñe-Valyrītsor, all the Little Valyrians from Qohor.

Etymology. Qohorōñe from Qohor and the suffix –ōñe (adj. II); Valyrītsor from the word Valyria and the diminutive suffix –ītsos (2sol) in the collective.

Remarks. Lemur-like primates with silver-white fur and purple eyes, living in the forest of Qohor.

Lannenka-Kēlior, all the lions of the Lannisters.

Etymology. Lannenka from Lann the Clever, founder of House Lannister whose sigil has a golden lion, and the suffix –enka (adj. I); Kēlior, from the collective of the word ‘kēlio’ (3lun; lion).

Ōgharenkys-Ñomber, all the great woolly elephants or mammoths.

Etymology. Ōgharenkys, from the word ‘ōghar’ (1aq; hair) and the suffix –enkys (adj. I); Ñomber, see above.

Remarks. Related to elephants, but more robust, with thick fur and curved tusks, from beyond the Wall. Giants usually ride them.

Sōnōñe-Gryver, all the snow bears.

Etymology. Sōnōñe, from the word ‘sōna’ (1lun; snow) and the suffix –ōñe; Gryver from the collective of the word ‘gryves’ (4sol; bear).

Remarks. Related to the brown bears, but adapted to survive in the cold environments beyond the Wall.

Μēremolrenkys-Epser, all the goats with a single horn or unicorns.

Etymology. Μēremolrenkys from the combination of the words ‘mēre’ (one) and ‘molry’ (2lun; horn) and the suffix –enkys (adj. I); Epser, from the nominative collective of the word ‘epses’ (4sol; goat).

Remarks. Goat-like animals with a single horn, believed to survive in Skagos and on the tall mountains of Ib. This disjointed distribution could be explained by two hypotheses: either they are native to one island and their presence on the other is explained by human interference; or this animal used to be widely distributed in the past (perhaps in times when the sea-level was lower and the two islands were connected to each other or to the mainland), and the present distributions are remnants.

Zōbritimpa-Anner, all the black-and-white horses or zorses.

Etymology. Zōbritimpa from the combination of the words ‘zōbrie’ (adj. III; black), ‘timpa’ (adj. I; white); Anner, from the nominative collective of the word ‘anne’ (4lun; horse).

Remarks. Related to horses, but with black and white stripes; they live in eastern Essos.

Valenka, the group of humans and human-like creatures.

Etymology. From the word ‘vala’ (1lun; man) and the suffix –enka (adj. I), meaning all-together ‘like humans’.

Remarks. This is the group that contains all human-like sentient species. Besides the group of humans, Valar (see below), there are several other species, mythical or not, that are most probably more closely related to the Valar than anything else. Although some of the species mentioned below could be myths and the product of fantasies and stories, I still prefer to properly name them. The distribution of Valenka is shown in Figure 4.

Figure 4. The distribution of known species of Valenka and Valar, the human-like species in the World of Ice and Fire.

Guēsōñe-Riñar, all the children from the forest.

Etymology. Guēsōñe from the word ‘guēsin’ (4lun; forest) and the suffix –ōñe; Riñar from the nominative collective of ‘riña’ (1lun; child).

Remarks. Dark and beautiful, less barbarous than the giants; renowned for working with obsidian and beautiful songs. Currently live beyond the Wall.

Rōvalar-Rōvalar, all the giants.

Etymology. Rōvalar (all the giants) from the nominative collective of ‘rōvala’ (1lun; giant). Both components of the name are identical for emphasis.

Remarks. Giants once had a broader distribution in the World of Ice and Fire, but currently are restricted to the lands north of the Wall.

Hagedornōñe-Annevalar, all the horsemen of Hagedorn, also known as the Centaurs.

Etymology. Hagedornōñe, in honor of the great Archmaester Hagedorn, who wrote that centaurs never existed and were simply mounted warriors; Annevalar, from the combination of the words ‘vala’ (1lun; man) and ‘anne’ (4 lun; horse), meaning horsemen in the nominative collective.

Remarks. Most probably, the specimens examined in the Citadel are artifacts of mixtures of skeletons of humans and horses, probably confused with the Dothraki. Even so, it is still possible, especially in a world of magic like the World of Ice and Fire, that they once existed. Supposed distribution in the eastern grasslands of Essos during the Dawn Age.

Theronōñe-Valītsor, all the little humans of Theron, also known as the Deep Ones.

Etymology. Theronōñe, in honor to Maester Theron who first wrote about these creatures; Valītsor from the word ‘vala’ (1lun; man) and the diminutive suffix –ītsos (2sol) in the nominative collective.

Remarks. Supposedly misshapen creatures that fathered the merlings (see below). Their exact distribution is not known, but reports mention the destruction of the Lorathi mazemakers by sea creatures and the sacrifice of sailors on the Thousand Islands to fish-headed gods, likely connected to the Deep Ones. As such, we can speculate that the Deep Ones had a Shivering Sea distribution.

Klihenka-Valar, all the fish-men, also known as merlings.

Etymology. Klihenka, from ‘klios’ (3sol; fish) and the suffix –enka (adj. I); for Valar, see below.

Remarks. Aquatic human/fish hybrids, with a cosmopolitan distribution. House Manderly has a merling at its sigil.

Guēsōñe-Dekurūptyr, all the walkers of the forest, also known as the Ifeqevron.

Etymology. Guēsōñe (of the forest) from the word ‘guēsin’ (4lun; forest); Dekurūptyr comes from the word ‘dekurūbagon’ (to walk) and the suffix –tys (2sol) to form the word ‘walker’ in the nominative collective.

Remarks. Ifeqevron means, in the Dothraki language, ‘those who walk in the woods’, which served as the inspiration behind the name in High Valyrian. They inhabit the great forest of the Kingdom of Ifeqevron in northern Essos, between Vaes Dothrak and the Ibben Islands.

Valar, the group containing all humans.

Etymology. From the nominative collective of the noun ‘vala’ (1lun; man), meaning ‘all the humans’.

Remarks. Besides the major ethnic groups of Valar described below (the First Men, the Andals, and the Rhoynars), there are other ‘species’ of Valar that deserve their own name, some of them clearly distinct (e.g., the Ibbenese and the Hairy Men) and others probably distinct from Sylvie-Valar, like the Valyrians. In other cases, we do not have enough information to discern if some ethnic groups are truly distinct from those mentioned above. The horselords Dothraki are, of course, the most important example, including the tribes around them (e.g., the Lhazareen, Jogos Nhai, Qathii). As the First Men originate from the grasslands of Essos, and the Andals were also a nomadic group that stretched eastward in Essos, it is likely that the origin of these groups could be found in them. In the absence of convincing evidence, I prefer not to name all these Sylvie-Valar groups for the moment.

Ibbenīha-Valar, all the Ibbenians.

Etymology. Valar, see above; Ibbenīha comes from the combination of the word Ibben, their island of origin, and the suffix –īha (adj. I), which would mean in the Common Tongue ‘Ibbenian’.

Remarks. They are included in their own species of Valar, as they are apparently unable to produce viable offspring with other species of humans.

Ōgharenka-Valar, all the Hairy Men.

Etymology. Valar, see above; Ōgharenka, from the word ‘ōghar’ (1aq; hair) and the suffix –enka (adj. I).

Remarks. As the Hairy Men are supposed to be closely related to the Ibbenians, I assume that they represent a distinct species of Valar. Some say that they originated in Ibben and then spread out to Essos, settling in places like Lorath.

Sothorīha-Valar, all the Sothorysians.

Etymology. Valar, see above; Sothorīha comes from the combination of the word Sothoryos, their island of origin, and the suffix –īha (adj. I), which would mean in the Common Tongue ‘Sothorysian’.

Remarks. As the humans from Sothoryos, or Brindled Men, were unable to produce viable offspring with other species of humans, I suppose that they represent a distinct species of Valar.

Jaedrōñe-Valar, all the humans from the Summer Islands.

Etymology. Jaedrōñe comes from the word ‘jaedria’ (Summer Islands; 1aq.), and the suffix –ōñe, in allusion to the Summer Islands, their place of origin; Valar, see above.

Remarks. They are included in their own species of Valar, as they, throughout their history, apparently lived isolated from the rest.

Sylvie-Valar, all the wise humans.

Etymology. Sylvie, from the nominative singular of the adjective ‘sylvie’ (adj. III; wise);  Valar see above.

