After a 13-year hiatus, the long-awaited sequel to James Cameron’s 2009 Avatar is out…

**Caption:** among the many interesting animals of *Avatar: the Way of Water* is the gar-like skimwing. It goes without saying that all the images used in this article are (c) 20th Century Studios, many taken from the **Avatar Wiki**.

I and my family saw it last night. It’s Avatar: the Way of Water, and… my god, it’s visually amazing. James Cameron has made no secret of the fact that these movies are love-letters to the natural world, to environments and spectacular creatures, and to indigenous cultures and peoples. We – as in, all of us, whatever sort of life we lead – need to maintain our connection to the rest of the natural world, to preserve and maintain what remains, and to me it’s obvious that this is the primary theme of these movies. It feels childishly naive to even say it, but colonisation, industrialisation, destruction and exploitation are very much pitted as bad in the two Avatar films; those fighting to save and preserve the natural world and its indigenous peoples are the opposite.

**Caption:** it should be obvious to anyone who follows me on social media that I talk often about those remaining green spaces close to me. We all need to do what we can to see that these places persist. I feel as if almost everything is at threat of destruction or deterioration.

That as it may, I am of course here for the creature design (not for the plot or storyline). Visually, Avatar: TWOW is simply incredible: huge congrats to everyone involved for the awe-inspiring and beautiful animals and environments they designed and built. But, alas, I’m not about to publish an article on these new creatures, at least not today. Instead, I’m using the release of Avatar: TWOW as an excuse to dust off and republish the 2010 Tet Zoo ver 2 article I published on the animals of the first movie, since now is as good a time as any. The article originally appeared here (looks like the reader comments have been saved, which is good). I’ve made a few minor editorial tweaks and corrections, and have also added new images. But here we go…

Tone and I recently went to see Avatar. I’ve been reading up on the movie for months and was really looking forward to seeing it. As implied above, I’m not here to talk about storylines and plot devices… you want to know about the creatures. And a lot of thought and time obviously went into the design of Pandora’s ecosystem and creatures. In part, I’d say that this was a success: a lot of people (even many not that interested in the natural world) have been very much taken in by the movie’s xenobiology, and let’s hope that this inspires them to become interested in, and passionate about, the biology and ecology of the real world. Without further ado, here are assorted musings on Pandora’s creatures. And I’d be interested in your take on them too, so feel free to discuss things in the comments below.

I should add here that the original article included a spoiler warning at this point. Seeing as I’m writing here about a movie that came out 13 years ago, I’m assuming that I no longer need to include one though.

Caption: in case there’s any doubt, I had no involvement at all with Avatar or its creature design, nor do I have any reason to think that those behind the making of Avatar: TWOW are aware of my writings on the first film. I have, however, worked closely with several people connected to the film, in particular animation supervisor Andy Jones.

Banshees and other fliers. In my opinion, the coolest creatures in the movie are the flying beasts: the dragonesque Mountain banshee and the awesome, gigantic Great leonopteryx. Banshees are bluish-green, long-necked creatures with membranous wings and a pair of hindlimbs that also sport flaring membranes (there are two kinds of banshee, but we only get to know Mountain banshee in the movie). Unlike most Pandoran creatures they aren’t hexapodal, and are assumed to have lost one of the rear limb pairs.

A prominent alula-like clawed digit (flanged with another membrane) allows the animal to climb and cling; it looks similar to the large thumb present in megabats. The distal parts of the wings are translucent. The hindlimbs also have a prominent, alula-like, clawed digit. There’s a propatagium, but the main part of the wings are not composed of a single, continuous sheet (as they are in pterosaurs); instead, there are several rods embedded within the main membrane that help it to fold and fan out. The wings thus combine elements of the pterosaur wing with bird primary feathers and bat fingers.

**Caption:** banshees and Na’vi have a long history of association. Na’vi riders of the Omaticaya clan form lifelong bonds with Mountain banshees (known to them as ikran); the image here shows Naytiri’s ikran Seze.

When the animals fly, they stick the hindlimbs out sideways from the body; while the hindlimbs have membranes, these aren’t connected to the body or forelimbs. As a consequence, the flight configuration reminded me of certain pterosaur reconstructions, since some authors have also shown pterosaur hindlimbs projecting outwards and backwards. The banshee tail is long and slim, with a horizontal vane at the tip. The superficial similarity between Ikran and certain pterosaurs have not gone unnoticed by palaeontologists, as is demonstrated by the naming of the Chinese Cretaceous pterosaur Ikrandraco avatar Wang et al., 2020.

