Research into pterosaurs—ancient flying reptiles whose fossils lie intermingled with dinosaurs—now approaches an upswing of what some predict will be a revolution of understanding.1 Better details of pterosaur anatomies are painting a different picture than the long-held perceptions of these creatures as clumsy fliers. In a recent feature in the UK's Observer, writer Mark Witton explained that many view pterosaurs as "gargoyles with lanky limbs." This bad reputation may have been

Juvenile, subadult and a big, old adult Rhamphorhynchus muensteri forage in a Jurassic lagoon. We know more about the position of this pterosaur in Mesozoic food webs than any other thanks to its excellent fossil record. The floating posture here is based on data from Hone and Henderson (2014). The Late Jurassic, Solnhofen Formation pterosaur Rhamphorhynchus muensteri is an exceptional flying reptile. We tend to overlook it a bit now - it's been known for almost two centuries, which is long enough to temper enthusiasm for any fossil species - but it's a remarkable animal for a number of reasons. Far from a typical example of the rhamphorhynchid lineage, it's the rhamphorhynchiest of all pterosaurs with a jaw full of large, conical teeth, elongate extensions to its jaw tips, exceptionally long and slender wings, delicate hindlimbs and walking digits, and a long, stiff tail famously adorned with a diamond or triangular shaped vane*. It also arguably has the best fossil record of any pterosaur. It's known from over 100 specimens, many of them being complete, articulated skeletons with at least some three-dimensionality, as well as providing excellent soft-tissues remains. Excepting embryos, we have complete growth series from tiny juveniles to chunky adults with 1.8 m wingspans, and its preservation is such that fine details of bones can be gleaned through careful mechanical or acid preparation. Its osteology is subsequently better known that any other pterosaur. The Cretaceous pterodactyloid Pteranodon might be known from more fossils (>1400), but these flattened, disarticulated remains are nowhere close to the fossil quality of Rhamphorhynchus. *We see these restored for numerous long tailed pterosaurs, but only Rhamphorhynchus is known, for fact, to have this 'classic' tail vane morphology. On top of all this, Rhamphorhynchus also provides greater direct insight into its daily behaviour than any other pterosaur. Multiple examples of gut content, coprolites, predator-prey associations and its inclusion in a ball of vomited spittle provide us with insight into what this pterosaur ate, where it was feeding, and what was eating it. In this, a second and concluding foray into the pterosaur palaeoecological record (click here for the first), let's take a look into how this animal slotted into Jurassic ecosystems. Snap my fish up What did Rhamphorhynchus eat? At least three specimens suggest that it foraged for fish of various sizes. The most famous example of such a fossil is sometimes referred to as the 'greedy guts Rhamphorhynchus', an animal which swallowed a fish almost as long as its own torso. This specimen represents a smallish individual, first described by Wellnhofer (1975), with a partly articulated, partly digested fish fossil preserved inside the posterior 60% of the rib cage (below). The orientation of the fish tail fin suggests the fish was swallowed head first, and we have to assume that some major distension of the throat occurred when doing so. That pterosaurs were capable of doing this isn't that surprising considering that extant pterosaur relatives - crocodylians and birds - have mobile throats which aid swallowing of big food items. The relative proportions of these gut contents suggests this animal must've bolted its food like these modern archosaurs, a behaviour rare among mammals but known to occur in at least one large, yellow, bipedal primate. This specimen also presents several elongate elements which defy easy explanation: they're suggested as other food items or even bits of preserved gut tissue by various authors (dark grey in the line drawing below). The fossil, interpretive drawing, and restored reality of Jurassic table manners. Disgraceful. Line drawing after Wellnhofer (1975). Fish dietary remains have been reported from two other Rhamphorhynchus specimens. A complete, relatively small fish has been found in the throat region of one specimen, also swallowed head-first, and various scales and bones occur within the rib cage of another (Frey and Tischlinger 2012; Hone et al. 2013). These three specimens do not represent the limit of Rhamphorhynchus gut content, but they are the limit of identifiable examples: some specimens are known with indeterminate bones or massive, disorganised crystal growths in their stomach regions which represent poorly preserved gut matter, but they aren't of much use to us here. The one, the only, pterosaur coprolite Of the many questions that keep pterosaur experts up at night - what are they related to? how did they work as functional organisms? how are they related to each other? - none has been greater than