Remarks. The First Men, the Andals and Rhoynars represent the three major ethnic groups in the World of Ice and Fire and we have evidence of their interbreeding producing viable offspring. As such, I include them in the same ‘species’, with different ‘subspecies’.

Sylvie-Ēlie-Valar, all the wise First Men.

Etymology. Ēlie comes from the adjective ‘ēlie’ (adj. III; first, primary).

Sylvie-Andalōñe-Valar, all the wise Andals.

Etymology. Andalōñe comes from the word for the Andals and the suffix –ōñe (adj. II).

Sylvie-Rhoynarīha-Valar, all the wise Rhoynarians.

Etymology. Rhoynarīha comes from Rhoynar and the suffix –īha (adj. I), denoting their place of origin.

Sylvie-Valyrīha-Valar, all the wise Valyrians.

Etymology. Valyrīha comes from Valyria and the suffix –īha (adj. I), denoting their place of origin. 

Sylvie-Timpa-Valar, all the wise white humans.

Etymology. Timpa comes from the adjective ‘timpa’ (adj. I; white).

Remarks. Although their origin remains unclear, they probably represent a variation of the First Men. As such, they are tentatively included in the same ‘species’, but in a different ‘subspecies’ (Fig. 5).

Figure 5. A typical specimen of Sylvie-Timpa-Valar, a white walker from beyond the Wall, from the Lands of Always Winter. Screen capture from Episode #8 (‘Hardhome’), Season #5, of Game of Thrones (HBO, 2011–present).

Hontenka, the group that contains all the birds.

Etymology. Comes from the stem of the nominative collective of the word ‘hontes’ (4sol; bird) and the suffix –enka (adj. I).

Remarks. This group contains all birds. Note that birds are not defined by their flight ability, which was developed independently in other groups, such as dragons and insects.

Bantenka-Lārar, all the crows of the night.

Etymology. Bantenka, from the word bantis (5sol; night) in honor of the Night’s Watch, whose members are called ‘crows’, and the suffix –enka; Lārar, from the collective of ‘lāra’ (1lun; crow).

Remarks. Iconic birds, mainly because of their association with the Night’s Watch.

Hontenkys-Dāryr, all the birds of the king, also known as the Eagle.

Etymology. Hontenkys, from the word ‘hontes’ (4sol; bird) and the suffix –enkys (adj. I); Dāryr, from the collective of the word dārys (2sol; king).

Udrenkys-Vōljer, all the ravens.

Etymology. Udrenkys, from the word ‘udir’ (5aq; word, news) and the suffix –enkys (adj. I); Vōljer, from the collective of the word ‘vōljes’ (4sol; raven).

Remarks. One of the animals with special importance to humans, as they are used in long-distance communication between settlements. They are usually under the care of the Maester of each castle.

Sōnenkys-Vōljer, all the ravens of the winter, also known as the White Ravens.

Etymology. Sōnenkys from the word ‘sōnar’ (1lun; winter) and the suffix –enkys (adj. I), in allusion to their use by the Maesters of the Citadel to announce the change of seasons; Vōljer, from the collective of the word vōljes (4sol; raven).

Remarks. A different species of raven, kept and raised in the Citadel. They are used to announce the changing of seasons in Westeros.

Sōnenkor-Vāedar, the song of the snow, also known as the Snow Shrike.

Etymology. Sōnenkor, from the word ‘sōna’ (1lun; snow) with the suffix –enkor (adj. I); Vāedar, from the nominative of the word ‘vāedar’ (1aq; song).

Remarks. Found mainly in the North, but go as south as the Riverlands.

Tīkunītsenka, the small winged animals.

Etymology. From ‘tīkun’ (3sol; wing) and the suffixes –ītsos (2 sol; diminutive) and –enka (adj. I).

Ānogro-Bībire-Zōbros, the purple, blood-sucking animal, or bloodfly.

Etymology. Ānogro, from the word ‘ānogar’ (1aq; blood) in the genitive; Bībire, from the verb ‘bībagon’ (to suck); Zōbros, from the substantive of the word ‘zōbrie’ (adj. III; purple). The name means the “bloodsucking purple one”.

Remarks. Bloodsucking, purple insect, living in marshes and ponds in Essos.

Kastys-Raeder, all the green scorpions, or manticores.

Etymology. Kastys, from the adjective ‘kasta’ (adj. I; blue, green), in allusion to the Jade Sea where this creature lives; Raeder, from the nominative collective of the noun ‘raedes’ (4sol; scorpion).

Remarks. They have a black carapace, a barbed tail, and a human-like face. Its sting is poisonous and causes heart attack in humans. They live in the islands of the Jade Sea.

Rīzenka, the group of reptile-like animals.

Etymology. From the word ‘rīza’ (1lun; reptile, lizard) and the suffix –enka.

Basiliskīha-Rīzar, all the Basiliskian reptiles.

Etymology. Basiliskīha, from Basilisk and the suffix –īha (adj. I), meaning “Basiliskian”; Rīzar from the collective of the noun ‘rīza’ (1lun; reptile, lizard).

Remarks. The basilisk is a venomous, large, reptile from the Basilisk Isles.

Drakarenkys-Zaldrīzer, all the fire dragons.

Etymology. Drakarenkys, from the word ‘drakarys’ (2sol; dragon-fire) and the suffix –enkys (adj. I); Zaldrīzer, from the nominative collective of the word ‘zaldrīzes’ (4sol; dragon).

Remarks. These magical creatures once lived in the entire World of Ice and Fire, with four limbs, two wings, strong jaws, sharp teeth and claws, horns, and a long pointed tail (Fig. 6); they breathe fire. Once the source of power for the Valyrian dragonlords and the Targaryens, they were considered extinct since the last dragon died in the 153 AC (After Conquest) following the events of the Dance of the Dragons. However, Daenerys Targaryen was recently able to hatch three dragon eggs.

Figure 6. Drogon, named after Khal Drogo, one of the two surviving Drakarenkys-Zaldrīzer, seen in the dragon pit of King’s Landing. Screen capture from Episode #7 (‘The Dragon and the Wolf’), Season #7, of Game of Thrones (HBO, 2011–present).

Suvenkys-Zaldrīzer, all the ice dragons.

Etymology. Suvenkys, from word ‘suvion’ (3ter; ice) and the suffix –enkys (adj. I); Zaldrīzer, see above.

Remarks. A mythical species of dragon that was larger than the fire dragons and breathed ice (Fig. 7). Rumor has it that the Night King was able to create a Suvenkys-Zaldrīzer beyond the Wall.

Figure 7. Viserion, named after Viserys Targaryen (brother of Daenerys Targaryen), the only known specimen of Suvenkys-Zaldrīzer in the World of Ice and Fire. Although seemingly identical to a Drakarenkys-Zaldrīzer, there is clear evidence that this species does not breathe fire. Scholars disagree if a Suvenkys-Zaldrīzer breaths ice or some kind of ‘icy fire’. Screen capture from Episode #7 (‘The Dragon and the Wolf’), Season #7, of Game of Thrones (HBO, 2011–present).

Tīkunoqittys-Zaldrīzer, all the dragons without wings, or firewyrms.

Etymology. Tīkunoqittys, from the nominative plural of the word ‘tīkun’ (3sol; wing) with the suffix –oqittys (adj. I; –less); Zaldrizer, see above.

Remarks. Wingless fire dragons from the Valyrian peninsula. Extinct.

Drakaroqittys-Zaldrīzer, all the fireless dragons, or wyverns.

Etymology. Drakaroqittys, from the word drakarys (2sol; dragon-fire) and the suffix –oqittys (adj. I; less); Zaldrīzer, see above.

Remarks. Related to dragons but fireless, surviving in Sothyryos.

Rīdōñe-Rīskelior, all the lizard-lions of the Reeds.

Etymology. Rīdōñe, meaning ‘of the Reed’, in honor to House Reed, whose sigil has a black lizard-lion, and the suffix –ōñe (adj. II); Rīskelior, from the word ‘rīza’ (1lun; reptile, lizard) and the word ‘kēlio’ (3lun; lion) in the collective.

Remarks. Crocodile-like lizards with large teeth that live in the streams and swamps of the Neck.

Qarthōñor-Qintrir, all the turtles of Qarth, or phantom tortoises.

Etymology. Qarthōñor, from the city of Qarth and the suffix –ōñe (adj. II); Qintrir, from the nominative col of the noun ‘qintir’ (5aq; turtle).

Tegōñior-Qintrir, all the terrestrial turtles.

Embōñior-Qintrir, all the marine turtles.

