Shaping hexapod shapes: high browsers

 First of all, I apologise for taking so long to post something. I have been much too busy writing large reviews. Some of you probably hoped that I was busy dealing with publishers to shape The Book. If only! At present it seems that sending manuscripts to publishers in the way they want you to has as much apparent effect as shouting in a vacuum. Do you remember the slogan 'In space no-one can hear you scream?' It's like that. If you didn't know the slogan, it's from the first Alien film (1979). Rather like that particular alien I intend to keep coming back, though.

Nevertheless, I did toy with the directions evolution might take the hexapod design into. The fun of doing something like this is that you must follow the rules, and these state that the basic anatomy is a given, exactly like in real biological evolution. In this case, the challenge was to evolve a high-browsing animal, something like a giraffe or possibly a sauropod. One way to get an animal's mouth high up into the air is of course to enlarge the entire animal but that is rather boring. Other solutions would be to stretch all animal parts vertically, or to only stretch parts close to the mouth, such as the mouth parts themselves, the skull and  and the neck. I chose somewhere in between the last two choices: legs are elongated, but not as much as the neck. Or necks.

Hexapods do not have vertebral columns in the sense of one chain made out of a large number of short bones. Instead, they have two parallel chains together forming a 'scala' (ladder). In the neck the original two series of bones forming the stiles of the ladder merged into one structure while the rungs disappeared altogether. This happened early on, when hexapods were sea dwellers, as an obvious way to expand the range of the mouth at little cost. With a long mobile neck you can increase the 'feeding envelope', which is the volume the mouth can reach while the body stays still. The same idea applies to sauropod necks and rusp mouthparts (see here). As you may have noticed from earlier sketches (here and here), hexapods first have a proximal neck of two segments, then a sensocranium, a distal neck of two segments and finally the jaw apparatus. 

Click to enlarge; copyright Gert van Dijk

 

Click to enlarge; copyright Gert van Dijk

Here are two sketches in which I played with such a high browsing hexapod scheme. I also stretched the sensocranium. Normally that is just a bulb sitting above where proximal and distal necks approximate one another. Here the neurocranium is truly a part of the chain of segments. If you look closely, you will see some curved lines gradually following the angled neck bones. That is the asymmetrical oesophagus, situated on one side of the animal, not in the middle below the bones. See here for more on that odd feature.     

Elongating the neck in this way involves elongating the individual bones. I felt that such animals should be able to hold their necks horizontally, not just vertically. Such a horizontal posture will stress such long bones though. Here's why: take one such long bone and assume that the proximal end (near the body) is fixed. All bones attached at the distal end then act as a weight to pull that distal end down. This weight tends to bend the bone down, which stretches the top margin of the bone while simultaneously compressing the bottom margin of the bone. Bone tissue is usually better at withstanding compression than tension, while tendons have opposite characteristics. A typical vertebrate trick to solve these stresses is to string strong tendons along the upper surface of the backbone to take care of those tensile forces, leaving the bones themselves to deal with compression. The trick works better when the tendons are at some distance from the centres of the vertebrae, and that is what neural spines, the bone projections sticking out of the top of vertebrae are for: they keep the tendons at a distance.

Click to enlarge; from Preuschoft & Klein 2013

The image above shows a scheme with vertebrae, spines and a big tendon in place for a sauropod. In the drawing, the neck vertebrae are fairly long, so we are getting close to  hexapod anatomy. But do extra-long bones pose additional problems? 

Click to enlarge; copyright Gert van Dijk

I am still considering that and can at present only offer some thoughts. Let's start with one spine on each bone; there are tendons running from the spine that bridge the joint on either  side (A). That means the tendon is attached at a sharp angle to the bone, whereas it would work better if the attachment were nearly vertically. That can be done by making the spine longer, or, in this case, by placing it near a joint (B). That works well for the closest joint but worse for the farther one, so that tendon now attaches to the bone itself and no longer crosses a joint. It still looks as if the bones would have to withstand lots of bending forces, which we do not want. Very well, let's duplicate the 'spine & tendon structure' so we have a spine at each end of the bone (C). But just to be safe we may add an additional ligament crossing all elements (D). So there we are.

Click to enlarge; copyright Gert van Dijk

A new problem now may be that flexing the neck might bring one spine into contact with the other, preventing the movement. That can be solved by moving the spine away from the joint again, but there is another solution. We already allowed the oesophagus to be asymmetrically placed to one side of the animal. Could we also place the distal and proximal spines off-centre, so one is displaced towards the right and the other towards the left? That is shown in the image above.

Do the rules allow that? We are all used to the fact that vertebrate skeletons are nicely symmetrical when our intestines are not all that symmetrical. I wonder why; anyway, there are skeletal exceptions, such as narwal teeth and crabs that have one big and one small claw. My guess is that large anatomical asymmetries between walking legs would make both the mechanics and the neurological control of walking extremely difficult, without offering any advantages whatsoever. (Mind you, quite a few neurological functions are already asymmetrical even when the underlying anatomy is nicely symmetrical. Humans have handedness, but also 'footedness' and 'eyeness'; bees have 'antennaness', and I could continue). Anyway, anatomical asymmetry in a nonlocomotory part should not cause any major problems. So perhaps we can add a new rule stating that skeletal asymmetry occur in Hexapods. I must think some more about that and will try to find out whether anyone has already solved the riddle why vertebrate innards are more asymmetrical than their skeletons.

What's a head? Or a neck...

Yes, it's a silly title. Isn't a head, rather obviously, that part of an animal where vision and hearing are gathered, along with their associated neural processing units, as well as air and food intakes, along with whichever specialised organs that takes? For air intake, you could read 'nose', and for 'food intake' there is the whole complex assembly of jaws, teeth, a tongue as well as an oesophagus. 

That seems right for humans and other terrestrial vertebrates, but a few moments' thought reveals that 'obviously' does not belong in this 'definition'. Fish have heads but take in water, not air, so the 'air intake' should become an 'inlet for gas exchange'. The hearing organs of many arthropods are not in their heads, and so the associated bits of brain need not be in the head either. The description supposes that the rest of the animal is distinct from the head. But in octopuses there is no distinction between head and body, with the limbs ('arms' that are not accompanied by 'legs') attached directly to the head/body unit. For spiders a similar point can be made that the head is merged with part of the body. 

