Birds with clubs and other smashers: clavigerism

I intended to write just one blog entry every two weeks this year, and am already breaking that rule. I could not resist, after reading a paper on an extinct Terran bird. The similarities to some Furahan creatures were simply too interesting to leave it alone, so there you are.

The bird, a flightless ibis, was described in a paper entitled 'The bizarre wing of the Jamaican flightless ibis Xenicibis xympithecus : a unique vertebrate adaptation' (Longrich NT, Olson SL; Proc R Soc B 2011, online 5 January 2011).

Click to enlarge; Longrich NT, Olson SL; Proc R Soc B 2011, online 5 January 2011

And here you see a reconstruction of the bird (top) and a comparison of its wing bones (bottom) with that of a still living ibis species. The authors make a case that the bird was flightless, and go on to say that the wing bones are odd for flightless birds. That holds for the hand in particular, with its thick and curved bone. The authors think it functioned as a club, and provide various anatomical reasons why they think so. After that, they discuss the club some more:

"We therefore propose that the wing of Xenicibis functioned as a club or flail. Several features of the limb would have facilitated this function. Kinetic energy is the product of mass and velocity squared; accordingly, weapons such as clubs and flails have a long handle to increase the angular velocity of the club, and are heavily weighted to increase the mass accelerated by the swing, and the centre of mass is near the end of the club, where the angular velocity is highest. Precisely this design is seen in the hand of Xenicibis, where the end of the wing is massive, and the proximal metacarpus and long forelimb could act as a handle. "

The authors of the paper also compare this odd ibis design with the front legs of mantis shrimps (Stomatopods), that function in a similar manner. The video above shows a slowed down version of a strike, and incidentally allows the shape of the club to be appreciated as well. It is taken from the lab of Sheila Patek, whose work on the biomechanics of their legs featured in this blog previously.

Click to enlarge; copyright Gert van Dijk

A long time ago mantis shrimps were the inspiration for Furahan neocarrnivores. You will find more material on them on the Furaha site (go to the land page). One of them is shown here (one of its commoner names is 'pugile'). Compare its front legs to those of mantis shrimps and to the ibis; I think that the ibis still had a long way to go before its clubs measured up to those of other club bearers. Then again, they were probably meant to hit other ibises, not kill prey animals, and it might not be advantageous in the long run to kill fellow ibises; merely sending them off might be good enough.

Stomatopods, Xenicibis and Furahan neocarnovores may all be said to have developed their clubs from what originally were locomotor limbs. In the Stomatopod and Neocarnivore cases, the limbs in question were used to walk with, and in the Xenicibis case its ancestors used them to fly with. In all cases their freedom from locomotion opened the door for a new function, and interestingly the new function was a weapon in all three cases. I think that Xenicibis therefore qualifies as much as the other two as an example of 'centaurism' (more on centaurism here and here).

I am delighted that we now have an example of 'raptorial centaurism' concerning a fairly large animal on Earth. Both Stomatopods and Furahan Neocarnivores have evolved various kinds of front legs, functioning as different types of weapons: there are 'smashers' and 'spearers' in both cases (Neocarnivores have webs or basket-like thingies as well). It would be great if more diligent paleontological work would uncover a 'spearer' on Earth as well, but I will not hold my breath. Meanwhile, perhaps it is time to coin a phrase to start differentiating between the various kinds of weapons limbs may turn into. 'Smashers' sounds good, but is limited to English, and that won't do. Luckily we still have Latin. A 'claviger' is an existing word for club carrier, with the plural 'clavigeri'. Turning it into a principle would result in 'clavigerism'. I don't expect a follow-up paper on Xenicibis to use the word; then again, the authors used the interesting phrase 'volant species' for birds that actually fly, so perhaps they can appreciate a bit of biogeekery...

Strandbeesten and mantis shrimps

Actually, 'Strandbeesten and stomatopods' might have sounded better, but would be even more incomprehensible, and a blog is supposed to attract readers, not frighten them away. Based on how many readers were attracted by previous posts, I should probably use 'The Future is Wild' and particularly 'Avatar' a lot more often in post titles (and no, Furaha was NOT modelled on Avatar; it is much older). Right; now that I've got that out of the way, back to the strandbeesten.

