Damselfly Wing Animation

I’m working on the art for a swooping melee-attack damselfly right now, and wanted to get a more realistic wing animation.  This turned out to be fairly challenging, first because insect wings move so quickly.  Even a 12x speed video won’t capture more than a blur.  There’s plenty of dragonfly slow-mo videos shot with professional cameras, but damselfly videos are harder to find.

Why damselflies?  Because they rest their wings folded atop their bodies, looking better for a 2D perspective in our game.  Dragonfly wings would look odd, seen edge-on.  

My search for a reference took me to some fascinating places online.  Japanese researchers back in the 1980′s had studied the flight mechanics of damselfly wings, down to the forces acting on individual sections! 
Check this out:

University of Tokyo, Flight Mechanics of a Damselfly -  AKIRA AZUMA, SOICHI AZUMA, ISAO WATANABE AND TOYOHIKO FURUTA  

 https://jeb.biologists.org/content/jexbio/116/1/79.full.pdf
The black and white gif I included is from photos taken from this paper.

Next, I found at a video that illustrated damselfly darting with fluid mechanics simulations. 
https://youtu.be/erd8FibfPR4

(the other gif is carefully taken from this video and reassembled so it stands still, for use as a reference).  

It’s like the damselfly swims through the air, using two sets of paddles - and still, the movement appears strange and counterintuitive.  

The only reference I couldn’t find was on the color shift of an ebony damselfly’s wings.  I had to fudge it, half from blurry video frames, half trying to not look too busy in a pixel-art animation.

This has been fun, challenging, and a deep dive into a rabbit hole I otherwise would have overlooked!