Our lives with pterosaurs, part 1

Not many mornings ago the lovely Georgia Maclean-Henry and I were discussing the the topic of 21^st century pterosaurs. Not, you understand, as a discussion of whether the reports of late-surviving, cryptid pterosaurs are genuine (they almost certainly aren’t, for reasons discussed in Darren Naish’s assassination of this idea), but a hypothetical premise that pterosaurs were commonplace components of our modern fauna, and what they would be like to live with. Though obviously speculative and completely juvenile, I thought this may be fun to blog on and discuss with others, so feel free to chime in at the end of the post with your own ideas. Who knows, we may even learn something in the process. (Image, above, shows what we're all now thinking).

Before we get going, though, some ground rules. Aside from the fact that we’re ignoring pterosaur extinction in this discussion, we’re basing everything else on fact as much as possible. For instance, we’re not ignoring the extinctions of specific pterosaur groups: if they went extinct before the terminal Cretaceous (when pterosaurs as a whole got the evolutionary chop), then they can’t exist in the modern day. Pterosaurs are also the only animals we’re hauling into the Modern: the biosphere is otherwise exactly as it is now, so there are no tyrannosaurs or anything running around as well. Also, the goal here is to consider pterosaurs as real animals, not hyper-aggressive movie monsters, so we don’t need to pay any attention to their Modern interactions with people in virtually all Silver Screen outings (which invariably boil down to said people being attacked and/or eaten) and start with a clean slate of ideas. Got that? On we go, then.

UPDATE: 24/04/12
Just one last rule, following on Mike Taylor's comment, below. I'm also focusing on pterosaurs as we know them in the fossil record, not as we may twist them through selective breeding or other genetic tampering. I guess this is an exercise in simply crashing pterosaurs into the Recent, considering what basic pterosaur palaeobiology would lend itself to in our modern world.

Roll call

The first part of this exercise, of course, is to determine what pterosaurs we would have running around today. Which lineages were present at the end of the Cretaceous that could, potentially, have survived until Recent times? Because pterosaur fossils are found within spitting distance, geologically speaking, of Tertiary rocks we assume that the last of their kind died out in the same mass extinction event that ruined the weekends for 75 per cent of life 65 Ma (Buffetaut et al. 1996), but the majority of pterosaur types were not witness to this event. Pterosaur faunas of the uppermost Cretaceous are almost entirely dominated by azhdarchids, the often gigantic, toothless and long-necked forms made famous by the likes of Quetzalcoatlus and Hatzegopteryx (see sketch, above, for a general guide to their appearance). These famous genera, incidentally, are some of the last pterosaurs we find in the fossil record, so giant pterosaurs are very much in for our consideration here. An incomplete nyctosaur humerus (if you’re not familiar with nyctosaurs, think Pteranodon, but weirder) from Mexico is the only record of non-azhdarchid pterosaurs in Maastrichtian strata (that is, name of the time interval representing the last 5 million years of the Cretaceous, 70-65 Ma), compared to literally dozens of azhdarchid occurrences (Price 1953). The pterosaur fossil record is noted for its incompleteness and preservational biases (Butler et al. 2009), but their reduced diversity at the end of the Cretaceous may not be an artifact of the fossil record as the number of pterosaur-bearing rock units at this time is relatively high, but diversity remains low. In short, then, while we may be able to identify dozens of different pterosaur groups across their evolutionary history, it seems that only the azhdarchids and nyctosaurs would have any hope of meeting us in the modern. (Image, below, shows a phylogenetic tree of pterosaurs using the major clades of Lü et al. [2010] mapped across time. The squiggly line shows the number of pterosaur-bearing rock units throughout the Mesozoic [borrowed from Butler et al. 2009] From my book).