Qelbōñior-Qintrir, all the aquatic turtles.

Etymology. The first components are formed from the adjectives ‘tegōñe’ (adj. II; terrestrial), ‘embōñe’ (adj. II; marine), and ‘qelbōñe’ (adj. II; aquatic, from the river); Qintrir, see above.

Remarks. Reptile-like animals, whose body is enclosed within a bony shell; they can reach large sizes and have a cosmopolitan distribution. Although probably there are dozens of different species of turtles in the World of Ice and Fire, they are grouped here under three species only, based on their preferred habitat. Further work should focus on describing the various species of turtles included in each of these above-named groups.

Martino-Qintrir, the turtle of Martin, also known as the Old Man of the River.

Etymology. Martino, genitive of Martin, in honor of G.R.R. Martin, the author of the Song of Ice and Fire series; Qintrir, see above.

Remarks. The Old Man of the River is a sacred giant turtle that lived in the river Rhoyne, and is worshiped by the Rhoynars. G.R.R. Martin has publicly expressed his love of turtles and the role that they played in the development of the World of Ice and Fire[2], so this species is named after him.

Embenka, all the sea-dwelling animals.

Etymology. From the noun ‘embar’ (1aq; sea) and the suffix –enka (adj. I).

Grējojōñor-Uēhor, all the great squids of the Greyjoys, or krakens.

Etymology. Grējojōñor, in allusion to House Greyjoy, whose sigil bears a golden kraken, with the suffix –ōñe (adj. II); Uēhor, from the word ‘uēs’ (3sol; squid) in the nominative collective.

Remarks. A kind of giant squid, supposedly living in the sea south of Dorne.

Embrōñe-Jēnqañōgher, all the sea creatures with eight arms, also known as octopods.

Etymology. Embrōñe, from the genitive collective of the word ‘embar’ (1aq; sea) with the suffix –ōñe (adj. II); Jēnqañōgher, from the combination of the words ‘jēnqa’ (eight) and ‘ñōghe’ (4lun; arm) in the collective.

Qaedrāzmar-Qaedrāzmar, all the great whales, or leviathans.

Etymology. Qaedrāzmar, from the word ‘qaedar’ (1aq; whale) and the augmentative suffix –āzma (1lun) in the collective.

Remarks. An enormous grey whale, among the most ancient creatures of the World of Ice and Fire. Found in the Shivering Sea.

Naggōñe-Embrōñe-Zaldrīzer, all the sea dragons of Nagga.

Etymology. Naggōñe, of Nagga, the mythical sea dragon, with the suffix –ōñe (adj. II); Embrōñe, from the word ‘embar’ (1aq; sea) and the suffix –ōñe (adj. II); Zaldrīzer, see above.

Remarks. A sea dragon, feeding on krakens and leviathans. Supposedly extinct since the Age of Heroes, although some believe it still survives in the Sunset Sea.


This is only the first account on the names of some of the most important animals of the World of Ice and Fire. Many more kinds of beings remain lacking formal names, including most domesticated animals and plants. Future work should focus on refining this system of taxonomy and describing the remarkable living and extinct diversity of Westeros and Essos.


Dothraki Wiki (Tongues of Ice and Fire Wiki). (2018) Learning High Valyrian. Available from: (Date of access: 27/Apr/2018).

E_v_a_n. (2017) The Full Taxonomy of Ice and Fire. Subreddit “A Song of Ice and Fire”. Available from: (Date of access: 27/Apr/2018).

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

International Commission on Zoological Nomenclature (ICZN). (2012) Amendment of Articles 8, 9, 10, 21 and 78 of the International Code of Zoological Nomenclature to expand and refine methods of publication. ZooKeys 219: 1–10.

Martin, G.R.R. (1996) A Game of Thrones. Bantam Books, New York.

Martin, G.R.R. (1999) A Clash of Kings. Bantam Books, New York.

Martin, G.R.R. (2000) A Storm of Swords. Bantam Books, New York.

Martin, G.R.R. (2005) A Feast for Crows. Bantam Books, New York.

Martin, G.R.R. (2011) A Dance with Dragons. Bantam Books, New York.

Martin, G.R.R.; Garcia, E.; Antonsson, L. (2014) The World of Ice and Fire: the Untold History of Westeros and the Game of Thrones. Bantam Books, New York.

Peterson, D.J. (2013) Valar Dohaeris. Dothraki: a Language of Fire and Blood. Available from: ris/ (Date of access: 27/Apr/2018).

Wiki of Ice and Fire, A. (2018) Bestiary. Available from: Bestiary (Date of access: 27/Apr/2018).


I would like to thank the Dothraki Wiki community for making available the rules, grammar and dictionary of High Valyrian. I thank the Reddit communities of the Song of Ice and Fire and Game of Thrones for inspiration and comments. Special thanks to the redditors u/hm0119 and u/jackm0ve for their interest to jump in and name some species of their own; these names have not been included herein. I would like to deeply thank the editor of the JGS, Rodrigo B. Salvador, and the rest of the editorial board for useful comments that greatly improved this manuscript. I would like to express my gratitude to David J. Peterson, the creator of the Valyrian and Dothraki languages, who reviewed an earlier version of the manuscript; he managed not only to point out the numerous mistakes I made in the formation of the words in my early version but also to provide valuable lessons through his critical review. His comments and suggestions also made the entire system much more consistent and uniform. Of course, I am solely responsible for any mistakes in the formation of the High Valyrian names. This project has been developed in my free time, but was inspired by the importance of zoological nomenclature and the art of coining species names. I would like to thank my family for their understanding and support when I spend time with projects like this.


Evangelos Vlachos is a big fan of the World of Ice and Fire and, just like G.R.R. Martin, a huge fan of turtles and tortoises. He is currently a CONICET researcher in the Museo Paleontológico Egidio Feruglio, in Trelew, Chubut, Argentina, working on fossil turtles and tortoises.

[1] See the ICZN’s website ( for detailed information.

[2] From ing/game-of-thrones-turtles/

Check other articles from this volume


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: (Date of access: 11/Jan/2018).

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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


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

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

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

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

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


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


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

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

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

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

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

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

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

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

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

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

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The plants of J.R.R. Tolkien’s legendarium

Walter S. Judd1,2 & Graham A. Judd3

1Department of Biology, University of Florida. Gainesville, FL, U.S.A.

2Florida Museum of Natural History. Gainesville, FL, U.S.A.

3Independent researcher. South Saint Paul, MN, U.S.A.

Emails: lyonia (at) ufl (dot) edu; gjudd (at) me (dot) com

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Many readers of The Hobbit or The Lord of the Rings believe that the events of these books occur in an imaginary world and thus have no connection with the world around us. However, John Ronald Reuel Tolkien sought to correct this misconception, stating that Middle-earth “is just the use of Middle English middle-erde (or erthe), altered from Old English Middangeard: the name for the inhabited lands of Men ‘between the seas.’” He went on to say that “imaginatively this ‘history’ is supposed to take place in a period of the actual Old World of this planet” (Tolkien, 1981, Letter No. 165). His writings should not be considered escapist, but instead are meant to reconnect us to important elements of our internal and cultural landscape. They should also influence how we interact with other individuals and with the world in which we live — including the landscapes of our natural environment — and especially plants! The importance of plants in the Tolkien’s Middle-earth is thus considered in detail in our book, Flora of Middle-Earth: Plants of J.R.R. Tolkien’s Legendarium, recently published by Oxford University Press (Judd & Judd, 2017), which we introduce here, along with an introduction to the importance of plants in connection with Tolkien’s imaginative world.

Flora of Middle-Earth (book cover, showing forest of Lothlórien)

Our book focuses on one of the major components of our environment — the Green Plants — organisms to which many in our modern, highly technological world have become blind (Wandersee & Schussler, 2001; Allen, 2003). Indeed, some have argued that we are now disconnected from the entire natural world (Yoon, 2009). Plants are ecologically diverse and range dramatically in size — from microscopic, aquatic, green algae to the tallest flowering trees or conifers. They are critically important in maintaining a healthy biosphere — and in fact, without plants, animal (and, of course, human) life would be impossible. They provide our food, construction materials for our homes, add beauty to our surroundings, and even provide the air we breathe. In Tolkien’s legendarium, plants are the primary concern of Yavanna Kementári, the Giver of Fruits and wife of Aulë, who has lordship over all the substances of which the Earth is made. As related in The Silmarillion, she is the “lover of all things that grow in the earth, and all their countless forms she holds in her mind, from the trees like towers in forests … to the moss upon stones or the small and secret things in the mould” (Valaquenta: p. 27). Understandably, she is held in great reverence by the elves, as are the natural environments she oversees. We believe Tolkien’s reference was comparable.