what their poop was like. Expelled waste is a common form of fossils in some localities, but remained entirely elusive for flying reptiles until last year when Dave Hone and colleagues (2015) identified the first pterosaur coprolite dropping out of a complete Rhamphorhynchus specimen. Finally, pterosaur workers can sleep easy. Rhamphorhynchus muensteri specimen with coprolite (cp). From Hone et al. (2015). The compact shape of these remains and their proximity to the pelvic region suggest that they were expelled from the pterosaur shortly after it settled on the seabed of the Solnhofen lagoon. Some of the coprolite content is pretty indistinct, but one portion preserves thousands of tiny spines, likely indigestible bits of a recent meal. Exactly what they are is difficult to say - you can get a good look here if you'd like to figure them out - but a number of alternatives were considered by Hone et al. (2015). The tentative, albeit still problematic, suggestion is that they represent cephalopod hooklets. This situation is somewhat frustrating, as this coprolite shows that Rhamphorhynchus was not just a fish eater, but doesn't really tell us much else. Still, now that we know what we're looking for, perhaps more pterosaur coprolites might start coming to light. Fossilised food chains Several Rhamphorhynchus specimens reveal it was prey to other Jurassic animals. Among the least commonly discussed is a small pellet produced by something like a fish or crocodylomorph which contains several Rhamphorhynchus wing bones (Schweigert et al. 2001). This is a rare example of Rhamphorhynchus from the Nusplingen Limestone, a unit lithologically similar to Solnhofen but slightly older. As is so often the case with such fossils, the identity of the pellet maker remains elusive. My own suspicions are of a piscine origin, as modern crocodylians don't tend to spit out bones (they digest them, only regurgitating hair, feathers and other keratinous tissues that are difficult to break down). That may not have been true for fossil crocs, of course. Rhamphorhynchus vs. Aspidorhynchus. I guess we should call this one a draw? From Frey and Tischlinger (2012). Among the most remarkable Rhamphorhynchus fossils are five instances where it is preserved alongside the predatory fish Aspidorhynchus acutirostris, a long-bodied species that is often much larger than its Rhamphorhynhus prey (Frey and Tischlinger 2012; Weber 2013). These fossils are palaeoecologically notable for three reasons. Firstly, large Solnhofen vertebrates are hardly ever associated, and yet we have five instances of this same pterosaur and fish species being found in touching, or near touching, proximity. Secondly, all five are exquisitely preserved - one example includes a 'mummified' pterosaur with wing membranes, and all are completely articulated. Thirdly, the Rhamphorhynchus are invariably positioned around the skull of the fish, as if a specific, repeated behaviour saw these animals preserved together. These particulars make chance association of these animals unlikely and imply Aspidorhynchus sought out Rhamphorhynchus as food, probably hunting live specimens rather than scavenging floating corpses (as indicated by the excellent preservation of the individual pterosaurs). Frey and Tischlinger (2012) provide a plausible scenario for the deaths of these animals: they reason the Aspidorhynchus tackled prey that became entangled in their jaws before accidentally entering the toxic bottom waters of the Solnhofen waterways. In these anoxic depths the tangled pair would die pretty quickly (if the pterosaur wasn't already dead, of course), leaving us with perfectly preserved bungled predatory acts. The icing on the cake of these specimens is that one of these Rhamphorhynchus specimens has already been mentioned here - that individual with a fish in its throat (Frey and Tischlinger 2012). In this specimen at least, we can assume the pterosaur ate a fish shortly before being attacked itself: a rare instance of a fossil food chain. The bigger picture of Rhamphorhynchus palaeoecology Collectively, we have 10 specimens of Rhamphorhynchus telling us something about its position in Mesozoic food webs. That's not bad going for a pterosaur, a famously rare type of fossil, and is actually a pretty good palaeoecological record for any fossil vertebrate. 10 specimens is not enough to tell us everything about the lifestyle of a fossil animal, but does allow us to paint a general picture. They show us that Rhamphorhynchus was adapted to foraging on pelagic prey - often small, probably live fish - and that it must have spent a good amount of time in or around water, as it was clearly an attractant to aquatic predators. These specimens gel neatly with general models of Rhamphorhynchus lifestyle interpreted from their functional anatomy. It's generally thought that Rhamphorhynchus was adapted for life along shorelines - basically a Mesozoic gull. It's not uncommon to be suspicious of such claims nowadays, it being realised that the 'Mesozoic seabird equivalent' almost became a trope, or at least a tremendous over-generalisation, of