I could go on, but the point is that a 'head' is not as clear an entity as you might think. A 'head' is the result of 'cephalisation', described by Wikipedia as 'an evolutionary trend in which... the mouth, sense organs, and nerve ganglia become concentrated at the front end of an animal, producing a head region.' I like 'head region': it provides the looseness we apparently need to describe what a head is. 

Furahan 'Scalates' certainly underwent cephalisation, and their heads definitely contain eyes (four of them) and ears, in terrestrial forms (also four) along with enough brain to do the heavy duty processing these organs require. But the 'intake for gas exchange' is not in the head at all. The intake for food is there, with jaws (originally six, later four) at the front of the animal, so clearly these are in the head. Well, that depends... 

Click to enlarge; copyright Gert van Dijk

A clade of Furahan Fishes ('Fishes V') developed a 'neck' in the form of a two-bone articulated connection between the body and the 'head region', allowing the latter considerable freedom of movement relative to the body. The thing is that a similar 'neck' also evolved between the 'head part' housing eyes and ears, and the mouth region. Are there then two heads, or is the entire region one head? 

The two necks, usually called the proximal (closest to the body) and distal (farthest from the body) neck, each consist of two long bones with a joint in between. The two 'heads' are the 'sensocranium' and the 'orocranium'. The image above shows a fragment of a painting showing a species of Fishes V with just that arrangement. You cannot see the joints in the necks clearly in these streamlined Fishes, but they are there. Those long neck bones form a big difference from the kind of vertebral columns we are used to, with their string of many small bones. Early Scalates never had a central string of small bones for evolution to play with; instead, they had a 'ladder'. The resulting neck movements look rather like those of a vertebrate arm or leg, with sharp angles, not at all like the curves of a lizard tail or a giraffe or sauropod neck. But that ungainly look need not be a functional handicap; despite the sticklike nature of human arms, baseball and darts players manage to land small objects with incredible precision quite a distance away. Of course, the trick is having a good brain in control.

We now only need to discuss bending of other organs in those necks, such as the oesophagus. An oesophagus relies on peristalsis making it flexible and elastic, so the structure itself should be able to withstand folding for a while. The nerves and arteries in our shoulders and elbows have to withstand bending too and usually do so fine. (Admittedly, if you spend too long in one position local pressure may pinch blood vessels, so people occasionally find that an arm or a leg, or just one nerve, has 'gone to sleep'.) If the oesophagus would be folded, it would probably be incapable of propelling food. If so, the distal neck has to be straightened between bites to allow the animal to swallow which is not a problem and looks interesting. Perhaps a different problem is that the length of the oesophagus may have to vary with neck position. If the oesophagus lies in front of or below the neck bones, as it does in humans, bending the neck backwards and upwards requires the oesophagus to lengthen. It has to be elastic anyway, so some degree of lengthening should not be a problem. 

Click to enlarge; copyright Katrina van Grauw

There is another solution though. In birds, the trachea (windpipe) and oesophagus do not commonly lie in front of the neck vertebrae, but to one side. This is not the exception but the rule, in fact, as shown by this paper. Most often the trachea and oesophagus lie toon the right side, but that can differ between individuals, with recorded examples on the trachea on one side and the oesophagus on the other. If birds bending their generally long necks, the oesophagus does not have to follow the bones. The oesophagus literally cuts corners. 

Click to enlarge; source here

The two images above show this odd anatomy very clearly, with a heron as an example. One is from the excellent work 'The unfeathered bird' by Katrina van Grauw, and the other is from the paper mentioned above. The oesophagus runs almost in a straight line when the neck is curved, which also means that the oesophagus is then much shorter than when the neck is fully extended. It has to be very elastic. The explanation for this arrangement is that the trachea and oesophagus can move so freely because they are not restrained by muscles as is the case in mammals. Some birds can swallow enormous prey (herons again) which also requires that the oesophagus has freedom of movement. 

Click to enlarge; copyright Gert van Dijk

I am revising an older image of a herd of large hexapods thundering into view. To get a better idea of their heads, I sculpted one in Zbrush. 

Click to enlarge; copyright Gert van Dijk

 

Click to enlarge; copyright Gert van Dijk

 

As you can see, I played with a lateral position of the oesophagus. The oesophagus forms a distinct bulge on the left side of the distal neck, cutting corners in bird fashion. The sensocranium sports shields and horns to impress others of its own species. When the animals start bashing one another's heads and necks, the necks should be protected too, so the middle joint of the proximal neck also bears some shields. So here is the somewhat baroque Latifrons augustus ('elevated broad-brow').

A sample of The Book (and a new painting)

It's time for a quick update. As you may know, The Book is ready to find a home with a publisher. Unfortunately, one or two of the publishers I would be very happy with warn aspiring authors away by saying that they pick their own candidates and have plenty of those to choose from, thank you very much. Others say they want submissions on paper, not digital, which might be a way to reduce the number of proposals they receive. If so, good, as perhaps my proposal may then have fewer others to compete with. Anyway, I had a few samples printed on good quality paper. Here is a quick view of the sample of The Book.

Some of you may have spotted that the name on the sample reads 'Nastrarruzzo', not 'Nastrazzurro'. I had to start an official business to market The Book, (and perhaps sell prints too), and wanted to avoid  possible problems with companies with similar names. That's why my firm is called the 'Nastrarruzzo Compagnie'. 

Click to enlarge; copyright Gert van Dijk

While all this is going on, I do work on the occasional new painting. Actually, there are about four in various stages of completion, but that is usual for me. Here is a fragment of one I just completed. The animal is a hexapod arboreal species, a swobbler (it was originally a 'wobbler', but I made a typo while storing the file, and the unintended 's' made the name more interesting, so I kept it. The animal is not entirely serious...

You may be wondering what the things on its back are. Well, hexapod skins vary a lot, and part of their basic integument is a sort of hair, but hair that is flattened, and often with multiple filaments, so it can range from feather-like structures, but decidedly heavier than true feathers, to a fur-like covering. The structure lends itself well to leaf-like excrescences. As hexapods are not at all restricted to the restricted palette of Earth's mammals, there is room for weird and colourful protuberances. 

The fruit is a baignac, of course. Don’t eat them if you are a human.

A work in progress: the pied stickler

It has been a while since I last posted something here. There were multiple reasons for that: the realisation that there are more than enough paintings for The Book lessened my enthusiasm to start new Furaha paintings. An interesting side effect of that is that I started painting a series of paintings completely unrelated to speculative biology, which was refreshing. So much so, in fact, that having finished three such paintings I became eager to do some more Furahan creatures.