**Caption:** several very nice banshee figures are available. These images show the McFarlane Toys banshees, released in 2022 to coincide with *Avatar: the Way of Water*.

The team behind Avatar‘s creatures put some thought into stuff like respiration, aerodynamics and gravity. Gravity is lower on Pandora than on Earth (it’s a moon orbiting the gas giant Polyphemus), and this has facilitated the evolution of giant fliers. It’s also noted that at least some Pandoran creatures have stronger bones (or bone analogues) than what we’re used to, as a sort of carbon fibre reinforces the animal’s tissues, and also makes them lighter. Banshees also have ‘intake valves’ – spiracle-like openings on their thorax – that allow them to take in more air than allowed through the head alone, and they’re said to have a unidirectional respiratory system, with used gas passing outwards via gill-like slits on the posterior thorax. This sort of stuff is fairly plausible and we might honestly expect alien creatures to have respiratory systems like this, rather than the nostrils and whatnot that we’re more familiar with.

**Caption:** banshees recall the Cretaceous dromaeosaurid *Microraptor* in having flight surfaces on two limb sets, rather than one, and in having what might be a lift-creating structure at the end of the tail as well. Spectacular *Microraptor* specimens (like IVPP V13352, shown here at left) have been known since around 2002. At right, a scale model (showing one of several possible hindlimb poses) of *Microraptor* produced for a study on flight performance in this animal, from **Dyke *et al*. (2013)**. Images: Colin Palmer; **Dyke *et al*. (2013)**.

While the banshees look pterosaur-ish, I’m curious to know if their designers were also inspired by the crow-sized, four-winged dromaeosaurid dinosaur Microraptor. Microraptor – famous for having really long feathers on its hindlimbs as well as its arms – has been imagined by some to ‘flap’ with its legs as well as with its arms (this may not be at all possible, but let’s avoid that debate for now), and the double-winged flight motion of the banshees reminded me of this hypothetical flight style. There’s also a scene where the two banshees ridden by Jake and Neytiri glide rapidly downwards in parallel with a vertical cliff face. This reminded me of some of the stunts practised by people who use proximity gliding suits.

**Caption:** great raptorial hooks and recurved, slightly heterodont teeth (or tooth analogues) are among the obvious predaceous specialisation of the Toruk. As is the case in the Thanator and Viperwolf, the teeth are black, which suggests that they are not composed of enamel and other materials familiar on Earth.

Toruk. The Great leonopteryx – known to the Na’vi as Toruk – is neat, and the movie is worth seeing for this creature alone in my opinion. A giant predator of banshees and other animals, it’s a vividly coloured, spectacularly crested hexapodal eagle-like monster, with terminal hooks on its rostrum and a pair of grasping posterior limbs. Some sequences in the film remind me of real-world sequences depicting the behaviour of giant raptors, most especially the incredible South American Harpy eagle Harpia harpyja.

**Caption:** Toruks can open and close the slots between their leading wing elements, and rotate the individual elements as required for vortex formation and so on.

Unlike banshees, the Toruk retains all three limb pairs. Its wings are also more complex than those of banshees: they’re mostly membranous, but there are three feather-like elements along the wing’s leading edge that can separate and create leading-edge slots. Again, there’s an alula-like digit with a huge, curved claw. We see clearly that the digit is used to help support the animal’s weight when it rests, with wings folded, on the ground. Rather than being directed laterally, the digit projects directly forwards when the animal is grounded.

**Caption:** the Toruk is absurdly big, with a wingspan that can exceed 25 m. It preys on other predators (mostly banshees) and is a solitary, large-brained animal with a central role in Na’vi lore.

Dark lateral ridges project from the sides of its head, shading its eyes (like many of the Pandoran creatures, it has more than two eyes: there are two smaller eyes behind the main ones). These ridges make its head superficially raptor-like. Enormous, sheet-like blue sagittal crests project both from the dorsal surface of the head and from the lower jaw. To my eyes, these make the creature superficially similar to a tapejarid pterosaur, in particular to the sail-crested Tupandactylus (for a now very dated ver 2 article on tapejarids, go here).