pre-21st century pterosaur science. But in this case, a gull-like lifestyle is a cogent hypothesis based on studies of wing shape, flight style, tooth and jaw apparatus, and limb function (e.g. Wellnhofer 1975; Hazlehurst and Rayner 1992; Witton 2008, 2015; Ösi 2011). Perhaps research on Rhamphorhynchus was involved in creating the stereotype of pterosaurs as Mesozoic seabirds, but we should not regard it as a victims of this stereotyping itself. But we should ask ourselves why Rhamphorhynchus palaeoecology is comparably well-represented in the fossil record. It might be something as simple as preservational conditions, but there are plenty of pterosaur Lagerstätten, some of them containing close relatives of Rhamphorhynchus in reasonable abundance, which provide no information about dietary preferences or interactions with predatory species. Is there something intrinsic to Rhamphorhynchus which makes it special? I find several reasons to wonder if Rhamphorhynchus was an atypically aquatic species, not only flying and feeding above water but actually routinely entering it. Firstly, Rhamphorhynchus has a hatchet-shaped deltopectoral crest (the process on the humerus which anchors the flight muscles), a small (or at least narrow) torso and short legs: these are features which Habib and Cunningham (2010) link to routine and efficient aquatic takeoff. Secondly, tests of pterosaur swimming suggest that Rhamphorhynchus had a pretty stable floating posture - not something that can be said for all pterosaurs (Hone and Henderson 2014). Thirdly, it also has comparably large but delicately constructed feet, which might suit paddling, as well as a slender set of forelimb bones (Witton 2015) which recall the streamlined arm bones of swimming and diving birds (Habib and Ruff 2008). Prolonged bouts of swimming might also account for its general abundance, excellent preservation and all-round good fossil record: being immersed in water puts it a step closer to being preserved than other, less aquatic species. Who knows: perhaps it even dived into Solnhofen's toxic bottom waters on occasion, explaining why so many specimens are excellently preserved? Hmmm.... perhaps this warrants further investigation. Coming soon: could dinosaurs - gasp - lie down on their sides? My take on the greatest of palaeoart debates. Rhamphorhynchus is supported by fish; this blog is supported by Patreon The paintings and words featured here are sponsored by the finest human beings on the planet, those folks who support me at Patreon. Backing my blog for as little as $1 a month helps me churn out researched and detailed articles and paintings to accompany them, and in return you get access to bonus blog content: additional commentary, in-progress views and high-resolution artwork, and even free prints. Accompanying this post, we're going to look at the bigger picture of pterosaur palaeoecology: azhdarchids and Rhamphorhynchus are just two lineages with palaeoecological records - what about the rest of Pterosauria? Sign up to Patreon to join that discussion! References Frey, E., & Tischlinger, H. (2012). The Late Jurassic pterosaur Rhamphorhynchus, a frequent victim of the ganoid fish Aspidorhynchus?. PloS one, 7(3), e31945. Habib, M. & Cunningham, J. 2010. Capacity for Water Launch in Anhanguera and Quetzalcoatlus. Acta Geoscientica Sinica. 31, 24-25 Habib, M. B., & Ruff, C. B. (2008). The effects of locomotion on the structural characteristics of avian limb bones. Zoological Journal of the Linnean Society, 153(3), 601-624. Hazlehurst, G. A., & Rayner, J. M. (1992). Flight characteristics of Triassic and Jurassic Pterosauria: an appraisal based on wing shape. Paleobiology, 18(04), 447-463. Hone, D. W., Habib, M. B., & Lamanna, M. C. (2013). An annotated and illustrated catalogue of Solnhofen (Upper Jurassic, Germany) pterosaur specimens at Carnegie Museum of Natural History. Annals of Carnegie Museum, 82(2), 165-191. 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. Hone, D., Henderson, D. M., Therrien, F., & Habib, M. B. (2015). A specimen of Rhamphorhynchus with soft tissue preservation, stomach contents and a putative coprolite. PeerJ, 3, e1191. Ősi, A. (2011). Feeding‐related characters in basal pterosaurs: implications for jaw mechanism, dental function and diet. Lethaia, 44(2), 136-152. Schweigert, G., Dietl, G. & Wild, R. (2001). Miscellanea aus dem Nusplinger Plattenkalk (Ober-Kimmeridgium, Schwäbische Alb) 3. Ein Speiballen mit Flugsaurierresten. Jahresberichte und Mitteilungen des Oberrheinischen Geologischen Vereines, 83, 357-364 Weber, F. (2013). Paléoécologie des ptérosaures 3. Les reptiles volants de Solnhofen, Allemagne. Fossiles. 14. 50-59. Wellnhofer, P. 1975. Die Rhamphorhynchoidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands. Palaeontoographica A, 149, 1-30. Witton, M. P. (2008). A new approach to determining pterosaur body mass and its implications for pterosaur flight. Zitteliana, 28, 143-159. Witton, M. P. (2015). Were early pterosaurs inept terrestrial locomotors? PeerJ, 3, e1018.

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