Another factor is that the number of views and replies has been fairly low lately. Is a blog like this one too old-fashioned, or perhaps too complex? No longer interesting? Or is it too difficult to leave a reply? I thought so when I wanted to write something on someone else's blog. Ideas and thoughts are welcome.

 

Click to enlarge; copyright Gert van Dijk

Anyway, here is a detail of a work in progress showing a 'pied stickler' (Perfixor artifex). The name means 'cunning impaler'. A part of the accompanying text follows.

Katarzyna Altanero, who devoted her life to the study of hexapod carnivores, described sticklers as ‘prototypical gregarious piluferentic centauraptors’. This statement shows that doctissimus Altanero was one of those people who, once they have learned something, think that people are born with that knowledge already in place. But the cumbersome jargon is correct. 

'Centauraptors’
These animals are obviously hexapods, but they have freed their front legs from all locomotion duties, a principle known on Furaha as ‘centaurism’. Here, the limbs have become weapons.  
Centaurism probably involves very quick evolution. The front limbs, liberated from walking, were free to quickly evolve suitable shapes for their new roles. At the same time the balance and locomotion of the animal changed drastically. The front part of the trunk became shorter and was tilted upwards, while the former middle legs increased in size and moved forwards.  
   These changes must have evolved hand in hand with specialisation to a specific way of catching prey. For instance, predators relying on speed should not carry massive heavy clubs, and those aiming to bring down armoured prey must have adequate weaponry and need only be faster than their prey. Scavenging centauraptors can be slow , allowing some of the heaviest weaponry of all.   

‘Piluferentic’
Centauraptor weapons betray an interesting array of forms. Some are pure clubs, others function only as ‘pointy end’ weapons, and in yet others the limb has more than one purpose. The limb type known as ‘axes’ have a sharp edge on the underside, useful to hack open a carcass, but the same weapon usually has a heavy bulge too, allowing it to double as a club to knock prey off their feet or to bludgeon the prey's head, blinding it.
   The stickler’s weapons are no exception to this ‘Swiss army knife’ approach. While its weapon is primarily shaped like a club, the former foot has become a sharp protrusion that can be swung into action. This sting explains the word ‘piluferens’, meaning 'lance carrying’.   

‘Gregarious’
As the word suggests, sticklers live in groups. In this species’ case, tasks differ between group members, allowing a fairly complex society that must run on advanced cognitive capacity. Do not underestimate sticklers; one may appear to study you with open curiosity, but its pack members may well be behind you, on their way to encircle you. They cannot digest humans but that makes little difference to the human involved.       
           

More alternate evolution in Japan: "What if extinct organisms continued to evolve?"

I am always interested in speculative biology projects, in particular those with well-done art. For some reason I was looking at the Japanese website of Amazon; I think I was looking at the Japanese edition of 'Demain; les animaux du futur', a French book I wrote about here and here. It is about life on Earth some 10 million in the future. As you may know, Amazon shows you lots of other books they hope you will buy. In this case it worked: one caught my attention, so I copied parts of the text into Google Translate so I would get a better idea what it was about. The translated title was: "What if extinct organisms continued to evolve?"

Click to enlarge; copyright Ken Tsuchiya, Masato Hattori

After looking at it a few times I gave in to temptation and bought it. The book was not that expensive, but postage from Japan to Europe more or less doubled the price, so check that first if you are interested. Thanks to Google Lens, I could point my phone at the pages and received an overlaid English translation. I you try that several times, you get different translations, so there is a 'Lost in Translation' sense here. I tried to contact the main author, Ken Tschuchiya, who is a geologist and palaeontologist as well as a science writer but was unable to reach him. However, the illustrator, Masato Hattori did respond, so I asked him a few questions. Unfortunately, but understandably, he redirected any enquiries regarding the reasoning behind the beasties to the writer. That left me to think of possibilities for myself, which is less certain but fun. I had a very similar experience looking at Dougal Dixon's Greenworld in Japanese. 

I asked Mr. Hattori how he made the images, guessing that he used ZBrush and Photoshop, which indeed he did. He modelled and rendered the animals in ZBrush and used Photoshop to blend the images into photographs. It is tricky to get the lighting, angles and contrast right, but Mr. Hattori certainly knows his business. Have a look at his own website here; there is more alternate evolution to see, in the form of dinosaurs running alongside present-day humans. 

Mr. Hattori was familiar with other works of speculative evolution, such as 'After Man' by Dougal Dixon and 'Demain, les animaux du futur' by Jean-Sébastien Steyer and Marc Boulay. The latter work is still not available in English, I think, but there is a Japanese version (among other translations). Mr. Hattori said he has a collection of such works and was in no small way influenced by them. 

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Let’s have a look at the book. Mr. Hattori said that the setting examines how archaic organisms would have evolved if they had not become extinct and had survived to the present day. In that respect it differs from 'After Man' and 'Demain', both of which describe Earth in the future. 

The cover of the book is shown at the top of this post; I like the composition and lighting very much. We are obviously looking at ammonites. I wonder if they differ much from their ancestors. Should we expect them to have diverged in shape given the very long time since their demise? Or would there be present-day ammonites that look just like their ancestors, as these seem to do? 

 

Click to enlarge; copyright Ken Tsuchiya, Masato Hattori

This is an obvious sea scorpion, again not looking that different from its ancestors. The world really needs more very large arthropods (I know, I know; they’re difficult...). I wonder whether such animals would have managed to compete successfully with sharks or bony fish for a long time. They don;y seem to have done so in our timeline. Such thoughts led me to think at first that the theme of the book was that the animals lived for a while beyond their extinction time, without necessarily making it all the way to the present. I also do not think that the authors meant for all animals to be alive at the same time. The book is arranged by geological era, with for some animals a point of departure at the end of the Permian. Later in the book we see alternate mammal evolution starting from recent ancestors, that would not have been there if the Permian extinction had not occurred. Each animal seems to be its own alternate history project. Anyway, I really like the energy of the image and the skilful use of iridescence. That is not an easy effect to achieve, I can tell you from experience. 

