Showing posts with label Ornithocheiridae. Show all posts
Showing posts with label Ornithocheiridae. Show all posts

Saturday, 7 March 2015

How Ornithocheirus simus and other pterosaurs took to the air... from water?

Aquatically-adapted ornithocheiroid Ornithocheirus simus takes off using aquatic quad launch, as hypothesised by Habib and Cunningham (2010). Prints of this painting - which might be the first illustration of this launch strategy - are available from my shop.

Many pterosaur lineages seem to have close ties with marine environments, as evidenced by biases in their fossil and taphonomic records and indications of frequent interactions with marine fish. It stands to reason that these animals would find themselves in water on occasion, and trackways made by swimming pterosaurs indicate they may have been quite at home in this medium. Recent studies by Dave Hone and Donald Henderson have cast doubt on the swimming ability of pterosaurs because their floating postures seem rather awkward, which they suggest might have impeded breathing while swimming (Hone and Henderson 2014). Their studies found that, rather than sitting atop the water with arcing necks like birds, front-heavy pterosaur bodies collapse the head and necks into the water, bringing the mouth and nostrils close to the water surface. Although initially sceptical of this idea, I must admit to at least agreeing that avian-like postures may be difficult for pterosaurs. Our expectation that they floated in a bird-like fashion - which I've illustrated several times (Witton 2013 and elsewhere) - is actually quite silly given their proportions and differences with bird morphology. Pterosaurs lack the well-muscled hindlimbs which depress the back end of bird bodies into water, as well as the flexible necks and small heads required to attain duck or gull like floating postures. Indeed, birds seem unique in their floating posture, whereas pterosaurs seem to have floated in a manner more typical of other animals. Does the proximity of their nostrils or mouths to the water surface impede their swimming ability? Maybe not, given that virtually all non-avian animals I can think of  - including aquatic species like otters, crocodylians, swimming rodents, etc. - float and swim with their nostrils close to the water. As long as they have enough control over their swimming ability to clear their nostrils for respiration, they were probably fine. I see no clear reason to think pterosaurs were less competent in water than other animals, and maintain the view that some - for functional reasons - probably needed to swim to obtain the pelagic prey they did/likely did consume (Witton 2013).

But what did pterosaurs do when they needed to leave the water? Could they fly from the water surface or did they have to seek land to take off from? Anyone who knows anything about current pterosaur research knows that the leading hypothesis on pterosaur takeoff is quadrupedal launching, where the hindlimbs mostly serve to provide forward momentum, and vertical heft was provided by the forelimbs (Habib 2008) - many species of bats including vampires molossids and some mystacinids use a similar mechanism. It's worth stressing that this idea is not only supported by anatomical characteristics and positive results from biomechanical studies, but also by the fact that the hindlimbs were too weak to launch sensibly-massed pterosaurs into the air. All studies favouring bipedal launch have had to circumvent this somehow, typically by under-estimating pterosaur masses by, probably, as much as 60%. This makes bipedal launch a real no-go, whereas quad-launch has, to my knowledge, has meet all tests and predictions. We typically discuss quad-launch in terrestrial contexts, but can it work on water?

Schematic of water-hopping quad-launch strategy, from Witton 2013. The floating posture in panel 1 might be incorrect, but the general thrust of the image is OK.

It turns out, probably yes. According to work by Habib and Cunningham (2010), a version of quad-launch works just fine in aquatic settings. It seems that many folks have difficulty imagining how this works, but it's really not too dissimilar to terrestrial quad-launching. As with any takeoff mechanism, the the trick to water-launching is a leap providing sufficient height and speed to facilitate wing use (flapping alone doesn't really get you anywhere, and is especially ineffective at larger sizes). In this respect, it is just like terrestrial launch. An added complication of water launch is escaping surface tension, the cohesive force operating at air-water interfaces. This is where differences between terrestrial and aquatic launch become apparent, because it seems most pterosaurs were incapable of overcoming surface tension from a 'standing' (er... stationary floating) start. Their half-submerged posture and the fluid nature of the medium they are pushing against likely prohibited generation of sufficient energy for a standing launch. The solution to this is a series of hops across the water surface (above), each one providing further water clearance and velocity than the last. These were achieved, as on land, by the combined efforts of both the legs and arms. The wings are not fully deployed at this point, although the arm motion is probably showing some similarities to a flight stroke. The early stage of this takeoff might  look a little like swimming with a particularly powerful butterfly-stroke, albeit one where the swimmer is emphasising vertical motion rather than horizontal. Eventually, the pterosaur is leaping across - not through - the water and is clear enough to push off fully from the water surface. For this final push, the wing is opened and flapping can start. It seems that some pterosaurs - principally ornithocheiroids - were very well adapted for these manoeuvres, showing the shoulder reinforcement, upper-arm strength and distal-limb adaptations you'd predict for water-hopping quad launchers (Habib and Cunningham 2010, Witton 2013).

Some especially powerful pterosaurs, however, probably didn't need to worry about water hops. Giant azhdarchids, which experts predict were not only fuelled by muscle, but raw awesomeness sucked out of the universe itself, were probably powerful enough to water launch without hopping (Habib and Cunningham 2010). Some also ducks have sufficient power to do this - check out the launches in this video for cracking, slow-mo examples (the second is best, at 20 seconds in. Hat tip to Mike Habib for the link).


