Rusps turn out to follow biological rules about the weaponisation of tails

I recently came across an interesting paper on the evolution of the use of tails as weapons in Earth animals. This turns out to be a fairly rare occurrence, and perhaps that rarity helps explains why animals with tail weapons are so spectacular. After all, we take the common for granted, and it is the departure from the common that attracts attention.

The glyptodont Doedicurus; click to enlarge. https://en.wikipedia.org/wiki/Doedicurus

A nice example on an animal with a tail that is obviously useful as a weapon is the glyptodon genus Doedicurus, a giant armadillo-like mammal, the size of a small car. Doedicurus was encased in strong armour and endowed with a tail with an impressive thickened club at the end.

Click to enlarge; Pinacosaurus Grangeri; Copyright Gregory S. Paul. Princeton field Guide to Dinosaurs, second edition

Ankylosaurs had the same idea, but much earlier. As far as their design was concerned, they went overboard in adding an array of large sharp spikes to their armour.

Click to enlarge; Spinophorosaurus nigerensis; Copyright Gregory S. Paul. Princeton field Guide to Dinosaurs, second edition

Some sauropods may also have had body armour as well as similar thick knobs on the end of their tails. Only one sauropod (Spinophorosaurus nigerensis) apparently sported pointy spikes on its tail, shown here as a juvenile, and drawn by Gregory Paul (I do not think I have to urge dinosaur enthusiasts to get his book 'The Princeton field guide to dinosaurs'). If these long tails were swept at high speed, the transfer of all that kinetic energy should do some real damage. But perhaps a simple threat, along the lines of 'Make my day, punk' would be enough to prevent an actual fight.

The paper in question has the title "The evolution of tail weaponization in amniotes" and is written by Victoria Arbour and Lindsay Zanno. The paper describes which features are the evolutionary precursors of the evolution of tail weapons. The authors performed a thorough statistical analysis of many body traits, and looked separately at four aspects of tail weaponry:  tail lashing, bony terminal tail spikes, a stiff distal tail, and an expanded tail tip.

Click to enlarge. Arbour & Zanno 2018

Here is a figure of the paper, showing these four aspects and the features they are associated with. The result of all this is that you are not likely to find tail weaponry in agile quick-footed predators. If you were designing just such an animal for your speculative biology project, you should probably pause to consider its likelihood. Tail weapons seem to be a last resort for large slow herbivores who already invested in body armour. The authors make the point that equipping heads with weapons occurred much more often. This seems odd because heads are already filled with important structures that should not be damaged, whereas damage to a tail is probably much less risky, so you would expect 'anterior armatification' to be less common that 'posterior armatification'(I could not resist latinising 'weaponisation'). The authors do not speculate why this should be so, but I wonder whether the effective use of weapons requires excellent motor control, something that in turn depends on excellent sensory control, meaning sight. If so, the animal's body may simply be in the way, so it cannot see well enough where to place the sting in its tail.
   At any rate, the authors state that armour in mammals evolved in those animals that are neither small enough to hide nor large enough to deter predators by size alone, and that live in open environments. Close combat with a predator must be a risky business, so the best strategy may simply be running away faster than a predator. And if flight is your main strategy, heavy armour is not going to help. But  a wholly new set of constraints must come into play if you have no chance to outrun your predator to start with. Defensive features such as large size and armour then may become useful, and it seems that active weaponry is the last feature on the list to evolve.

Click to enlarge; copyright Gert van Dijk

So glyptodons, ankylosaurs, stegosaurs  and some sauropods all fit the 'big slow armoured' description to various degrees. And so do Furahan rusps! The image above shows half a rusp from an unfinished painting (for more on rusps, use the blog's search function). From my very first rusp sketch on, rusps were large, had thick hides and used their whips as active weapons. Of course rusps have front as well as hind whips, so the word 'tail' is not applicable at all, but the point is clear; rusp whips are analogous to the 'weaponised' tails of Earth. Those early rusp sketches predated the paper as well the posts in this blog about rusps by many years. I do not remember exactly how much of the rusp body plan came about consciously. I think that I started with a long body shape. Add to that some wondering why many Earth animals are so vulnerable at their rear and sometimes along their middle as well. As the earliest sketches show eyes on middle rusps segments as well, rusps must have started with a weak encephalisation tendency. From there on the double encephalisation seems natural. Note that the posterior whip is well controlled by its own ring brain, with excellent visual information available to direct the strikes. But part of all this may have come about through largely unconscious associations while sketching. Once a design is on paper, it is often hard to say where it came from. Regardless, it is nice that the meme 'rusps have whips' can now be attributed to a firmly established biological principle.

Much as I like the paper, there is a minor matter that might have made it even nicer. Rather than 'tail weaponisation', the authors could easily have used a word that is both relevant and fun: a tail weapon is a 'thagomiser'.

Click to enlarge; copyright

The first use of 'thagomizer' is shown above (this blog uses British spelling, so I assumed the word would become 'thagomiser' in the UK; the rules aren't always clear...).
   It was published as one of Gary Larson's Far Side cartoons in May, 1982. Actually, this colour image stems from a later luxury edition of all Far Side cartoons. Poor Thag Simmons. For 'Far Side' fans, a caveman called 'Thag' occurs at least once more, and one cartoon, taking place in modern times, featured a 'Mr Thagerson'.
  At first glance the word thagomiser seems to indicate 'to turn an object, animal or person into thag', but the real meaning is obviously a 'structure to kill animals or persons, in particular Thag Simmons'. The word has later been picked up in the scientific community to describe the tail weapons of stegosaurs, and apparently of stegosaurs only. I propose to widen its use to all tail weapons.
   As an author of scientific papers myself I realise that the use of humour in scientific papers can be tricky as it is often frowned upon, and you never wish to harm your chances of getting a paper accepted. (I once inserted the phrase 'This resistance is futile' in one of my own scientific papers as an irreverent reference to Star Trek, but I do not think anyone ever noticed).

If we use 'thagomiser' as a word for 'tail weapon', the paper could have been called "The evolution of thagomizers in amniotes", which would be clear, succinct and elegant, but admittedly probably too flippant for a serious paper. Once 'thagomiser' is an accepted word, can we resist to stop there? The tendency to evolve a thagomiser then might become 'thagomiserificability', and the transition process from 'nonthagomiseriness' (not having a thagomiser) to 'orthothagomiserity' (having a proper thagomiser) is 'thagomogrification'. Obviously.