 

Click to enlarge; copyright Ken Tsuchiya, Masato Hattori

This one is decidedly funny. It is a caricature of Diplocaulus, the famous Carboniferous/Permian amphibian with a very odd 'boomerang' skull. I wondered how broad the lower jaw of Diplocaulus was, so I tried to find accurate anatomical drawings or photographs of Diplocaulus' mandible but couldn’t find high-quality images. The mouth seemed to be small though. According to the book, the extinction at the end of the Permian allowed a descendent to develop in a different manner. Young diplocaulids had no broad 'horns', but these developed as the animal grew. In this descendent, the moth broadened with growth too, resulting in a mouth that is as wide as the skull. It also seems to have gone for complete flattening of the body. Another image shows it lying on a lake floor, opening its mouth. I guess it might suck in prey that way. 

 

Click to enlarge; copyright Ken Tsuchiya, Masato Hattori

Here we have an animal looking very much like a river dolphin; it even has a 'melon', that bulge on the forehead that apparently helps whales produce sonar. But this is no whale: it's got scales! What is this animal descended form? 

Click to enlarge; copyright Ken Tsuchiya, Masato Hattori

Turning over the page reveals its ancestral line: it’s a spinosaur descendent! You may be familiar with the recent discussion about how well spinosaurs were adapted to swimming: some said they are, countered by some who say they aren’t. The authors of the If-book decided to let it take to the water. I wonder about the melon: could, and would, spinosaurs have developed sonar? I do not know enough about the starting point of hearing in mammals and dinosaurs to have an opinion on that, but it is a nice idea. So here we are: a 'river spinophin'. 

Click to enlarge; copyright Ken Tsuchiya, Masato Hattori

Here is the last example of what the book has to offer: this is obviously a predatory dinosaur, but not an agile one at all. Its genus name 'Megapubis' means you should take a good look at what it has between its legs: yes, that is a massive pubic bone, and it is sitting on it. Also look at the nearly columnar legs, only a bit longer than the pubis. This beast must spend its days resting and doing not much. It can’t even scratch itself, as its arms have atrophied completely: the species name 'acheirus' means 'no hands'. The animal is a scavenger, happy to wait until some animal has the decency to keel over dead. 

I will leave the rest of the book to buyers. There are 25 species in all, with quite a few reconstructions and additional images. The book is great fun, and the images are well worth looking at closely. The text seems quite interesting too; a pity I can't speak Japanese. Will there be an English version? Mr. Hattori would like that, and so would I,  but he wasn't aware of translation projects at present.

Avatar 2: The Way of Water II: whales, gill mantles, corals...

In the previous post on 'The Way of Water (TWOW) I discussed the skimwing, and had to conclude that it couldn’t work as an enlarged flying fish because of scaling effects. Simply said, they overdid it and made it impossibly large. Making animals too large is a recurring theme on his blog. The underlying reason is probably that the effects of scaling on mass and weight are not intuitive at all. Speaking for myself, even after years of considering animal size, I still sit back often and think that 'That is too much and can’t be right…'. For example, would you have guessed than making a man twice his original height of 1.75 meter would increase his weight from 80 kg to a staggering 1440 kg?    
 

But anyway, today's species will not involve any calculations at all. Let's have a look. 

 

Click to enlarge; copyright 20th century studios

The tulkun
The tulkun looks so much like a whale that we will just call it that here. To quote from the book ‘The Art of Avatar TWOW' by Tara Bennett: “… it’s very much written like a whale and functions like a whale. But how whalelike do you make it?” 

Click to enlarge; copyright 20th century studios

Well, apparently some early designs were not whale-like enough, and the book offers some glimpses what might have been. Here's one. I would have preferred something more alien, but the audience would probably not have connected the resulting much more alien shapes  with the emotions associated with whales on Earth (I am referring to the sympathy and enthusiasm for whales in most of the West, not the sentiment "Let's have a big whale steak!"). The Avatar film makers obviously intended to evoke whale enthusiasm. I like whales too, but felt that the film makers overdid it when tapping into sympathy towards whales. This particular whale species happens to have a clear yellow fluid in its head that stops human ageing, serving as background to have a scene in which villains enthusiastically harpoon the whales to get that mystery substance.

Do we need to discuss the plausibility of an alien biochemistry producing a substance for its own good that by an apparent coincidence also just happens to stop human ageing? Human ageing does not appear to be a process depending on just one simple biochemical trigger (although that cannot be dismissed with complete certainty). If there would be one, you can expect a massive effort to identify the substance, and shortly afterwards it could be manufactured on Earth simply and cheaply. Let's also not discuss how you would find out that some fluid inside an alien animal's head stops human ageing. Did someone stumble upon this liquid and thought "Let's just see what happens if I inject myself with this"?

The tulkun is supposed to be extremely intelligent: it appears capable of understanding the spoken language of a completely different species. That's quite something! Humans take a very long time to learn another human language, but maybe we are just more stupid than Pandoran whales. But even the brightest whale could only extract meaning from spoken words after gathering a massive amount of data to correlate specific sounds with objects, questions, tenses, nouns, verbs, etc. etc. Another big question is how and why intelligence would evolve in such a being. I personally do not think that you need hands or something similar to develop intelligence. I always rather liked the view that social interactions played an important role in evolving human intelligence (including gossiping), and perhaps such interactions were important for these whales too.  

Anyway, the tulkun looks like an Earth whale. So much so that I do not doubt it would function well as an organism, except for one thing: why is it not streamlined? It can obviously move fast, so streamlining is not a luxury, but a necessity. The animal has a rough texture with  grooves and ridges. There is also its headgear that doesn’t help streamlining either. Are these details there only there to make us think that it’s a whale, but not one of ours? If so, I am a bit disappointed. Streamlining fast aquatic animals should be compulsory to achieve some biological believability.

 

Click to enlarge; copyright 20th century studios

The gill mantle
The gill mantle is an animal that latches onto the Metkayina (partially aquatic Na’avi), meaning Pandora’s native humanoids. The gill mantle allows them to breathe underwater. That is quite a feat. If the Matkayina’s metabolism is like that of Earth mammals, they need lots of oxygen and a way to get rid of CO2 (and, seeing they can dive deeply, ways to deal with pressure to avoid the bends). The gill mantle can do all that, it seems, which is quite a feat. The problem here is that gas exchange needs a very large area, usually obtained by folding the gas exchange surfaces right down to the microscopic level. For instance, estimates of the total area of human lungs amount to 100 square meters. But the mantle looks smooth; I don’t see any kind of subdivision offering the requested area, nor does it transparent nature suggest that the mantle houses the rather considerable blood flow needed: in humans, the same amount of blood flows through the lungs as through the rest of the body, meaning some 4 litres per minute.
But the real tricky question is why any animal would evolve such a feature? How does it benefit from this ability and how did its evolution start? 