So – even before I’d seen the movie – I’d decided that the banshees and Great leonopteryx were inspired by (1) microraptors, (2) tapejarids and (3) proximity gliding suits, plus with a bit of raptor and megabat thrown in too. While there might be some truth in this, the creatures are actually more complex, with inspiration for their design apparently drawn from even more diverse sources. Neville Page – mostly responsible for the creature’s design – has noted how he worked to emulate the smooth lines and streamlined shapes of creatures such as Great white sharks. It’s evident from the way their jaws open that banshees have some similarities with teleost fishes: when the animals gape, a maxilla analogue is pulled downwards from the upper jaw. A neat detail is that the entire tooth row in the upper jaw is mobile: when the jaws are closed, the teeth fold posteriorly into slots on the ventral surface of the maxilla analogue, but they swing anteroventrally when the jaws are open.

**Caption:** at left, a proximity gliding suit or wingsuit. Certain of the scenes in *Avatar* are reminiscent of wingsuit flight sequences. At right: the amazingly elaborate skull of the tapejarid pterosaur *Tupandactylus imperator*, showing the extent of the soft tissue and the associated branched, filamentous structures on the rear spar. Images: Wingsuiting, CC BY-SA 4.0 (from **here**); Cincotta *et al*. (2022), CC BY 4.0 (from **here**).

Also on the teleost-like anatomy, we have good views in the movie of a Great leonopteryx skull (the Omaticaya clan of the Na’vi keep one in their giant hometree for symbolic or ceremonial reasons: the creature is important in their mythology). It doesn’t look tetrapod-like, but has the corrugated, gnarly bone texture you normally associate with fish. Having said that, there are some bird-like aspects to the skull as well. The rostral hooks recall the hooked tips of raptor bills, for example. Some articles state that manta rays, skates and plesiosaurs were also inspirational in the design of these creatures, and Wayne Barlowe (who was involved in creature design early on) is on record as saying that he based the banshee’s sleek design on sports cars. The patterns and colours of these animals were apparently inspired by those of birds, poison-dart frogs and monarch butterflies.

Hammerhead titanothere. Pandora is home to at least one mega-herbivore, the spectacular, enormous Hammerhead titanothere. The fact that it’s called a titanothere might explain the origin of its design. Titanotheres, generally called brontotheres (or brontotheriids) these days, are rhino-like Eocene perissodactyls: they’ve been discussed on Tet Zoo a few times. Go here, here and here.

**Caption:** the giant brontothere *Megacerops* (previously known as *Brontotherium*, *Menodus*, *Brontops* or *Titanotherium*) is diverse in horn form, this varying according to age and sex. I like this image (taken at Badlands National Park) because it shows an especially big-horned individual. Image: Dave Fuentes, **from here**.

The hammerhead-like, err, head is a nice idea, but – for my tastes – too ‘familiar’ given that we all recognise this shape. They could have gone for the same idea (after all, a battering-ram head could well be useful to a giant terrestrial herbivore: note that the Hammerhead titanothere does not have eyes on the lateral ends of its ‘hammer’), but made it far freakier and more alien. The animal uses a fan-like cluster of flag-like structures on the top of its head as a signalling device. I liked the fact that (ordinary) guns wouldn’t be much use when confronting an animal this large and formidable: shoot it, and it will still be coming right for you. The stampede scene was awesome and very satisfying.

**Caption:** Hammerhead titanothere from the encounter scene with Jake. Note that the eyes are present posteroventral to the hammer-like cranial structure. Titanotheres are social animals, adults varying in the colour of their fan-like cranial structures. They reach 11 m in length.

Thanator: super-predator. The Thanator – a large hexapodal terrestrial predator – is another of my favourites from the movie. I initially thought from the trailers that its designers had been looking at gorgonopsians, but, nope, no good reason for thinking this. Cameron has said that it was meant to be a sort of super-panther.

Caption: Thanator in lateral view, with retracted lips and elevated soft cranial flaps. The opposable nature of the digits on the anterior limbs is obvious here, as is their heavily muscled form.

It’s dark, with smooth, mostly leathery skin, and dextrous muscular limbs with long, sharp claws borne on semi-opposable digits. Its head is long with a sort of naked, protruding rostrum and large anterior fangs. Its eyes are large and it has peculiar flap-like and soft, spike-like structures arranged around the back of its head. Not sure what these are for, though they might be used in visual display. In one scene, it grabs Jake’s back-pack. Jake slips out of the back-pack and escapes, and when the Thanator notices, it opens it jaws, raises its paddle-shaped tail, and flares the flaps and spikes outwards. That’s some nice body language there, implying intelligence and mood.

