The pterosaurs got going something like 230 million years ago. They died out with the dinosaurs 65 million years ago. Over their history they came in all sizes, from Rhamphorhynchus who was the size of a sparrow to Quetzalcoatlus with a wingspan of variously estimated as being 30-40 feet. Spread out it was a similar size to a t-rex, and on the ground its estimated height was close to a giraffe's. The largest bird ever, Argentavis was much, much smaller than that.
However the birds arose about 150 million years ago. Feathers were a big advantage in flight, and over time the birds took over a lot of what the pterosaurs were doing. But the pterosaurs did not go away. Instead they wound up being in niches for very big flying animals. This is competition through specialization, which I talked about some time ago when I discussed the Neanderthals.
This coexistence provides evidence that birds were generally better fliers, but there was a niche for very big fliers that the pterosaurs were better at. The question I'm curious about is why the pterosaurs were better at being big fliers than birds were.
I have a theory. But before I can explain it I need to provide some background.
Wings have evolved in vertebrates three times in pterosaurs, birds, and bats. All three started with the basic vertebrate limb structure and found different ways of constructing a wing out of it. In both bats and birds the arm bones form part of the wing. Now an important fact about vertebrate bones is that different bones grow at different rates as you grow. In particular arm bones start off shorter and catch up later. The result is that in birds and bats, babies have the wrong proportions for their wings to be useful. Therefore baby birds and bats can't learn to fly until they have achieved a significant fraction of their full size.
Pterosaurs were different. Their wings were entirely constructed from wrist and hand bones. (Fully half the wing was supported by an elongated 4th finger.) Hand bones stay in proportion your whole life. Comparisons of fossils of pterosaurs at different ages in the same species verifies that their wings always had good proportions for flight. Furthermore we have fossils from baby pterosaurs that died miles out at sea, which is direct evidence that they flew young.
What does this have to do with the eventual size of the animals? Well birds cannot learn to fly until they are near full growth. Which means that they need intensive care from their parents until they reach that growth. This care is a significant fact of life for bird species, and is why most types of birds have both parents providing care. Unlike most mammals where the mother is generally capable of taking care of young on her own. The larger the bird is, the harder this care is to provide.
By contrast pterosaurs were probably able to take care of themselves at a much younger age, and smaller size. Which means that they were free to grow for a lot longer, to a lot larger size, without unduly taxing their parents. (In truth we don't have any data indicating how much or little parental care baby pterosaurs got. But I suspect it was less than birds get.) And, I believe, that is why they were able to get so much larger than birds.
Random trivia I came across in preparing this post. The reason bats can't fly during the day is that their wings are vulnerable to sunburn. There is evidence that pterosaurs had a protective layer so they didn't have this issue. Also birds have stiffer wings than bats do, which provides better lift and less maneuverability. Pterosaurs had more joints in their wings than birds do, but didn't have finger bones inside of the structure of their wings like bats, which suggests to me that their wings would have been somewhere between.
And my whole train of thought was started by watching National Geographic - Sky Monsters. If you're interested in pterosaurs, it is a worthwhile video.
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