Click to enlarge; copyright 20th century studios

Coral
You might be surprised at my choice, but the image in the TWOW book that caught my attention longer than all other ones was not one showing a big spectacular animal, but a coral fan concept by Jonathan Bach. Here it is (page 191). It is very well wrought. The connections between its branches caught my eye.  The ends of the arches are merged, forming a complete connected structure. This means that the arches do not form the typical branching structure familiar to Earth life, from lungs to corals and oak trees, but a net. The easiest way to form a biological net is probably to start with a sheet, or a series of planes in 3D space, and then to make holes in it, leaving just the arches. However, plants and Earth corals grow branches with ends, and branch ends do not typically merge if they touch one another. There is some evidence that Earth trees can do that though, and I designed Furahan thorn shrubs that did so, resulting in a truly impregnable wall of thorns. I have not used that design yet, because I felt that little hooks would work just as well and would not require an organism to open its tissues to the outside world. 

Click to enlarge; copyright 20th century studios

The ilu
I am not yet certain what to think about the ilu yet. It looks a bit like a plesiosaur with a small head and a long neck. But whereas plesiosaurs have slender fairly stiff fins with rounded narrow tips, as do turtles, seals, and mosasaurs, the ilu has very long somewhat floppy fins that appear to get broader towards their ends. Are there Earth analogues for such a design? If not, is there a good reason for that? I'll have to think more on that, so perhaps there will be third post on TWOW.

So, what did I think of the animals in TWOW? I felt the same as I did about the first film in the series: the images were very pretty. I will watch the film several times and enjoy it every time. Still, I again felt a bit disappointed about the lack of biological plausibility. Would the audience really object to more plausible or more alien shapes?   

     

A hexapod muscle study

Some time ago I wrote about the Great Hexapod Revolution, which I might also have called the Quite Considerable Hexapod Revision. At any rate, that particular revolution or revision has conceptually been completed for quite some time, so I am busy revising old paintings as well as coming up with new ones. The latter are not really necessary as The Book is basically done.

Yes, The Book is done.

Basically.

The 'done' part means that there are easily enough double pages to fill a book, so rather than producing more I am shifting emphasis towards finding a publisher. I do not expect immediate success, which explains 'basically': I might meanwhile just as well keep on thinking about Furahan creatures.

Let's review the hexapod revision. Their skeleton still reflects that of early representatives of the clade. There is no vertebral column, here defined as many short similar bones placed end to end running from front to back in the vertical plane dividing left and right halves of the animal (technically, the sagittal plane). The most bare bones version of that skeleton (sorry for that one) would remind you of a foldable ladder. As the clade started with animals without legs, it would be strange to use the name Hexapods ('six-leggers') for all of them. I chose the new name 'Scalata' instead, based on the Latin word 'scala': ladder. The word 'Scalata' is technically correct while 'scalates' is suitable for colloquial use. Hexapods then become a subgroup, consisting of scalates with legs. 

Click to enlarge; copyright Gert van Dijk

The next big step was deciding the shape of the legs. I built on the zigzag principle, in which successive major leg segments bend one way at the topmost joint, the other way at the second joint, and so on. By reversing direction, no joint is ever really far away from a line perpendicular from the hip down. Being close to that line reduces the force needed to keep the joints in those positions, meaning muscle power. have a look at these posts here and here. The image above was taken from these earlier posts and explains that principle.

The least force to keep the segments in place is needed when all segments are stacked vertically, making the leg into a column. That is a fine way to conserve energy but does not produce athletic animals. Vertical leg bones are typically found in large non-athletic animals: think of elephants and sauropods. In smaller animals all segments can be closer to the horizontal than the vertical position, because fighting gravity costs relatively much less (for scaling effects, see here and here). The actual position of the leg bones will depend on mass and athleticism.
             
I suggested in earlier posts (here and here) that it wouldn’t really make a difference whether the legs started by being angled forwards ('zig') or backwards ('zag') at the hip joint. Mammals are peculiar in having their front legs start with a zag and hind legs with a zig. This is a consequence of how they re-engineered their original sprawling posture: front legs rotated backwards, with elbows pointing back, and hind legs forwards, with knees pointing forwards.

Should that reversal be seen as a natural 'law' or as an evolutionary coincidence that became locked in place? I could not think of any physical reason for this pattern and so had freedom to decide what to do with scalate legs. All three pairs of legs underwent the same rotation, which is simple and keeps them out of each other's way. The top segments all point backwards.   

 

Click to enlarge; copyright Gert van Dijk

Another decision was how to join the legs to the scala. Should the hip joint allow movements in all directions, or should they restrict movement in one or more directions? Should the joint be so 'open' that all positions need to be controlled by expensive muscle activity, or do we let bones and ligaments take up some of the stresses? I decided to give the joint surface a 'roof' in the hip to push against, transferring weight. The image above shows three possible patterns: in A, the bone sits directly underneath the spherical joint, allowing three-axial rotations and simple weight-bearing. In B, the joint does the same, but the shaft of the bone is shifted a bit to the side, allowing room for gut, eggs, or whatever. In C, the joint restricts rotations around the axis running down the bone and the bone extends a bit past the joint. That sturdy upwards spur can be used to attach muscles to; that's the hexapod hip joint.  

The main propulsion force involves swinging the upper leg segments thighs backwards: retroflexion. The thigh has a limited range of motion, from an angled pointing just a bit forwards to a much larger backwards angle. To work over that range, hexapods have one large muscle starting behind the joint and attaching to the hip bone below the joint, exactly like human buttock muscles. But another big muscle originates in front of the hip and inserts on the spur above the joint. These two muscles act in concert to pull the leg back: they are 'agonists'.

That range of motion has consequences for where muscles can produce the most force. The force exerted by muscle fibres is most effective if these fibres make a right angle with a line from the insertion site to the axis. If that angle is not 90 degrees, only the component of the force that is at a right angle is useful to rotate the bone; the remainder just presses the bone into the joint or pulls it out of it. When the bone rotates, the effective force component changes with the rotation angle. You would want to place a muscle in such a way that most muscle fibres do useful work over most of the movement range. 