With 65 million years separating us from the last pterosaurs, it is not unreasonable to assume that they may have developed into rather different forms by the time modern man appeared. Or would they? Evolutionary stasis spanning 9 – 10 Ma has recently been proposed for several non-pterodactyloid pterosaur clades (Lü et al. 2012) and, although admittedly suggested by rather fragmentary remains, several pterodactyloid lineages also do not appear to change dramatically over longer time frames. This may be true for azhdarchids as much as anything else: a vertebra representing the oldest known azhdarchid is known from Berriasian rocks of Romania (140 Ma) (Dyke et al. 2010) and looks, so far as I can see, no different from the vertebrae of Maastrichtian forms. Note that azhdarchid necks are very derived compared to those of other pterosaurs, so this comparison of their cervical anatomy suggests that the group was already fairly ‘evolved’ very early on in the Cretaceous. Maybe, then, modern pterosaurs would not be so dissimilar from the forms we know in the fossil record.

Bird brains

What sort of behaviour would we expect of our modern pterosaurs? To best answer this we may want to assess some likely basic aspects of pterosaur physiology and neurology, as this may provide  an insight into how active and intelligent they may have been. There’s scant discussion of pterosaur physiology in pterosaur literature, but their flight adaptations, erect carriage (in at least pterodactyloids, and probably some non-pterodactyloids too), insulating fuzz and relatively large brains all seem to correlate with modern animals that have elevated metabolisms. Pterosaur brains are known from specimens spanning much of their phylogenetic range, and they all seem fairly bird-like, but especially so in later forms (e.g. Witmer et al. 2003; image and caption, below, from this study). There are some differences, such as the pterosaur flocculus (the region of the brain primarily dedicated to motor coordination) being relatively enormous, (perhaps because the muscle-laden wing membranes of pterosaurs were being directly controlled and shaped during flight, requiring some extra computing power [Unwin 2005]), and bird brains are, on the whole, a little larger, but they are otherwise fairly similar. 

It may not be unreasonable, then, to predict that all pterosaurs – including our hypothetical modern ones – would be active, fairly intelligent beasties that, with warm bodies and big brains to fuel, may spend much of their time foraging. This leads us to a further analogy with birds: the requirement for lots of food does not sit well with flight, as a full belly is more mass to shift about. Hence, pterosaurs – like birds – may have dumped their waste as often as possible, presumably in the same form of acidic paste that common to all archosaurs. Such waste can be very damaging to architecture and car paint, so the existence of giant pterosaurs dropping vast quantities of crap on our cool stuff is not an appealing one. Plus, we’ve all been hit by stray bird guano on occasion, which is unpleasant enough, but imagine the same experience when the offending animal is several hundred times the size…

My pet pterosaur, and pterosteaks

As with most things in life, it probably wouldn't be long before the economic potential of Modern pterosaurs was tested. Could we farm them for meat and eggs, or breed them as household pets? Pterosaurs would probably be lousy sources of food for several reasons. The amount of meat offered from pterosaur carcasses is tiny compared to their overall size, providing minimal returns to pterosaur farmers for the space required to rear them. Pterosaurs have tiny, tiny bodies, with their edible soft tissues tightly concentrated around them. Even the biggest azhdarchids probably only had bodies 70 cm long (Witton and Habib 2010) with around 60 kg of flight muscle (Paul 2002), despite standing tall enough to look into a first floor window. Ornithocheiroids are even more disproportionate, with torsos barely longer than their humeri (near-enough the shortest bones in their wings). Some pterosaurs may offer better options, such as the relatively long bodied ctenochasmatoids, but they were long gone before the KT boundary, and therefore out of the game here. 