**Caption:** a Thanator encounters Naytiri, adopts a conciliatory pose, and invites her to ride it into battle. Alas, this doesn’t end well for the Thanator.

While I think that the Thanator is a very neat looking creature, it does exhibit ‘Hollywood super-predator syndrome’ to a degree, and acts like an unstoppable, psychotic whirlwind, smashing through vegetation, tearing tree roots up, and altogether doing everything possible in order to kill and eat the object of its attention. It’s not even deterred by a barrage of automatic gunfire, and almost pursues Jake right off the end of a cliff. It also appears to be incredibly intelligent, grabbing guns and discarding them during combat, for example. We all like predators in movies to be super-predators with super-powers, and on the one hand you might argue that creatures like the Thanator are so over the top that they really create the wrong impression as to what real predators are (mostly) like. One day I want to see a movie where the predator is a conservative coward that faints when confronted with a gun.

On the other hand, we know for sure that certain predators can be unbelievably gung-ho and bold on occasion, Exhibit A being the various tales that involve man-killing big cats that have moved into dwellings and vehicles to kill people. And I should add that I only recently read Stephen Herrero’s 1985 Bear Attacks: Their Causes and Avoidance. Oh. My. God…

**Caption:** I really like this poseable Thanator figure, made by Mattel in 2011. I’d love to own one but have never seen it for sale. I’m not sure it was ever available here in the UK.

I do dislike one other thing about the Thanator: its roars sound exactly the same as the tyrannosaurs in the Jurassic Park movies. Having said all this, it’s an awesome creature and its two major appearances in the film are among the highlights.

Direhorses and viperwolves: not my favourites. As for some of the other creatures, I confess that I found them to be less interesting: they were just too similar in form or behaviour to real animals, or, in other words, too derivative. I refer in particular to direhorses and viperwolves. I didn’t like the Direhorse because it just seemed all too much like, well, a horse, though a very big, alien-ish horse… though still a horse.

**Caption:** direhorses can bond with Na’vi through integration of their neural whips. This image is from a battle scene where a Na’vi cavalry battles a heavily armed RDA squadron involving infantry augmented with an AMP (amplified mobility platform) unit.

To their credit, Cameron et al. did try and make the animal fit in with the rest of Pandora: direhorses have the same thorax spiracles as the banshees, and – in keeping with the planet’s riotous and super-elaborate flora – we see in the movie that direhorses aren’t grazers; instead, they’re nectarivorous, and feed with a long proboscis from flowers. Needless to say, the evolution of gigantic nectar-feeders like this could only work in an ecosystem where flowers are enormous, permanent, and produce huge quantities of nectar.

**Caption:** it should be obvious that the direhorse face superficially recalls that of a giant anteater. Horizontal pupils on the largest eye set are obvious, as is an anterodorsally projecting cranial crest (continuous with a tall ridge present along the midline of the neck) and the two parallel neural whips.

The viperwolves were the least compelling creatures for me. They’re long-bodied, slinky, black-skinned hexapods with hand-like feet and facial tissue that can be retracted right off their scary looking, pointed teeth. They have floppy neural whips at the back of the head. We see a viperwolf group behave in an extremely tenacious way when Jake is alone in the forest for the first time; despite his use of fire and other aggressive tactics, he’s not succeeding in preventing their attack until Neytiri comes to his rescue. I did initially think that this was unrealistic, but I now think that this appraisal is unfair given how social canids will behave when harassing animals the size of people and large deer. Whatever, I didn’t especially like the design. They remind me of small, black versions of Falkor the luck dragon in The Neverending Story.

**Caption:** viperwolves are sharp-toothed, lithely built predators that share highly mobile lips, large and pointed teeth (or tooth homologues) and dextrous digits on their anterior limbs with thanators. Viperwolves are co-operating, pack-hunting predators.

The Na’vi and others. Finally, the creatures that feature most strongly in the movie are of course the blue humanoid Na’vi. In many ways these are the easiest of the creatures to criticise, if – that is – you’re like me and think that the odds of human-like creatures evolving independently of us are vanishingly small and downright improbable. On the one hand, you can argue that successful alien films can work fine when the creatures don’t look at all human. And – given that humans are meant to be remotely piloting genetically modified Na’vi bodies – it wouldn’t have mattered to the story what the Na’vi looked like.