 

This animation shows a muscle placed to the front (at left) of the bone, inserting at the spur above the axis of rotation. The bone rotates through its working range, which is shown three times. The four  panels show fibres at different sites of origin. The red lines show the parts of the force that do the actual rotation. It is obvious that fibres starting high above the spur are not much use and can even pull in the wrong direction. The most useful fibres start at the level of the joint or lower, so this is where the muscle should be. I have not shown the other muscle, the one pulling on the thigh below the axis while starting behind the joint (at right). The principle is the same, but now the most useful part lies at the top.      

As I like Latin anatomical nomenclature for its simplicity, I named these muscles ('simple' is here based on the premise that the names are in another language you have to learn anyway). For the front legs, the front muscle is the musculus retractor artus primi anterior, and the hind one is the m. retractor artus primi posterior. For the middle and hind legs, replace primi with secundi or tertii.
 
There are of course muscles that work in the other direction, the 'antagonists', but these are weaker and run the other way, lying underneath the big 'retroflexion' muscles.

Click to enlarge; copyright Gert van Dijk

Click to enlarge; copyright Gert van Dijk

The two images above shows the result of some experimental ZBrush sculpting. Here you see a general hexapod with some main muscles shown. I still find ZBrush extremely non-intuitive, but am very slowly feeling my way around it.

This animal is probably the size of a horse. Note that the middle legs are sturdier than the others. That is because that is where most of the mass is! The middle legs are also wider apart, to allow room for a possibly sizeable gut and also for the front and hind legs. Of course,  the scheme underwent substantial changes in particular with those predators that freed their front limbs from locomotion ('centaurism') and turned them into weapons. maybe I’ll show those anatomical changes too, one day.           

Avatar 2 The Way Of Water, or do skimwings tip the scale?

Many readers will by now have seen the second film in the Avatar series, 'The Way of Water' (TWOW). I felt that the story of the film resembled that of the first film a bit too much, but never mind that; this is not a blog about cinema, but about speculative biology. Luckily, TWOW offers new species to enjoy, to watch again and to think about. I used the book ‘The Art of Avatar TWOW' by Tara Bennett to write this post. There will probably be two posts on TWOW; this one will be about the skimwing, chosen because it has a lot to offer from a biomechanical point of view. It  will be a long post and there are equations at the end, so you have been warned...

That book contains quotes and explanations that confirm a conclusion I had drawn from viewing the first film, and that is that the shapes and form of the life forms on Pandora  are primarily governed by audience appeal, with biological plausibility taking a definite second place. I deplored that second place when I wrote my post about the first film, posted 13 years ago to the day. I still do, but now accept that the people in charge of making films think this is what the audience wants. They may be right; but I, and I guess many readers of this blog, are not typical in this regard. We like our science well-done, not rare. Relegating plausibility to second place is acceptable as long as film makers do not claim that the life forms they present are biologically sound. I got the impression that they did make that claim for the first film, but for TWOW the book acknowledges that the director's opinion of audience appeal came first. 

Click to enlarge; copyright 20th century studios

The goal: can we answer the question whether skimwings can 'taxi' as shown?  

 The skimwing is basically a gigantic flying fish: it has a long slender body and two fins that double as wings. Like flying fish, the skimwing is fast enough to partially leave the water, with just the tail in the water to propel it. Flying fish use this stage to accelerate and leave the water altogether, but skimwings do not do that; they just taxi along.
    Could the skimwing as shown really taxi in the way flying fish do? That is not an easy question because of all the factors that are involved. Let's consider the problems.
 

1. While taxiing, the animal's tail provides thrust propelling the animal forwards, but the tail may also produce some upwards force. If so, the wings need to provide less lift than if all the upwards force is due to lift. In this post, I will assume that the tail only provides forward thrust, so the wings are responsible for all the upwards force. But I will get back to this matter in the end. 

2. As for lift, we will use conventional equations. See this post for an introduction. We are not on Earth, so lift is altered by the higher atmospheric density on Pandora, where the story takes place (1.2 times that of Earth).

3. The animal's weight is also other than it would be on Earth because the gravity constant of Pandora is lower than the one of Earth (it is said to be 0.8 of that of Earth). 

4. A more difficult effect to deal with is the 'ground effect', which means that flying objects (animals and aircraft) experience extra lift if they fly close to the ground. The literature makes it clear that flying fish use the ground effect, and the book states that skimwings do so too. 

 

Click to enlarge; copyright 20th century studios

5. Finally, there is a literally enormous difference between skimwings and flying fish: flying fish are only about 35 cm long and I estimate skimwing length to be 12.2 to 13.5 meter. I derived that estimate using the image above, in which the skimwing appears to be 4.5 times as long as an adult Na'vi. The internet tells me that Na’vi are 2.7 to 3 meters tall. I used a skimwing length of 12.5 m as a reasonable estimate.      

I am not an aeronautical engineer, so what follows should be seen as nothing more than a layperson's attempt to understand how all the above factors might work together.  But first, what do we know about flying fish?     

 

Click to enlarge; from Wikipedia

Flying fish can swim, 'taxi' and glide
 
Flying fish are often said to take to the air to escape predators (Socha 2015). That may well be true but has not been proven; other animals jump out of the water for a variety of reasons. Squid may glide above water to escape predators too but also to catch prey (Socha 2015);  whales apparently breach to impress other whales, while dolphins may do so to achieve greater overall speed. As jumping out of the water must take a lot of energy this seems a strange way to save energy, but moving through air obviously offers less resistance than through water, so that gain may offset the additional energy needed to leave the water. 

There is another odd effect going on here, and that is that swimming just below the surface of the water costs more than swimming in deeper water. Why? Well, all swimmers push water out of the way to make room for themselves, but in shallow water some of that displaced water moves upwards, which costs more energy than if the water only moves sideways. That additional energy cost may just tip the balance, making leaving the water more efficient than pushing all that water up (Socha 2015).
    