Tolkien’s descriptions of Middle-earth are richly detailed, including succinct verbal sketches of many of its plants, and thus create a realistic stage for his dramas. His detailed treatment of plants plays a major role in the creation of this stage — providing the distinctive landscapes and natural locales of Middle-earth — from the tundra and ice-fields of the north, to the extensive prairies of Rohan, and the coniferous forests of Dorthonion, as well as the broad-leaved forests of Doriath or Fangorn and wetlands such as the Gladden Fields. The dominant species within each plant community are always mentioned, especially the trees, which Tolkien, like Yavanna, held most dear (see The Silmarillion: chapter 2). Thus, it is critical for our appreciation and understanding of Middle-earth to envision these scenes accurately. These plants, however, do more than merely provide descriptive detail, enhancing the veracity of the tales of Middle-earth. The plants within Tolkien’s legendarium are actually part of the story, and in ways that are more deeply significant than merely evident in the actions of Ents — anthropomorphized trees — that “speak on behalf of all things that have roots, and punish those that wrong them” (The Silmarillion: p. 45). Their significance can be seen in the numerous connections between plants and important individuals in the myths and history of Middle-earth. For example, in the First Age (and earlier), how are we to understand the Two Trees of Valinor, fashioned by Yavanna, and why is it important that Thingol, the elven ruler of Doriath, was called the king of beech, oak, and elm? Why was his daughter, Lúthien, when first observed by Beren, dancing among the hemlock-umbels under the beeches of Neldoreth? And what is the link between her feet and the leaves of lindens? Why did hawthorns obscure the entrance to the Hidden Kingdom of Gondolin? During the Second Age, why did the elves give Aldarion, soon to become the sixth king of Númenor, a White Tree — Nimloth — and what is the connection between this tree and the White Trees of Gondor? Why did the elves bring to Númenor several different fragrant trees from Eressëa — and what did these trees look like?  In the Third Age, how was pipe-weed integral to the culture of the Shire, and why was athelas (kingsfoil) useful in the hands of the king of Gondor? How did these two herbs get to Middle-earth? What is the connection of willows and the Withywindle valley (in the Old Forest), and should willows, therefore, be viewed negatively? Why does Quickbeam love rowan-trees, and why were mallorn-trees important to Galadriel and the elves of Lothlórien? What did mallorn-trees look like? And finally, how should we envision the herbs elanor and niphredil, and what made these two plants so sacred to the elves? Of course many additional questions come quickly to mind, and we deal with these in our book.

It is obvious from even a cursory reading of The Lord of the Rings that the book was written by a person who was botanically knowledgeable — but more than that — a writer who really loved plants! (In fact his introduction to the world of plants occurred very early in his life when he was taught botany by his mother.) But we don’t need to merely accept this from our interpretations of his writings. Tolkien tells us of his appreciation of plants. He said in his letter to the Houghton Mifflin Co.: “I am (obviously) much in love with plants and above all trees, and always have been; and I find human maltreatment of them as hard to bear as some find ill-treatment of animals” (Tolkien, 1981: Letter No. 164). We agree: his love of plants is obvious, and it is apparent on nearly every page of The Hobbit or The Lord of the Rings. Only a writer whose eyes were open to the diversity of the natural world could have accomplished such a task — closely integrating plants into his imagined world, and, as a result, including nearly all the trees of England (and also most European trees) within the Middle-earth of the First through the Third Ages. Because the species of trees (as well as shrubs and herbs) growing in England and other European regions are for the most part members of widely distributed genera that also occur in temperate North America and Asia, especially eastern and southeastern Asia, we can find the plants of Tolkien’s Middle-earth in the forests and fields around our homes. Thus, a major goal of this book, in addition to increasing our appreciation of the imagined landscapes of Middle-earth, is to increase our respect for and understanding of the plants that grow in the natural environments that exist around us. Tolkien appreciated the beauty and diversity of the natural world, and its destruction through urbanization and industrialization angered him (unfortunately, modern followers of Saruman are not hard to find!). Thus, one of our goals is to increase the visibility of and love for plants in our modern culture. And, taking the Ents (i.e., sentient trees, indwelt by spirits “summoned from afar”; The Silmarillion: p. 45) as our role-models, we hope to foster the desire to protect the forests and meadows near our homes (and across the world). Finally, the wild plants of forest and field are not our only concern. In our book we have also described the cultivated plants of vegetable and flower gardens as well as agricultural fields, addressing the interesting and long history of plants and people (or hobbits and elves!). We should appreciate not only wild plants (as do the Ents) but also the plants of orchards and cultivated fields (like the Entwives). In the end, the fact that an investigation of the plants of Tolkien’s Middle-earth reconnects us with the plants of our own world should not be surprising. Tolkien, in his essay On Fairy-Stories, said that “Recovery” is one of the goals of fantasy, and by this he meant “a re-gaining — regaining of a clear view” and “seeing things as we are (or were) meant to see them.” Thus, in “experiencing the fantastic, we recover a fresh view of the unfantastic, a view too long dulled by familiarity” (Flieger, 2002: chapter 3).

If the plants of Tolkien’s legendarium are the trees, shrubs, and herbs of our own world, one might ask: What about plants such as elanor, niphredil, alfirin, simbelmynë, mallorn-trees, or the White Tree of Gondor? Are these simply the creation of Tolkien’s imagination, or do they also have links to our own world. The answer, we think, is both — certainly these plants, as Tolkien explained, “are lit by a light that would not be seen ever in a growing plant” (Tolkien, 1981: Letter No. 312) in our world — so they arise, some more and others less, out of his imagination and are used in specific ways in the story in order to clarify aspects of elven, human, or hobbit culture. They are artistic creations, enhancing the wonder and mystery of Tolkien’s imaginative world. But it is also important to keep in mind that perhaps all of the imaginative plants of Middle-earth are based, at least in part, on species of our own world. For example, Tolkien suggested that niphredil — if seen in the light of our world — would be “simply a delicate kin of a snowdrop,” while elanor would be “a pimpernel (perhaps a little enlarged) growing sun-golden flowers and star-silver ones on the same plant” (Tolkien, 1981: Letter No. 312). As early as 1956, Tolkien commented that “Botanists want a more accurate description of the mallorn, of elanor, niphredil, alfirin, mallos, and simbelmynë” (Tolkien, 1981: Letter No. 187), and we trust that many readers today have a similar desire. We have, therefore, done the necessary detective work to connect these imaginative plants with their sources and provide such accurate descriptions. We believe that this botanical knowledge will enrich the experience of those who have read (or are reading) Tolkien’s works. Our book explores the interactions between plants and the speaking-peoples of Middle-earth — such as humans, hobbits, elves, or ents — whether such plants are the common oaks, pines, or grasses found in the sunlight of our world or are those plants lit by a more imaginative light, such as niphredil or elanor. Thus, we attempt in our book to synthesize information from diverse realms: Tolkien’s writings, etymology (the evolution of words), botany and plant systematics (the study of plants and their evolutionary relationships), and artistic endeavors. We hope that Tolkien would approve of our attempt, as he suggested that the gold and silver light of Valinor, pouring from the Two Trees (Telperion and Laurelin), represents the “light of art undivorced from reason, that sees things both scientifically … and imaginatively” (Tolkien, 1981: Letter No. 131).

In the book we provide detailed treatments of the 141 plants of Middle-earth, and for each of the 100 most important plants of Tolkien’s imaginative world, we include (1) the common and scientific names, along with an indication of the family to which the plant  belongs; (2) a brief quote from one of Tolkien’s works in which the plant is referenced; (3) a discussion of the significance of the plant in the context of Tolkien’s legendarium; (4) the etymology, relating to both the English common name and the Latin (or Latinized) scientific name, and where relevant, the name in one or more of the languages of Middle-earth; (5) a brief description of the plant’s geographical distribution and ecology; (6) its economic importance; and (7) a brief description of the plant. Most of these also are provided with a woodcut-style illustration (as an aid to identification), along with an inset illustrating one of the events in the history of Middle-earth in which the plant played a role.

Niphredil (based upon the snowdrop, Galanthus nivalis, in the plant family Amaryllidaceae) with inset (vignette) showing Aragorn and Arwen on Cerin Amroth. Illustration from Flora of Middle-Earth.