**Caption:** a scene showing several of the main Na’vi characters. It’s hard to make people care about characters in a film where those characters aren’t relatable.

On the other hand, this movie is about warfare between cultures, about allegiances, and – I suppose – about cool-looking shit, so it figures that good looking ‘people’ need to feature large in the story. Originally, the Na’vi were going to look weirder, with gills, fins and other structures, but over time they were made to look more human simply to appeal more to the audience. Female Na’vi have breasts, specifically for this reason, apparently. So, yeah, the Na’vi are nothing more than attractive, semi-naked blue people with cat-like features… let’s let it go. When we’re introduced to their spiritual beliefs and practises, the Na’vi incorporate details inspired by or based on those of various indigenous groups of people, and it’s not really possible to think about the plot without comparing it to the conflicts that have occurred between invading European cultures and indigenous ones.

The idea in the movie is that Pandora includes a lineage of primate-like animals whose evolution has closely paralleled primate evolution on Earth. Early in the movie, we see the hexapodal, arboreal Prolemuris (they also make a guest appearance in Avatar: TWOW). This animal is to the Na’vi what lemurs are to us (distant cousins that share a common ancestor), and this explains why its two more anterior limb pairs are partially fused. Presumably, the arms of the four-limbed Na’vi represent two, fully fused original limb pairs (though, if this is true, it might be odd that the Na’vi have only four fingers).

**Caption:** Prolemuris is a social, arboreal, primate-like Pandoran creature. Unlike the majority of Pandoran animals, they only have a single pair of eyes and only one neural whip. Both features are shared with the Na’vi.

Several other creatures feature in the movie. The hexapede is a deer-like animal with a fan-like cranial structure, while we also see a small, arboreal hexapod that looks like a cross between a leaf-tailed gecko and a frog. When disturbed, it unfolds a giant glowing spiral-shaped structure on its back and takes flight while spinning and emitting bioluminescence. Bioluminescence is a major theme on Pandora, with virtually all of the plants glowing in the dark.

A linkage of some sort unites most (or all?) living things on Pandora, and this proves key to the victory of the Na’vi over the invading humans. The Na’vi ‘store’ their cultural heritage in the memory of special trees, and it seems that memories and even personalities of individuals can be uploaded and downloaded via biological links with these plants. The Na’vi also have neural links with the creatures they ride (direhorses and banshees) and have to plug into them via a tentacle-like structure (termed the queue) on their heads. I don’t want to over-analyse any of this, but the link between the Pandoran creatures and the trees reminded me of the mycorrhizal networks known to involve certain trees and nearby fungi. I wonder if this is what inspired the whole idea?

**Caption:** mycorhizzal fungi have a mutualistic relationship with plants via a subterranean network. Inter-plant chemical messages relating to predation, disease and perhaps environmental stress are sent across the network. Image: Charlotte Roy, Salsero35, Nefronus, CC BY-SA (**original here**).

Finally, I loved the technology featured in the movie. Avatar is set in 2154 and I think the tech looks plausible based on the way things are going (yeah, if our culture lasts that long). The aircraft looked and behaved plausibly. The fat, heavily armed C-21 Dragon Assault Ship has an aesthetic charm, and the gigantic Valkyrie reminded me a lot of the Hercules and other bombers it was probably inspired by. The AMP suits look like evolved, combat versions of the cargo-loaders we know and love from Aliens. Though, I think we can doubt that an exosuit would be kitted out with a giant KNIFE.

**Caption:** Ellen Ripley using a P5000 load-lifter, a scene from *Aliens*, of course. *Aliens* and *Avatar* almost certainly occur in the same universe… though I don’t know if this has been specifically confirmed.

The Avatar movies really are the visual feasts that we’ve been promised. Both the 2009 original and the 2022 sequel look great, and the cool animals – the Mountain banshee, Great leonopteryx, Thanator and so on – make them well worth a viewing, or two. I ended up seeing the first film several times in the cinema. Avatar: the Way of Water is brand-new at the time of writing (December 2022) and my first and only viewing is still fresh in my mind. Additional movies in the series are due to appear over coming years, meaning that the series will be a mainstay in discussions about creature design and speculative biology for some time yet.

That’s where we’ll end things for now. Say what you like about the plots of these films… I cannot help but like them in view of the stupendous creature design they include, and their emphasis of our empathy with the natural world.