Taxiing
Flying fish accelerate under water, break though the water's surface, and then accelerate some more with only their tail underwater, from 10 to 20 meters per second (Socha 2015). That's from 36 to 72 km/h. Calculations show that they need 350 Watt to swim underwater at 10 m/s, but only 36 Watts while taxiing; taxiing is therefore quite efficient (Deng 2019). So far so good!
    Flying fish then leave the water and glide, held aloft by their two or four wings (some species use both pectoral and pelvic wings, others just the pectoral wings). Although some papers describe the flight path as relatively flat (Fish 1989, Socha 2015), meaning at a constant height above the surface, this can only be an approximation. The reason is that flying fish have no propulsion while in the air, so the glide is either completely passive or else lift is helped in some way. In a true passive glide losing height is physically unavoidable (Socha 2015).
     This suggests that their lift is indeed helped. Updraughts can help (Fish 1989) and so can the ground effect. In fact, the relatively flat trajectory itself suggests that the ground effect does help (Socha 2015). Wind tunnel experiments with stuffed flying fish (Really? Yes, really) showed that the ground effect reduced drag (let's say that ‘drag’ is the force impeding forwards motion) by 14%. As lift stayed the same, the so-called lift-to-drag ratio was improved, which is not bad at all (Park 2010). Remember that number of 14%, because we will need it later.
     As flying fish know their physics they do come down to the water and may repeat the procedure: taxi, glide, land, etc. They may cover distances of some 400 meters in 30 seconds (Park 2010), suggesting a mean speed of 48 km/h.       
    
Gliding
The wings proved to be like bird wings, in particular as regards wing loading. 'Wing loading' is calculated as the area of the wings divided by weight of the animal, so it tells you how much kg a square m of wing carries. A low value makes flying easier. The wings of flying fish  are designed for high lift and low drag (Fish 1989). The four-winged species have lower wing loading, pointing to increased lift at low speeds, than the two-winged species.

Scaling flying animals

Now it gets more complicated. But not that much, so hang on! I have discussed scaling winged animals before, but I will repeat the main thoughts here. What we will do is to take Earth’s flying fish, scale them up, export them to Pandora to accommodate the different atmosphere and gravity, calculate their weight there, and then see whether they can produce enough lift to keep that weight aloft in Pandoran air.
   
Lift is proportional to just three relevant parameters (there are two more: angle of attack and a constant, but if we keep these the same throughout we can ignore them).

• density of the air (D), in kg per cubic meter
• area (A) of the wing, in square meters
• square of speed (S^2), in meter per second (I cannot use superscript, which is why I used '^2' to indicate a squared speed)

LIFT = D x A x S^2

If a flying animal flies at a stable height and does not sink, the amount of lift it generates must equal its weight. We can calculate weight W as the product of the gravity constant G and the mass of the animal M.

WEIGHT = G x M

So remember this for stable flight:

LIFT = WEIGHT

Now we need another look at scaling; (see here for flying and here and here for an introduction). Suppose we take an animal with length L and make its length, height and width all twice as large as before (that's called 'isometric scaling'). Its length becomes 2L. However, the area of the wings is the product of length and width of the wings, and as each became twice as long, the area becomes four times (2x2) as large. However, the volume of the animal has three dimensions, so that becomes eight times (2x2x2) as large. The mass corresponds to the volume and also becomes eight times as large. The lesson here is that mass increases more than area, and that is a problem. 

Click to enlarge; copyright Gert van Dijk

Weight becomes eight times as large, but the four times larger wing area will only get you four times the lift. To fly stably, lift must equal weight, so we must find a way to achieve eight times the original amount of lift. In the scheme, above, the first way to do so is labelled 'Enlarged flying fish 1'. In that option, squared velocity was made twice its original value. To do that, velocity itself needs to become 1.4 times larger (1.4 is about the square root of 2). Hence, the larger animal has to fly faster if its wings stay in proportion with the body. But to land and take off, the animal needs to be able to fly at low speeds too: you cannot fly fast all the time.

Is flying faster the only solution? No, we can also choose to have the enlarged animal fly at the same speed as its predecessor ('Enlarged flying fish 2'). We still need to achieve eight times the lift, so we need to make the wing area eight times larger. If we make the length and width of the wing each 2.83 times larger (2.83 is about the square root of 8) we get that. Mind you, this solution will reach a dead end at some point because the enlarged wings will also add weight, which needs to be lifted, etc. I discussed how much weight such larger wings will add to the animal in an earlier post (here); it is dramatic! 
         
To conclude, we have a choice of achieving more lift for the larger animal by either enlarging wing area or flying faster; either solution will reach a limit at some point. On Earth, larger birds combine larger wing area as well as higher speed to achieve that higher lift. That combination can only be an evolutionary compromise between the costs of high speed (difficulty in taking off and landing) and large wings (more weight). There must be a physical limit where flight is no longer feasible for an animal. Does scaling up the skimwing tip the scales? (Sorry for that one, but I could not help myself) 

The case of the skimwing

The length of a skimwing is at 12.5 meter 35.7 times larger than that of a flying fish. If we multiply the dimensions of a flying fish by 37.5, its wing area becomes 1275 times larger and its mass becomes about 45,500 times larger. Wow!

We now need to do an analysis as explained above, but the animal gets to be not twice the size, but 37.5 times. The results of this thought experiment are in the appendix, for those who want numbers. If we choose to scale the animal isometrically, meaning that the proportions of the animal stay the same, then it must taxi not at 20 m/s (72 km/h) as the flying fish manages to do, but at a staggering 351 km/h. That is wholly unrealistic; for one, the Na'vi sitting on top would be blown off…

How about the other approach, meaning making the wings larger? Well, isometric scaling made the wing length and width each 35.7 times larger. It turns out that we need to make each 174 times larger instead! I did not bother calculating how much weight that would add.

Save the skimwing!

But the animal might derive part of the upwards force from beating its tail. True, but there is a good reason why tail walking dolphins are not large. It is, once again, scaling: the force needed to push against the water depends on the cross section of the muscles, meaning area, and needs to equal the animal's weight. It is the exact same problem as before: force increases with the square of length and weight with the third power. Turning to tail walking instead of gliding trades one unsurmountable scaling effect for another.

But how about the ground effect? Well, in flying fish that reduced drag by 14%. While that is not the same as increasing lift, it may be treated that way. The appendix shows that a gratis 14% increase in lift still do not result in a viable animal, and neither did halving the mass of the skimwing: that won’t fly (sorry for that one too).

Conclusion
 
What a pity. It seems that the designers did not realise how much that third power puts a brake on scaling up animals. Admittedly, those effects are not immediately obvious, but it's not rocket science either. The simple conclusion is that 12.5-meter-long taxiing flying fish are too large to work, even under Pandora's favourable gravity and atmosphere.