It is our goal that the inset illustrations (vignettes) be functional, decorative, and fit visually into the lore of Middle Earth. By abstracting the images with a woodblock aesthetic, Graham, the second author and illustrator, was able to simplify the complexity of the plant pictured, providing a clearer view of the diagnostic features of each plant than a photograph would have offered. In his botanical illustrations, only the information needed to identify each plant is provided, and this same concept inspired his approach to the vignettes and narratives depicted. The tales and lore of Arda have been imagined by all of us, conceived and casted in movies, and depicted by talented and amazing artists. From the Hildebrandt brothers to Cor Blok, these artists and actors have shaded our original conceptions of what these characters, such as Bilbo or Gandalf, look like. Because of this we seek to create an abstracted view, offering silhouettes rife with symbols, pulling heavily on descriptions from the Tolkien’s books to color our conceptualization of these well-fabricated characters. Keeping Tolkien’s concerns in mind, we do not want to infringe on the viewer’s ideation of the characters, but we feel it is very important to provide the framework for people to see the narrative, while still allowing them to project their own conceptualizations onto the image.

Traditionally, when we think of fantasy illustrations, we think of images framed like classic historical paintings or Greek dramas. By focusing on the flora over the fauna, we had to restructure how we approached the composition of each scene. So often plants are only the background that our grand actors stride across, but in contrast, we want to highlight how these narratives played out in the botanically rich and vibrant world that Tolkien imagined. This led Graham to a fundamental restructuring of the composition of each image, so the action or drama of the characters is often deemphasized, with the vignette focusing on how the action would have settled into the environment.

In conclusion, we hope that our book will create a visual reference — and legitimacy — for both the plants growing in our forests, meadows, and marshes, as well as those that we have received as gifts from Tolkien’s imagination.


Allen, W. (2003) Plant blindness. Bioscience 53(10): 926.

Flieger, V. (2002) Splintered Light: Logos and Language in Tolkien’s World. 2nd edition. Kent State University Press, Ohio.

Judd, W.S. & Judd, G.A. (2017) Flora of Middle-Earth: Plants of J. R. R. Tolkien’s Legendarium. Oxford University Press, New York.

Tolkien, J.R.R. (1977) The Silmarillion. George Allen and Unwin, London. [Cited here from the 2nd Edition, 2001, Houghton Mifflin, Boston].

Tolkien, J.R.R. (1981) The Letters of J. R. R. Tolkien, edited by H. Carpenter, with the assistance of C. Tolkien. George Allen and Unwin, London / Houghton Mifflin, Boston.

Wandersee, J. & Schussler, E. (2001) Towards a theory of plant blindness. Plant Science Bulletin 47(1): 2–9.

Yoon, C.K. (2009) Naming Nature: The Clash between Instinct and Science. W.W. Norton & Co., New York.


Walter S. Judd is a Distinguished Professor Emeritus in the Department of Biology, University of Florida and also has an affiliate appointment in the Florida Museum of Natural History. His research focuses on the systematics and evolution of the flowering plants. He has published over 230 refereed articles and has described numerous new species of plants. He is also a co-author of the textbook, Plant Systematics: A Phylogenetic Approach.

Graham A. Judd has a MFA in Printmaking and received a Jerome Foundation Fellowship for Emerging Printmakers at Highpoint Center for Printmaking. He currently teaches at Augsberg College and Minneapolis College of Art and Design.

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Arthropod diversity in Pokémon

André W. Prado* & Thiago F. A. Almeida

Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil.

*Email: awp03 (at) hotmail (dot) com

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Like most regular children in the 2000’s, we were obsessed with Pokémon games and anime series. The experience of exploring new environments, discovering new creatures and collecting them, always fascinated us. Maybe this was a sign of what we would become: zoologists. During college, as we got to know ever more about animal biodiversity, we could not help but notice several similarities between Pokémon and real animals. Today, as an arachnologist and an entomologist, and still Pokémon fans, our interest in arthropods and admiration for this franchise were the main motivations for this study.


Animal diversity has always been debated and represented in different types of media. Since the Pleistocene, humans depict animals in their paintings (Aubert et al., 2014), likely reflecting an age-old fascination with nature that still endures. Or, as E. O. Wilson puts it in his “biophilia hypothesis”: “humans have an innate desire to catalog, understand, and spend time with other life-forms” (Wilson, 1984). Given this, studies relating Zoology and culture, especially pop culture, are becoming more and more common recently. Just to name some examples including arthropods, Coelho (2000, 2004) studied insect references in lyrics and cover art of rock music albums, Castanheira et al. (2015) analyzed the representation of arthropods in cinematographic productions, Salvador (2016) studied the biology of giant centipedes in the Gears of War game franchise, and Da-Silva & Campos (2017) analyzed the representation of ants in the Ant-Man movie. There are even some science outreach works about the Pokémon franchise as the analysis of the ichthyological diversity in the Pokémon world (Mendes et al., 2017) and the study of the group of birds popularly called “robins” represented in the game (Tomotani, 2014).


Arthropods correspond to the largest part of the known biotic diversity in the world, counting with over 80% of animal diversity (Zhang, 2011a). With lots of morphological variation, the phylum Arthropoda is divided into five subphyla: Trilobitomorpha (the trilobites, now extinct); Chelicerata (arachnids, horseshoe crabs, and others); Crustacea (shrimps, lobsters, crabs, barnacles and woodlice); Hexapoda (insects) and Myriapoda (centipedes and millipedes). With a high biomass, terrestrial arthropods can be easily seen in a variety of environments, and their presence affects us in several ways.

Although arthropods can inspire fear as venomous creatures or disease vectors, actually most of them are either harmless or important for our own well-being and survival. For instance, many groups of insects are extremely important pollinators and without them, agriculture would collapse. Moreover, terrestrial arthropods have a considerable role as bioindicators for assessing environmental quality (Andersen, 1990; Brown, 1997; Fischer, 2000; Ferrier et al., 2004) and some even have remarkable medicinal uses (Kumar et al., 2015).


The word “Pokémon” is a contraction from the Japanese “Pocket Monsters” (ポケモン). The idea consists in fictional creatures – the eponymous Pokémon – that humans can capture and train to do all sorts of chores, the main one of which is fighting each other. Created by Satoshi Tajiri, Pokémon was originally a game released in 1996, but its tremendous success soon spawned an anime series, mangas, animated movies, a card game, and countless ”goodies” (toys, accessories, clothing, candies, etc.). Developed by Game Freak and published by Nintendo, today Pokémon is one of the most successful game franchises in history, with more than 270 million of overall game copies sold around the world (The Pokémon Company, 2017).

The anime series was released in 1997 and was an instant success with kids, remaining so to this day. Many episodes have an environmental tone, showing how humans can affect the habitats and biodiversity of Pokémon, and emphasizing the importance of collecting for species preservation (Bainbridge, 2013). As a game franchise, Pokémon reached mainly teenagers, which remains a loyal customer base to this day. Today, the games are in their seventh generation (“Gen VII”) and each generation adds a new territory to be explored and several new creatures to be caught. As of now, there are 802 creatures, but some new ones have already been announced for the second game of Gen VII.

The creator of Pokémon, Satoshi Tajiri, loved to collect bugs when he was young, which likely influenced his creation. The Pokémon are mostly inspired by animals and plants and some of them have particular features that can be related to certain real species. In this way, Pokémon biodiversity can be seen as a virtual sample of natural biodiversity.


The main objective of this study is to survey all Pokémon inspired by arthropods, up to Gen VII, and conduct a comparative biological classification of them until the taxonomic level of “Order”, if possible[1]. Considering the Pokémon world as a simulation of our own natural world, we also investigate if the different arthropod groups have the same real-world representativeness in Pokémon. This can be done by analyzing the proportion of species of each group.


The sources of information used for this study are: Bulbapedia (https://bulbapedia. and The Official Pokémon Website ( The Pokémon were classified by Type, Generation, and by their respective taxonomic levels in real-world Biology: Phylum, Subphylum, Class and Order.

The classification into real-world taxonomic levels was made by analyzing morphological and behavioral characters present in the Pokémon species, and comparing them to the relevant animal groups (Fig. 1). Morphological characters were obtained by observing official illustrations and game models. Behavioral characters were obtained from the Pokédex entries of each Pokémon species. Some Pokémon species presented arthropod’s features that were too imprecise to be related to a certain subphyla or order, or their design included features from more than one group of arthropods (for instance, Venonat and Whirlipede). In these cases, the species were marked as “undetermined Subphylum/Order”; regardless, we always classified them to the most accurate level possible.