Keeping ourselves stocked with pterosaurs may require a lot of careful planning as their development times appear more extended than we're accustomed to with modern livestock. Because pterosaurs lay parchment-shelled eggs like most modern reptiles, it’s assumed that they required similarly long incubation periods of two or three months (Unwin and Deeming 2008). Once hatched, it seems that neonate pterosaurs did not rocket to full adult size like modern birds (a trait we’ve artificially enhanced in poultry to have large, fully-grown chickens within weeks of hatching), instead slowing their growth rates once they reach half size (Chinsamy et al. 2008). It's predicted that, for some pterosaurs, this threshold may take several years to reach (Bennett 1995; Chinsamy et al. 2008) As such, we could be looking at several years between pterosaur generations, which is a little on the slow side for big business. We don’t know much about pterosaur clutch sizes or reproductive rates, so it’s not clear how many animals you’d need to sustain a harvestable, breeding population but, regardless, it seems that you’d need a pretty substantial operation to get any profit out of space-demanding animals with awkward reproductive mechanisms.

So, pterosaurs would probably make for lousy food sources, but what about pets? It would certainly be cool to keep your own little azhdarchid that you could take out for a flap, train to fetch the morning paper and perform tricks, but the ‘little’ part may be a problem. Pterosaurs are said to demonstrate Cope’s Rule, the controversial idea that the average body size of individuals within a given lineage will increase over time (Hone and Benton 2007; see graph from this study, above, showing the increase in average pterosaur wingspans over time). Whether you agree with the notion of Cope’s Rule or not, it’s hard to ignore the steady increase in average pterosaur body size throughout the Mesozoic, leading to the smallest known Maastrichtian taxon (the oddly-proportioned Montanazhdarcho) being 2.5 m across the wings. A 2.5 m span may seem small compared to its 10 m span contemporaries but, for a homeowner, it would still be far too large to have in the house. Standing upright, said diminutive azhdarchid would have a shoulder height of over a metre and, with its long neck, be nearly as tall – if not taller - as you. That’s hardly a little animal, and probably one that would scare the bejesus out any other pets you have, and may even see them as potential lunch. Perhaps best to leave the pterosaur wrangling to zoos, then.

The biggie: could I ride a pterosaur to work?

Almost certainly the most important consideration in this concept: were pterosaurs strong enough fliers that we could saddle them up fly them places? Well, possibly.

Pterosaur.Net regulars are no doubt aware that some pterosaur workers now think that pterosaurs launched quadrupedally, using their powerful flight muscles to propel themselves into the air (Habib 2008). Part of the rationale for this idea is the strength of the forelimbs compared to the hindlimbs, as the launching limbs tends to be proportionally large in any flying vertebrate you care to look at over a certain mechanical threshold. As with most animal skeletons, it seems that the pterosaur forelimbs came equipped with large mechanical safety factors to accommodate for any atypically heavy loads that may be placed on the limbs. The humeral safety factors against bending in the largest azhdarchids – which we would possess in the Modern in our hypothetical scenario here, remember – are around 2.5 – 1.8, depending on how heavy you consider the animal to be between 180 - 250 kg (Witton and Habib 2010). Thus, the pterosaur skeleton could take weight of a person without crumpling, but could it take off? It seems so: Marden (1994) calculated that a giant azhdarchid would find launch no more strenuous than a 1 kg vulture, suggesting that one could, theoretically, take on the extra burden of a person on its back. Perhaps only relatively small folks would be suitable pterosaur jockeys to reduce the strain as much as possible but, hey, that’s still something, right?

This is not the end of the story, however. While the azhdarchid may be able to sustain flight with a jockey when flapping vigorously, it would not be able to endure this indefinitely. Mike Habib predicted for our 2010 study that a giant would have a few minutes of burst flight, tops, before it had to rest in a gliding phase. To avoid merely landing at the end of this, an alternative source of lift would be needed, and this is where a potential fly in our ointment appears. Long distance travel for azhdarchids was probably achieved by soaring (Witton and Habib 2010), which would be reliant – as it is with modern birds and bats – on climbing to high altitudes (many thousands of metres in some cases) on uplifts of air before gliding on. This would be a significant problem for our jockeys. Mammals are far less tolerant of hypoxia than birds (and, perhaps, by extension, pterosaurs) and, at altitudes that even little birds like sparrows are alert and lively, mammals are comatose (Faraci 1991). Hence, to fly with azhdarchids we may have needed to curb their flight styles a bit, keeping them at lower altitudes and, presumably, making more frequent use of areas of uplift. Alternatively, we supply them with oxygen tanks and warm clothing to keep them alive, but this all adds weight and reduces our azhdarchid's flight ability. Hmm... perhaps this is more complex than we thought.