Does it matter? The science in TWOW seems fairly typical for how Hollywood treats science, meaning with rather limited respect for plausibility or accuracy. That's not good news; actually, it's not news at all. However, there is good news: the film shows a very profound love for the natural world. If that helps make people care about nature, I'm all for it. 

As far as that love for nature is concerned, consider this: the skimwing differs in only one important aspect from flying fish: its size, and it is exactly that difference that makes the skimwing impossible. But everything else that makes Pandoran skimwings fascinating was already fascinating about flying fish, right here on Earth.     

Acknowledgement

Abbydon made insightful comments on a first draft of this post. He also added remarks about wave effects that would make using skimwings as public transportation rather unreliable.   

 
Selected references
FE Fish. Wing design and scaling of flying fish with regard to flight performance. J Zool Lond 1990; 221: 391-403
Park H, Choi H. Aerodynamic characteristics of flying fish in gliding flight. J Exp Biol 2010; 213: 3269-3279
Deng J, Wang S, Zhang L. Why does a flying fish taxi on sea surface before taking off? A hydrodynamic interpretation. https://doi.org/10.1101/765560
Socha JJ, Jafari F, Munk Y, Vyrnes G. How animals glide: form trajectory to morphology. Can J Zool 93: 901–924 (2015) dx.doi.org/10.1139/cjz-2014-0013  
Sullivan TN, Meyers MA, Arzt E. Scaling of bird wings and feathers for efficient flight. Sci. Adv. 2019;5: eaat4269 

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Appendix

 

Abbreviations and parameters

            FF                    Flying Fish

            SW                   SkimWing

            Length             L

            Area (wings)     A

            Volume             V

            Mass                M

            Weight             W

            Speed               S

            Density of air    D

            (local values for Pandoran SW and Earth FF)

                                    D_SW = 1.2 ∙ D_FF

            Gravity constant G (local values for SW and FF)

                                    G_SW = 0.8 ∙ G_FF

            Lift                   LIFT

 

----------------------------------------------------------------------

 

Take the length of the animal of the FF as base value of 1; the SW is 35.7 times longer.

            L_FF = 1

            L_SW = 35.7 ∙ L_FF

This results in the following for area and volume for SW:

            A_SW = (35.7)² ∙ A_FF         or         A_SW = 1275 ∙ A_FF

            V_SW = (35.7)³ ∙ V_FF        or         V_SW = 45500 ∙ V_FF

For weight we get:

            W_SW = V_SW ∙ G_SW                   Pandoran values

            W_FF = V_FF ∙ G_FF                        Earth values

For lift we get:

            LIFT_FF = D_FF ∙ S_FF² ∙ A_FF

            LIFT_SW = D_SW ∙ S_SW² ∙ A_SW

We can now start filling in values for Pandora using Earth parameters, not altering wing area A and speed S on Pandora yet:

            for lift:              LIFT_SW = (1.2 ∙ D_FF) ∙ S_SW² ∙ A_SW                     

            for weight         W_SW = V_SW ∙ G_SW

                                    W_SW = V_SW ∙ (0.8 ∙ G_FF)

                                    W_SW = (45500 ∙ V_FF )∙ (0.8 ∙ G_FF)

                                    W_SW = 36400 ∙ V_FF ∙ G_FF

                                    W_SW = 36400 ∙W_FF

If the flight is stable, weight must equal lift, so it follows that:

            LIFT_SW                                     =          36400 ∙ LIFT_FF

            (1.2 ∙ D_FF) ∙ S_SW² ∙ A_SW     =                 36400 ∙  D_FF ∙ S_FF² ∙ A_FF

             S_SW² ∙ A_SW                                                        =                 30333 ∙  S_FF² ∙ A_FF

 

We have SW parameters on the left and Earth FF parameters on the right and can play with this relationship.

 

Variant 1 assumes isometric scaling, making  A_SW = 1275 ∙ A_FF

             S_SW² ∙ 1275 ∙ A_FF                  =                 30333 ∙  S_FF² ∙ A_FF

             S_SW²                                                 =                 23.8 ∙  S_FF²

We know that the speed of FF = 20 m/s, so

            S_SW²                                                   =                 23.8 ∙  400  = 9516

            S_SW                               =          97.6 m/s or 351 km/h           

This is obviously a ridiculous speed and cannot work. Perhaps larger wings may do the trick if we abandon isometric scaling.              

 

Variant 2 starts a few steps back.

            S_SW² ∙ A_FF                                      =                 30333 ∙  S_FF² ∙ A_FF

If we assume that the skimwing has the same speed as Flying fish, then  S_SW² = S_FF²

            A_FF                                                      =                 30333 ∙ A_FF

This means that the width and lengtn of the wing have to increase by the square root of 30333, meaning they become 174 times larger instead of the original 35.7 times. This is ridiculous, and the real situation would be worse because the arger windg wopuld weigh a lot more, which he haven’t accounted for yet!

 

Are there ways out?

 

The ground effect seems to reduce drag by 14%. That is not the same as increasing lift by 14% but assume that lift is indeed increased by this amount. 

            S_SW² ∙ 1275 ∙ A_FF                    =                 0.86  ∙ 30333 ∙  S_FF² ∙ A_FF

            S_SW²                                                   =                 20.5 ∙  S_FF²

            S_SW²                                                   =                 20.5 ∙  400  = 8200

            S_SW                               =          90.6 m/s or 326 km/h           

or

            A_FF                                                      =                 26086 ∙ A_FF

The wing's length and width have to become 161 times larger. Ground effect does not save the skimwing, and that holds for an isometric as well as for a non-isometruc approach.  

 

What if we make the skimwing relatively more slender than the FF, by halving the volume of its body? This results in:

            S_SW² ∙ A_SW                                    =                 15167 ∙  S_FF² ∙ A_FF

            S_SW² ∙ 1275 ∙ A_FF                    =                 15167∙  S_FF² ∙ A_FF

             S_SW²                                                 =                 11.9 ∙  S_FF²

We know that the speed of FF = 20 m/s

            S_SW²                                                   =                 11.9 ∙  400  = 4758

            S_SW                               =          69.0 m/s or 248.3 km/h           

If we assume that the skimwing has the same speed as Flying fish, then  S_SW² = S_FF²

            A_FF                                                      =                 15167 ∙ A_FF

This means that the length and width of the wing must become 123 times larger instead of the original 35.7 times. This is still preposterous.