The biodiversity data used for comparison to the natural world were retrieved from Zhang (2011b).

Figure 1. Arthropod-like Pokémon and the real-world species that inspired them. A. Caterpie. B. Papilio xuthus Linnaeus, 1767. C. Binacle. D. Lepas anatifera Linnaeus, 1758. E. Kabuto. F. Tachypleus gigas (Müller, 1785). G. Anorith. H. Anomalocaris sp. I–J. Kabutops. K. Dimeropyge speyeri Chatterton, 1994. Images A, C, E, G, I, J are official artwork from Pokémon games (extracted from Bulbapedia); images B, D, F extracted from Wikimedia Commons; image H reproduced from Collins (1996); image K reproduced from Chatterton (1994).


We found a total of 91 Pokémon species inspired by arthropods, representing 11.3% of all Pokémon creatures. Most of them (19) belongs to Gen III, corresponding to 14.1% of the total in this generation (Fig. 2, Table 1).

Figure 2. Proportion of Pokémon inspired by arthropods (red) compared to the other monsters (dark grey) from each generation of the game. Total number of Pokémon per generation is shown above each bar.
Table 1. Pokémon inspired by arthropods, with their Pokédex number, Generation, Type(s) and their pertinent biological classification. Horizontal lines separate the game Generations. Symbols: *Wormadam secondary Type might be Steel, Grass or Ground; †fossil group; “???” indicates an undetermined taxonomic position.


Most of the Pokémon species could be classified into the four main living subphyla of Arthropoda: Hexapoda (Figs. 3A–H), Crustacea (Figs. 3I–M), Chelicerata (Figs. 3N–R) and Myriapoda (Figs. 3S–U). The three exceptions were: Kabutops, Anorith and Armaldo (Figs. 3V–X). The former was allocated to the entirely fossil subphylum Trilobitomorpha. The latter two were allocated into another fossil group, with an uncertain position inside Arthropoda (or even an external group, according to some researchers). They belong to the Class Dinocaridida, Order Radiodonta (this ranking is still highly debated, though) and are popularly known as “terror shrimps”.

The Arthropoda subphylum that inspired most of the Pokémon species was Hexapoda, with 62 pokémon, followed by Crustacea (12), Chelicerata (11) and Myriapoda (3) (Figs. 4–5).

The taxonomical order that inspired most of the arthropod Pokémon was Lepidoptera, represented by 21 species. This can be explained by the huge visual appeal and beauty of butterflies and moths. This explanation can be also applied to the large number of Pokémon inspired by the order Coleoptera (13 species), the beetles, animals with an astounding variation of colors and shape. The third order in diversity is Decapoda (10 species), represented by crabs and shrimps.

Figure 3. Examples of Pokémon inspired by arthropods, separated according to subphyla. A–H. Hexapoda: A–B. Hymenoptera (Beedrill, Durant); C. Coleoptera (Ledyba); D. Odonata (Yanma); E. Phasmatodea (Leavanny); F. Hemiptera (Surskit); G. Lepidoptera (Vivillon); H. Mantodea (Scyther). I–M. Crustacea: I–L. Decapoda (Dweeble, Clauncher, Krabby, Corphish); M. Pedunculata (Binacle). N–R. Chelicerata: N–O. Araneae (Spinarak, Galvantula); P–Q. Scorpiones (Gligar, Drapion); R. Xiphosura (Kabuto). S–U. Myriapoda: S–T. Chilopoda (Venipede, Scolipede); U. undetermined order (Whirlipede). V–X. extinct taxa: V. Proetida (Kabutops); W–X. Radiodonta (Anorith, Armaldo). The illustrations are official artwork from the games; images were extracted from Bulbapedia.
Figure 4. Representativeness (in proportion) of Pokémon species inspired by each Arthropoda subphylum. *Dinocaridida is usually considered a class, with uncertain position in Arthropoda.
Figure 5. Number of Pokémon species inspired by each order inside each subphylum of Arthropoda. *Dinocaridida is usually considered a class, with uncertain position in Arthropoda. “???” indicates an undetermined order.


The large number of Pokémon inspired by Hexapoda is congruent with the high diversity of this group in the natural world (Table 2). The fact that there was more Pokémon inspired in Crustacea (Table 3) than in Chelicerata (Table 4) is at odds with natural diversity, but can be related to the very frequent contact that Japanese people have with aquatic animals, which are one of the country’s main food sources (Ashkenazi & Jacob, 2003). The few specimens of Myriapoda in the game are proportionally congruent with their diversity in nature (Table 5).

The comparison between natural and Pokémon diversity shows that the Pokémon world presents higher representativeness of arthropod-like creatures that are more familiar to people or that have a greater visual appeal. The latter is the case of Lepidoptera (Fig. 5), whose diversity in the Pokémon world is much higher than the second place (Coleoptera). However, beetles are the most diverse insect (and overall animal) group in the real world, with approximately 387,000 species, while lepidopterans count “just” with around 157,000 species (Zhang, 2011b). Proportionally, butterflies and moths represent 33.9% of Hexapoda in Pokémon, while in nature this percentage is much closer to that of Coleoptera within Hexapoda (37.6%) rather than the proportion of Lepidoptera (15.3%) (Table 2).

Table 2. Comparison between the diversity of Pokémon species inspired by Hexapoda orders and their respective representativeness in the natural world (Zhang, 2011b).
Table 3. Comparison between the diversity of Pokémon species inspired by Crustacea orders and their respective representativeness in the natural world (Zhang, 2011b).
Table 4. Comparison between the diversity of Pokémon species inspired by Chelicerata orders and their respective representativeness in the natural world (Zhang, 2011b).
Table 5. Comparison between the diversity of Pokémon species inspired by Myriapoda orders and their respective representativeness in the natural world (Zhang, 2011b).


The large number of Pokémon inspired by arthropods indicates that this group, even though not as charismatic as mammalians or birds, still plays an important role in pop culture. The visual appeal and the everyday contact seems to be important aspects that ensure a higher diversity to certain arthropod-like groups in Pokémon. Nevertheless, the Pokémon world still seems to be a good virtual sample of the natural world and this kind of representation can be an interesting source for educational purposes, helping young people to know other type of animals that they do not usually have much contact with, including extinct species.


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André Prado has a bachelor’s degree in Biological Sciences by UFRJ (Rio de Janeiro) and a master’s degree in Zoology by Museu Nacional (Rio de Janeiro). He is a great enthusiast of Cultural Zoology, studying especially the role of animals in cinema.

Thiago Avelar has a licentiate degree in Biological Sciences by UFRJ (Rio de Janeiro) and is currently a high school teacher (Colégio e Curso Miguel Couto, Rio de Janeiro). He was a Fairy Type Elite Four in the extinct Pokémon League Brazil. 

[1] Biological classification organizes species into groups. From the largest to the smallest group: Domain, Kingdom, Phylum, Class, Order, Family, Genus, Species. Sometimes subcategories can exist inside one of these, like a “Subphylum” or “Subspecies”.

Check other articles from this volume


Making a vampire

Veronika N. Laine

Netherlands Institute of Ecology (NIOO-KNAW). Wageningen, The Netherlands.

Email: veronika.laine (at) gmail (dot) com

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The modern vampire is often portrayed as a human transformed into a vampire due to a bloodthirsty spirit[1], demons[2], viruses and other pathogens[3], magic or some unknown reason[4].  Neither fiction nor more realistic accounts have shed light on the precise molecular mechanisms of how the transformation happens until the novel trilogy and TV series called The Strain (Fig. 1) introduced some ways as to how the transformation could happen. In The Strain, parasitic worms carry a virus that causes the vampiric changes to happen through a modification in the expression of genes. This change even creates new organs such as the stinger.

Figure 1. Promotional poster of The Strain TV series, directed by Guillermo del Toro and Chuck Hogan. Image retrieved from: IMP Awards (http://www.impawards. com/).

For obvious reasons, no actual experimental studies have been conducted with vampires and so the exact origin and evolution of vampirism remains unknown. A full genome-wide association study or transcriptome analysis would be preferred to recognize the exact genes behind the vampiric traits, but getting enough samples from vampires will most likely be difficult. Thus, the “candidate gene” approach might be the best method for reaching some conclusions or, if there is enough material, a whole genome sequencing and comparison to human genomes.

