Thursday, 28 May 2015

Quadrupedal Launching In Bats And Pterosaurs

I can't recall when I first heard about 'quad-launching' as a serious suggestion for pterosaurs getting airborne, (I was under a rock, palaeontologically-speaking, between '06 and '12) though Mark Witton's excellent 'Pterosaurs - Natural History, Evolution, Anatomy' was the first time I remember anyone going to any effort to depict it pictorially. Indeed, all of his book's pterosaurs are shown mid-launch for their profile images, as if Mark is making a concerted effort to familiarise readers with the concept. Most of the other books on my shelves tend to hedge their bets, offering up a selection of methods, including (but not limited to) dropping from elevated perches, facing into the wind and spreading their wings, and taking a run up whilst flapping.

My biggest problem with quad-launching was that I found it hard to visualise. I've never seen anything get airborne like that. Given that birds are obligate bipeds and their legs are not connected to their wings by a continuous flight surface, they are free to either jump into the air, as with pigeons, or propel the animal along the ground with an energetic run-up, like swans and geese. Many palaeontologists agree that pterosaurs were obligate quadrupeds and that their fore-limbs and hind-limbs were, in life, connected by the wing membrane. Birds are, therefore, a poor analogue for launching pterosaurs, and it is for these, and other anatomical reasons, that palaeontologists believe that pterosaurs' primary launch method probably involved a highly-energetic 'push up'.

A recent post at Pterosaur Heresies again demonstrates its author's frustrations with the problems he sees with the forelimb launch mechanism. The article points out that vampire bats achieve a considerable height from an initial leap before they perform a single flap, and that pterosaurs would be unlikely to achieve such a feat. In a bid to attempt to understand bats taking off from the ground (only a few species can do this) I looked at video footage of a fringed myotis taking off. Adams et al, in their 2012 paper, looked at how bats use their uropatagium to facilitate launch, and made available the following video:

There are four video links in the online paper, showing launches from various angles. In order to get a better idea of what's going on, I rendered the bat as a very-basic stick figure, traced from screenshots of the first online video. The wings' tracings show the stroke, and the head shows the positions of the animal relative to the ground.

Sequence showing a bat (Myotis thysanodes) taking off from the ground, mapped from screen-shots of film footage. This section of the sequence totals around two-and-a-half seconds. (Sequence drawn by author, traced from footage available with Admas, Snode & Shaw 2012.)
In the next image, the seven stages are overlaid in order to get a slightly clearer view - though I think both diagrams are useful when taken in together. The bat accelerates quickly, with its wings in contact with the ground in stage 1-3 (in 1 and 2, they are still flush to the floor). In stage 4 it begins the upstroke, is preparing for its first proper downstroke at 5, and has achieved that downstroke by stage 7. It's already flying and is only a few inches off the ground. My understanding, at least for M. thysanodes, is that when it jumps its inertia carries it a little higher than it would appear when standing with its arms stretched out beneath it, but it's enough to get the first flap in, and by then it's already airborne.

The same bat's take-off sequence, overlaid in order to better show the small area required for a successful launch. Black numbers denote head positions during launch; red numbers denote left wingtip positions. (Sequence drawn by author, traced from footage available with Admas, Snode & Shaw 2012.)
About a year ago I began work on a graphic novel showing the birth, life and death of Nyctosaurus. I may have underestimated how long this would take to put together, so it's still filed under 'ongoing'. But in order to understand quad-launching, I put together a couple of graphics showing an adult Nyctosaurus getting airborne, both of which inspired the bat graphics:

Overlaid launch sequence for a male Nyctosaurus gracilis. (Copyright © 2014 Gareth Monger)
And the looong version:

Launch sequence for Nyctosaurus. Nicked from my deviantART profile, hence the whole lo-res thing. Copyright © 2014 Gareth Monger)
So there you go. Now that I've done the bat thing, I might refine the Nyctosaurus graphics. I might even put together a cel animation at some point. There's nothing overly scientific in all that, however it might prove useful for those of you out there who are into your leather-flappers and pterosaurs.


Adams RA, Snode ER, Shaw JB (2012) Flapping Tail Membrane in Bats Produces Potentially Important Thrust during Horizontal Takeoffs and Very Slow Flight. PLoS ONE 7(2): e32074. doi:10.1371/journal.pone.0032074

Elgin, R.A., Hone, D.W.E., and Frey, E. 2011. The extent of the pterosaur flight membrane. Acta Palaeontologica Polonica 56 (1): 99–111.

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