Gosh, look at the time. There’s a lot more we could mention about riding pterosaurs, but I think we’ll stop there for now. This has already gone on too long and I’ve not even covered the most exciting bit: living alongside wild pterosaurs. Would we be potential pterosaur prey? Could they be pests of annoyances to us? All things to be discussed soon...

References

Buffetaut, E., Clarke, J. B. and Le Lœuff, J. 1996. A terminal Cretaceous pterosaur from the Corbiéres (southern France) and the problem of pterosaur extinction. Bulletin de la Societe Geologique de France, 167, 753-759.

Butler, R. J., Barrett, P. M., Nowbath, S. & Upchurch, P. 2009. Estimating the effects of the rock record on pterosaur diversity patterns: implications for hypotheses of bird/pterosaur competitive replacement. Paleobiology, 35, 432-446.

Bennett, S. C. 1995. A statistical study of Rhamphorhynchus from the Solnhofen Limestone of Germany: year-classes of a single large species. Journal of Paleontology, 69, 569-580.

Chinsamy, A., Codorniu, L. and Chiappe, L. 2008. Developmental growth patterns of the filter-feeder pterosaur, Pterodaustro guiñazui. Biology Letters, 23, 282-285.

Dyke, G., J., Benton, M. J., Posmosanu, E. and Naish, D. 2010. Early Cretaceous (Berriasian) birds and pterosaurs from the Cornet Bauxite Mine, Romania. Palaeontology, 54, 79-95.

Faraci, F. M. 1991. Adaptations to hypoxia in birds: how to fly high. Annual Review of Physiology, 53, 59-70.

Habib, M.B. 2008. Comparative evidence for quadrupedal launch in pterosaurs. Zitteliana, B28, 161-168.

Hone, D. W. E. and Benton, M. J. 2007. Cope’s Rule in the Pterosauria, and differing perceptions of Cope’s Rule at different taxonomic levels. Journal of Evolutionary Biology, 20, 1164–1170.

Lü, J., Unwin, D. M., Jin, X., Liu, Y. and Ji, Q. 2010. Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull. Proceedings of the Royal Society B, 277, 383-389. 
Lü, J., Unwin, D. M., Zhou, B, Chunling, G, and Shen, C. 2012. A new rhamphorhynchid (Pterosauria: Rhamphorhynchidae) from the Middle/Upper Jurassic of Qinglong, Hebei Provine, China. Zootaxa, 3158, 1-19.

Marden, J. H. 1994. From damselflies to pterosaurs: how burst and sustainable flight performance scale with size. American Journal of Physiology, 266, 1077-1084.

Paul, G. S. 2002. Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds. John Hopkins University Press, Baltimore, 472 pp.

Price, L. I. 1953. A presença de Pterosáuria no Cretáceo superior do Estada da Paraiba. Divisão de Geologia e Mineralogia Notas Preliminares e Estudos, 71, 1-10.

Unwin, D. M. 2005. The Pterosaurs from Deep Time. Pi Press, New York, 347 pp.

Unwin, D. M. and Deeming, D. C. 2008. Pterosaur eggshell structure and its implications for pterosaur reproductive biology. Zitteliana, B28, 199-207.

Witmer, L. M., Chatterjee, S., Franzosa, J. and Rowe, T. 2003. Neuroanatomy of flying reptiles and implications for flight, posture and behaviour. Nature, 425, 950-953.

Witton, M. P. and Habib, M. B. 2010. On the size and flight diversity of giant pterosaurs, the use of birds as pterosaur analogues and comments on pterosaur flightlessness. PLoS ONE. 5, e13982.