In this article I will explore some thoughts on how we could make a vampire in the lab and which part of the genome we would need to alter in order to see the necessary changes. Imagine if genetic engineering would be so advanced that when you tweak little bits of the human genome here and there, you could make whatever traits, even vampiric ones, appear (or disappear) any way you like. Unfortunately, reality is seldom as easy, as it has been shown in movies such as Gattaca (Columbia Pictures, 1997), Splice (Warner Bros., 2009) and the X-Men series (20th Century Fox, 2000–2017), although the genome editing method CRISPR (Cong et al., 2013; Hsu et al., 2014) has lifted genomic modification to a completely new level and has already been used in removing diseases in humans (Ma et al., 2017). Alternatively, what if vampires already existed and we could get our hands on their genome sequence? Which genes would be affected by the transformation? Intriguingly, there are real life examples of species and conditions that could be thought of as vampiric and we can find potential candidate genes for vampirism from these traits. These “vampire building blocks” could then be used in constructing a lab vampire (at least hypothetically).

The myth of vampires has been around for thousands of years and the descriptions of vampirism vary between times and cultures. The vampires we know today date back to the 17th century and they have been covered by every platform in our popular culture. A good summary of the evolution of vampire myths can be found in Harris (2001).

The exact way in which humans transform into vampires depends on the source of the story you are reading and it often remains a mystery. In the extensive study of the science of vampirism, Dr. Pecos and Dr. Lomax (2001–2017) from the late Federal Vampire & Zombie Agency (FVZA) suspected that it is a human vampirism virus (HVV) that causes the transformation. The origin of the virus is suspected to be the vampire bats and their fleas, which sounds very plausible since bats are known to be carriers of many diseases such SARS, ebola and rabies (Biek et al., 2006; Smith & Wang, 2013), and it was also suggested in the movies Daybreakers and the Underworld series. Furthermore, rabies has been suggested to be the actual origin of the modern vampire myth (Gomez-Alonso, 1998).

In this article, I will present real life examples of vampiric traits and hypothesize possible molecular mechanisms and candidate genes that could be mutated after the transformation. I will concentrate on the following three vampiric traits that are common to many descriptions of vampires:

  1. Hematophagy (that is, feeding on blood)
  2. Immortality
  3. Sunlight avoidance


For many people, bloodsucking is the first vampiric trait that comes to mind. Blood is a nutritious fluid tissue, full of proteins and lipids and it is easy to consume. In nature, blood consumption has evolved in several unrelated species throughout the animal kingdom. Among invertebrates, leeches, mosquitos and fleas are the best known examples, and some fish (lampreys) are also known to feed on blood. There are several bird species that practice hematophagy, such as the oxpeckers, hood mockingbirds and vampire finches. Among mammals, the best known hematophagic species are the vampire bats.

Several changes in the genome are needed in order for animals to survive exclusively on blood. One of the key features is to prevent the victim’s blood from coagulating while feeding. In vampire bats the plasminogen activator (PA) genes have gone through gene duplication, domain loss and sequence evolution (Tellgren-Roth et al., 2009). These genes are expressed in the saliva glands of vampire bats and the proteins they produce help to process the blood of birds and mammals. In humans, these genes protect against heart attacks by producing proteins that clear the blood vessels by degrading blood clots. The hairy-legged vampire bat’s (Diphylla ecaudata) PA genes resemble the PA genes of the closely related non-blood feeding bat species. These bats feed on the blood of birds and it seems that the activation of PA in saliva glands is enough to keep the bird blood flowing. However, in the two bat species that feed on mammal blood, common vampire bats (Desmodus rotundus) and white-winged vampire bats (Diaemus youngi, which also feed on birds), the PA genes have gone through more extensive modifications in order to better tackle the natural inhibitors of PA proteins in mammal blood. A transcriptome and proteome study of common vampire bats found additional genes expressed in the salivary glands (Francischetti et al., 2013). Furthermore, by comparing vampire bats and leeches to non-blood feeding species, Phillips & Baker (2015) found additional genes related to blood feeding, such as ectonucleoside triphosphate diphosphohydrolase-1 (ENTPD1), which has not before been linked to secretory expression. They also suggest that alternative splicing of genes has been an important mechanism for these species to rapidly evolve to feeding on blood.

In addition to blood coagulation, the vampire bats needed to overcome the bitter taste of blood. Bitterness in nature often means that the substance is poisonous and should be avoided. However, in all of the three vampire bat species there is a greater percentage of non-functioning DNA in the bitter taste receptor genes than in other bat species. These results suggest that these genes have been relaxed from selective constraint in vampire bats, which has led to a reduction of bitter taste function (Hong & Zhao, 2014).

Lastly, the problem with consuming blood is the ratio between amount of nutrition needed and the liquid consumed.  A typical vampire bat can consume half of its weight in blood in one feeding. The blood is then rapidly processed and the excess liquids are urinated within two minutes of feeding in order for the bat to take flight. Conceivably, the same effect would not be very convenient for vampires. If the vampire weighed for example 70 kg, it would need to consume 35 kg of blood in one feeding and urinate the excess liquid almost immediately, because the bladder can only hold about half litre of liquids. Furthermore, humans have about 5 kg of blood on average, so vampires would need to suck dry about seven people per night and urinate between victims, something that has not been discussed or shown in vampire stories, except in The Strain, where vampires defecate the blood while drinking. To compensate for the low intake of nutrients, vampires might slow down their metabolism and go to a hibernation mode and thus avoid the need to suck several litres of blood in one go. It would also enable fasting through hard times. In many stories, vampires have managed to survive without blood for days (see below).


Vampires are often regarded as undead; they are dead but behave like living beings, which in turn gives them eternal “life”. In this paper, I am not going to discuss whether vampires have a heartbeat or if they breathe (for that we would need actual vampire specimens); I will instead concentrate on how actual immortality could be achieved by giving real life examples.

First, we need to define what immortality is. The concept of biological immortality means that there is no mortality from senescence, which is biological aging. This of course means that the organism is not truly immortal, it can die through injury or disease. Vampires are often presented as highly resilient beings who can survive disease and injuries, but there are things that still kill them, like sunlight, a wooden stake through the heart, fire or beheading.

What is then the ultimate cause of senescence? It is still unclear how the process of senescence happens exactly, since it is a very complex phenomenon. This subject is under heavy research, especially in regard to how we could slow down or even reverse aging (de Keizer, 2017; see movies Self/less [Focus Features, 2015] and Mr. Nobody [Wild Bunch, 2009] for further thoughts). The research has been concentrating on gene expression changes, chemical and DNA damage, and telomere shortening. Telomeres are repetitive regions at the end of chromosomes. Every time cells divide, the ends of the chromosomes are progressively clipped in the replication process. Because the repetitive sequences in the telomeres are not protein coding, the clipping does not affect cell functions. When the telomeres are gone after a certain number of divisions, the cells stop dividing (Hornsby, 2007). However, cells have ways of replenishing the telomeres with an enzyme called “telomerase reverse transcriptase”. The drawback is that the majority of adult somatic (that is, non-reproductive) cells do not express telomerase, but it can be found for example in embryonic stem cells, male sperm cells, epidermal cells and in most cancer cells. In vampires, this enzyme might be active also in the adult somatic cells but this might pose an increased cancer risk. However, vampires might have ways to avoid cancer, as discussed below.

The way senescence happens is not universal; there are species where aging is negligible or cannot even be detected. There are two well-known examples of truly immortal species, the immortal jellyfish (Turritopsis dohrnii) and the animals from the Hydra genus. The immortal jellyfish, originally from the Caribbean Sea and now spread around the world, can use the process known as transdifferentiation to rejuvenate itself from its sexually mature free-swimming medusa form to sessile polyp form when the conditions turn harsh for the animal. When conditions are suitable again, the immortal jellyfish again transforms to its medusa form. This cycle can in theory continue forever, making the species immortal in the biological sense. However, this does not save the jellyfish from predators and diseases. The immortal jellyfish also appeared in the TV series Blacklist (Sony Pictures Television, 2013–present), where its cells were injected into humans in order to generate immortality. In the real world, science is not that advanced yet and it is also highly unlikely that it would be this easy to achieve immortality.