I discussed Theo Jansen's imaginative mechanical walking machines before on this blog. Literally the word is Dutch for 'beach beasts'. If you do not know about them, read that entry and visit Mr Jansen's site, or just enter 'Theo Jansen' into Google or YouTube. His work came up in this blog because of my interest in animal locomotion. The problem he faced was how you can get a foot to move backwards along a straight line when on the ground, after which it has to be lifted, moved forward and put down again for the next step. For real animals this is not a big problem, as the various segments of a limb are all controlled by a nervous system telling each segment when to do what. As Jansen's devices lack a brain, he needed a purely mechanical system to achieve this sort of motion. In the end he came up with an intricate series of interconnected bars: if you start with a rotary motion of one bar, another bar, ending in a foot, produces a suitable movement. Very clever indeed. Such series of connected bars are called linkages. You can take a good look at his design on this particular site, which shows other linkages as well. When I wrote that post I had never seen a single strandbeest yet, and that has now been rectified. Mr Jansen works not that far from where I live, so it was a matter of time before I could visit one of his demonstrations nearby. This was the case last June, on a very cold and windy day. I will show a few videos I made that day.

This is a tiny strandbeest, of which there were three. If its sail is perpendicular to the wind direction, the little beast may walk with the wind. I tried pushing it forward as well, and found that it is not in fact that easy to move. While the 'beesten' are quite light, their joints were harder to move than I had expected. There is no lubrication, but the main problem seems to be that the entire shape deforms enough to put shearing forces on the joints. One result of this is that the poor beest tends to topple over. But never mind that, they are an amazing sight.

Here is a larger one following one of Mr Jansen's assistants.

And this is the major species present at the occasion. Not only did it have two bodies or trunks, an enormous number of legs, but also two waving membranes at the top that I think were designed to help propel it. These sails were reefed that day however, and the force of the wind on the body was enough to prod the beast onwards. Aren't they wonderful?

In my previous post I wondered how often linkages occurred in biology, but did not look up the matter. I have done a bit of research now, and found that there are quite a few examples. Fish jaws are probably the best-known example (see below). Other structures, such as sheep hocks and human knees are also counted as so-called four-bar linkages. In a four-bar linkage four stiff bars are linked together in a sort of circle by pivots. If you hold one bar still, and move another one, the remaining two must move in a fixed way. What that way is depends on how exactly they are connected. I felt that regarding the human knee as a four-bar linkage is bending the rules a bit, as two of the bars are ligaments rather than stiff bars. If you include connected series of bones as well as ligaments there are lots of linkages in biology; what I was looking for was linkages of bones involved in locomotion, but I have not yet seen any. Presumably a system with more mechanical freedom but with a smart nervous system to control it is simply superior. Still, the other ones are interesting.

Click to enlarge; Wainwright et al, Integr Comp Biol 2005; 45: 256-262

Here is an example of what fishes do with a four-bar linkage. The source is mentioned in the caption, and the colours are my addition. Fish use this kind of mechanism to move their jaws forward and to enlarge the volume in their mouths, sucking in water as well as their prey. There is at least one Furahan animal with a similar arrangement, and those are 'Fishes' too. The jaws of the sawjaw are connected, all four of them, by bars linking them to the neurocranium in a kind of circular linkage.

So where are the mantis shrimps, everyone's favourite Terran alien? When searching for linkage mechanisms I found that there is a four-bar mechanism in their 'raptorial appendages' as well! I like that: somehow you expect animals that not just spear or club their prey but can see depth with just one eye to be special in other respects as well, and mantis shrimps never seem to let us down when it comes to, well, weirdness.

Click to enlarge; Patek et al; Nature 2004; 428: 819

Here is a figure from the journal Nature, no less. The first author, Sheila Patek, has a lab where she studies all kinds of biomechanically interesting things, most notably mantis shrimps. Have a look, as there are quite a few videos and photographs. Under 'multimedia' you will find an inspired lecture she gave on 'TED', where she explains the striking mechanism of stomatopod raptorial appendages. Very interesting. She is not the only one interested in stomatopods either; here is another enthusiast.

So now you know why strandbeesten and stomatopods end up in the same post: they are connected by linkage (I could not resist that one). I guess both also score very highly when it comes to their ability to evoke a sense of wonder.



PS. This is post #100!