Recently, palaeoblogosphere regular Mike Traynor commissioned me to paint pterosaur water quad-launch - I think for the first time (if anyone knows different, please let me know). The results, a 6 m wingspan Ornithocheirus simus at the apex of the launch cycle, are above and, for fun, shown in progress from my Twitter feed below. If you'd like to own a copy of this image, you can: point your internet mobile at this page.





We're not done with aquatically-adapted animals just yet. Coming soon: the surprising aquatic adaptations of our own Mesozoic relatives.

References

  • Habib, M. B. (2008). Comparative evidence for quadrupedal launch in pterosaurs. Zitteliana, B28, 159-166.
  • Habib, M., & Cunningham, J., (2010). Capacity for water launch in Anhanguera and
  • Quetzalcoatlus. Acta Geosci. Sin. 31, 24–25.
  • Hone, D. W., & Henderson, D. M. (2014). The posture of floating pterosaurs: Ecological implications for inhabiting marine and freshwater habitats. Palaeogeography, Palaeoclimatology, Palaeoecology, 394, 89-98.
  • Witton, M. P. (2013). Pterosaurs: natural history, evolution, anatomy. Princeton University Press.

Friday, 8 February 2013

Ornithocheirus and Anhanguera: 4 m wingspans are rubbish

It's been a bit pterosaur-light around these parts since I opened the blog in November, with dinosaurs dominating most posts. This week, to start setting things right, we're returning to the warm, leathery-winged bosom of pterosaurs, with a painting from 2010 showing two of the most famous ornithocheirid pterosaurs, Anhanguera santanae (on the left) and Ornithocheirus mesembrinus (right). These Brazilian pterosaurs are both from the Lower Cretaceous Santana Formation, a fossil site renowned for its excellent, three-dimensionally preserved vertebrate fossils. Pterosaurs are the most common tetrapods in this unit, and ornithocheirids are a well known component of that fauna. In fact, they're probably the most extensively documented pterosaurs from the Santana, in part because of the thorough and beautifully illustrated descriptions by Peter Wellnhofer, including those for specimens of Ornithocheirus and Anhanguera (Wellnhofer 1987; 1991). 
Cock of the slight awkward walk: Ornithocheirus. mesembrinus out for a  stroll, possibly trying to accentuate it's bottom. Perhaps it works out. 
Ornithocheirids are unusually proportioned pterosaurs, bearing extremely robust and long wings, enormous heads but tiny bodies and legs. Only other ornithocheiroids, particularly members of Pteranodontia and (to a lesser extent) Istiodactylidae can boast similar proportions. These forms are considered closely related by some (e.g. Unwin 2003), suggesting that their unusual bauplan developed only once, and was taken to extremes by members of Ornithocheiridae and Pteranodontia. In all likelihood, this evolutionary emphasis on increasing the size of the wings and head reflects adaptations for long soaring flight over seas and oceans, while retaining long jaws to grab pelagic prey. This group of ocean-soaring pterosaurs also includes Nyctosaurus, which may be one of the most effective soaring animals to have ever lived. Nyctosaurus also achieves the accolade of being the cover star of my book, which I'm sure it would be much more excited about. 
Two cowardly Anhanguera santanae, being cowardly.
 The painting here shows a few ornithocheirids striding around, an activity that probably wasn't their favourite pastime. Their short trunk skeletons and hindlimbs make for very disproportionate frames, and their forelimbs are probably at the limit of being useful in terrestrial locomotion, beyond simply preventing them from falling over. The pair of Anhanguera on the left are clearly somewhat wary of the larger Ornithocheirus, but it's worth mentioning that they're hardly small. The wingspan of A. santanae is estimated at 4.15 m, which is fairly middling for a Cretaceous pterosaur, but dwarfs the largest flying animals we have today with their piddling 3 m wingspans. Ornithocheirus mesembrinus, by contrast, is one of the largest ornithocheiroids known with an estimated wingspan of 6 m (this, of course, contradicts what Kenneth Branagh told us in Walking with Dinosaurs, but evidence for Ornithocheirus, or any ornithocheirid for that matter, spanning 10 m has yet to be presented). The only ornithocheirid that may have intimidated O. mesembrinus was Coloborhynchus capito, which may have spanned up to 7.25 m (Martill and Unwin 2012), gigantic proportions comparable to those of the largest Pteranodon. Accordingly, when Ornithocheirus wanted to walk or fly somewhere, Anhanguera moved out of its way. 

And that will have to do today, I'm afraid. I've already gone on too long, and I'm much too busy to say anything about other things that are relevant here: colour choices, ornithocheirid rostral structure, ornithocheirid taxonomy and many other things. Perhaps another time, then. 

References

  • Martill, D. M. and Unwin, D. M. 2012. The world’s largest toothed pterosaur, NHMUK R481, an incomplete rostrum of Coloborhynchus capito (Seeley, 1870) from the Cambridge Greensand of England. Cretaceous Research. 
  • Wellnhofer, P. 1987. New crested pterosaurs from the Lower Cretaceous of Brazil. Mitteilungen der Bayerischen Staatsammlung für Paläontologie und Historische Geologie, 27, 175-186.
  • Wellnhofer, P. 1991. Weitere pterosaurierfunde aus der Santana-Formation (Apt) der Chapada do Araripe, Brasilien (Translated title: Additional pterosaur remains from the Santana Formation (Aptian) of the Chapada do Araripe, Brazil).  Palaeontographica Abt. A, 215, 43-101.