Hydras have been under more research than the immortal jellyfish. Hydras are simple freshwater animals (also cnidarians, like the immortal jellyfish) whose cells can continually divide and not undergo senescence. One gene, “Forkhead box O” (FOXO) has been extensively studied in hydras (and also in other species, like the nematode Caenorhabditis elegans, mice and humans) (Boehm et al., 2012; Martins et al., 2016). In hydras, this gene is the main player behind the renewal of the cells. In other species, this gene has been linked to aging and longevity in many studies. In an essay by Schaible & Sussman (2013), the authors suggested that during the evolution of the FOXO gene, its function changed from Hydra’s life span extending role to many other pathways related to maintenance, which altered the gene’s rejuvenating functions in multicellular eukaryotes such as humans. Thus it might be that in vampires this gene (or actually all the FOXO genes – mammals have four of these genes) have retained the original function of FOXOs.

In the mammalian world, naked mole rats (Heterocephalus glaber) and Brandt’s bats (Myotis brandtii) are exceptionally long-lived compared to other small sized mammals. Naked mole rats are known for some very peculiar characteristics. They can survive anoxic conditions, they have delayed ageing and live up to 32 years, and the species is highly resistant to cancer, among other things, making them a very interesting species for scientists to study. In studies of the longevity and cancer resistance of this species, scientists found that a gene called INK4, which is the most frequently mutated gene in human cancer, produced a new product through alternative splicing. This protein isoform (that is, protein variant), called pALT(INK4a/b), prevented the mutated cells from clustering together and thus made the naked mole rats more resilient to cancer (Tian et al., 2015). In another study by the same group, extremely high-molecular-mass hyaluronic acid was found in naked mole rat fibroblasts (the most common cells in the connective tissue of animals). The molecular weight was over five times larger than that of human or mouse hyaluronic acid. It was speculated that a higher concentration of hyaluronic acid evolved to keep the skin elastic in underground tunnels. In addition to skin elasticity, long hyaluronic acid molecules wrap around cells tightly, preventing tumor cells from replicating (Tian et al., 2013). Whole genome sequencing revealed additional genes that could be linked to longevity in this species (Kim et al., 2011).

Brandt’s bats are known to live for over 40 years, making it the most long-lived mammal of its size. In the whole genome study of the species, Seim et al. (2013) suggested that a combination of different adaptive characteristics such as hibernation, low reproductive rate, cave roosting and an altered growth hormone/insulin-like growth factor 1 axis could extend the Brandt’s bat’s lifespan. Furthermore, FOXO1 gene was expressed in high levels in Brandt’s bat suggesting a possible role also in the longevity of this species. Hibernation in general has been linked to survival of different species allowing them to withstand extreme conditions (Turbill et al., 2011; Wu & Storey, 2016). The molecular difference between hibernators and non-hibernators seems to be in gene regulation rather than a difference in the DNA sequence itself. Differential expression was detected in the genes that were involved in metabolic pathways, feeding behavior, and circadian rhythms (Faherty et al., 2016). Hibernation or some other kind of dormant state seems to be present in vampires as well, helping them to get through tough times. In the Vampire Chronicles by Anne Rice, the vampires go to a hibernation-related state to cope with changing times. In the Underworld movies, two of the elders are kept in hibernation while a third reigns over the vampires. The reign goes in cycles, each of elders having their turn over the vampires and slave lycans. This cycle has social reasons, but it also gives rest for the elders from their immortal life.


Vampires are creatures of the night and sunlight is often regarded as deadly to them; in many occasions, they burst into flames whenever in contact with sunlight. It is an adverse trait for vampires and most probably emerged through pleiotropism. Pleiotropism is a phenomenon where one gene affects two or more unrelated traits (Paaby & Rockman, 2013). Mutations in genes causing immortality or blood consumption could also cause death by sunlight (antagonistic pleiotropy). Real life examples of bursting into flames due to sunlight are obviously not found, but sun can cause problems to people with certain conditions. Sunlight can cause severe allergic reactions, people can suffer from blood disease called porphyria, or have a rare recessive genetic disorder called “xeroderma pigmentosum”.

“Sun allergy” is an umbrella term for a number of conditions where rash and blisters occur on skin that has been exposed to sunlight. Some people have a hereditary type of sun allergy, such as hereditary polymorphous light eruption, others a non-heritable type, such as solar urticaria. In some cases, symptoms only occur when triggered by another factor, such as certain medications or skin exposure to certain plants. The allergic reaction to sunlight occurs in the same way as in any other allergic reaction, although it is still not clear what the triggering component is. Somehow, the immune system recognizes the sun-altered skin as foreign to the body, which in turn activates the immune defences against it. If vampires suffer from sun allergy, could strong antihistamines and a high sun protection factor sunscreen help them survive under the sunlight, in the same way as people with sun allergies? As death is a very severe reaction to sunlight, it is likely that vampires do not suffer from a sun allergy but from something more serious.

Porphyria, a group of blood diseases, have been suggested as a possible explanation for vampire myths but these ideas have been rejected in later papers (Winkler & Anderson, 1990). However, the mechanism behind porphyria could still shed light on why sunlight would be poisonous for modern vampires. In the cutaneous forms of porphyria where the skin is mostly affected, sunlight can cause pain, blisters or open sores to the patients. The disease is often hereditary due to a mutation in one of the genes that make the heme molecule (a component of hemoglobin, the red pigment in our blood): ALAD, ALAS2, CPOX, FECH, HMBS, PPOX, UROD, or UROS (Badminton & Elder, 2005). These genes could also be suitable candidates for vampire sunlight avoidance.

There is an even more severe sunlight sensitivity illness, the rare hereditary condition called “xeroderma pigmentosum” (XP). In extreme cases, the patients need to avoid all exposure to sunlight as it can cause severe sunburn with redness and blistering. If not protected from the sun, people with XP have a high risk of developing skin cancer. XP patients’ eyes are also very sensitive to sunlight and some of the patients have neurological problems such as seizures and hearing loss. The condition is caused by mutations in the genes that repair DNA damage. This causes a deficiency in DNA repair after ultraviolet damage to cells, which in turn accumulates abnormalities to the DNA causing the cell to become cancerous or die. In most of XP cases, mutations occur in these four nucleotide excision repair related genes: POLH, XPA, XPC or ERCC2 (Schubert et al., 2014). In addition to porphyria genes, these are also potential candidates for vampires’ adverse reactions to sunlight.


Obviously, the transformation from human to vampire would affect many genes, some of the changes being bigger than others, which makes the genetic modification of human to vampire even more difficult. From the real life examples, the PA (blood coagulation) and FOXO (immortality) genes seem to be strong candidates. Furthermore, it is also possible to find more suitable genes to test and to investigate interactions between hematophagy, immortality and sun avoidance genes by using network analysis such as Genemania (Warde-Farley et al., 2010). For example, when inserting the human ortholog (roughly put, the equivalent gene) of bat PA gene, the plasminogen activator, tissue type (PLAT), the FOXO genes FOXO1 and FOXO3, and the four XP genes, POLH, XPA, XPC and ERCC2 to Genemania, it is possible to see how the genes are linked and what additional genes might be involved (Fig. 2).

Figure 2. Gene interaction network of the genes PLAT, FOXO1, FOXO3, POLH, XPA, XPC and ERCC2 done with Genemania. Showing 20 related genes with 27 total genes and 207 total links. Input genes are indicated with stripes.

In many of the traits mentioned above, we assumed that mutations in these candidate genes would be the cause of the vampiric traits. However, mutations are not the only possible cause. Epigenetic changes are functional changes in the genome that do not involve modifications in the DNA. Such mechanisms are, for example, DNA methylation and histone modification. External or environmental effects can cause DNA methylation and change gene expression. In vampires, both mutations and epigenetics could be possible players, causing changes and vampiric traits. Furthermore, if vampirism is caused by a virus or a parasite, we need to take into consideration the possible ways the pathogen could affect the human cells, which is a topic of its own.


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I would like to thank Dr. Olaf Thalmann and Angela Boeijen for insightful comments and Nina Haglund for language revision. 


Dr. Veronika Laine is a molecular biologist working currently with the great tit and she is especially interested in behavior, genes, pleiotropism, bats, kittens and vampires, especially Eric Northman. She plays too much video games.

[1] The Queen of the Damned, by Anne Rice (1988).

[2] Old folklore; Buffy the Vampire Slayer (20th Television, 1997–2003).

[3] Daybreakers (Lionsgate, 2010); the Underworld film series (Lakeshore Entertainment, 2003–2016); The Strain (20th Television, 2014–2017).

[4] Dracula, by Bram Stoker (1897); The Vampire Diaries (Warner Bros., 2009–2017); The Twilight Saga (Summit Entertainment, 2008–2012); True Blood (HBO Enterprises, 2008–2014).

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