Editing Pterosaur (section)

===Diet and feeding habits===
[[File:Dimorphodon.png|thumb|Modern interpretations of the diet of ''[[Dimorphodon]]'' have challenged traditional ideas of all pterosaurs being piscivorous]]
Traditionally, almost all pterosaurs were seen as surface-feeding piscivores or fish-eaters, a view that still dominates popular science. Today, many pterosaurs groups are thought to have been terrestrial carnivores, omnivores or insectivores.

Early-on it was recognised that the small [[Anurognathidae]] were nocturnal, aerial insectivores. With highly flexible joints on the wing finger, a broad, triangular wing shape, large eyes and short tail, these pterosaurs were likely analogous to [[nightjars]] or extant insectivorous bats, being capable of high manoeuvrability at relatively low speeds.<ref>{{cite journal | last1 = Bennett | first1 = S. C. | year = 2007 | title = A second specimen of the pterosaur ''Anurognathus ammoni'' | doi = 10.1007/bf02990250 | journal = Paläontologische Zeitschrift | volume = 81 | issue = 4| pages = 376–98| bibcode = 2007PalZ...81..376B | s2cid = 130685990 }}</ref>
[[File:Lusognathus.png|thumb|left|[[Ctenochasmatoid]] pterosaurs such as ''[[Lusognathus]]'' may have had specialised niches in freshwater ecosystems]]
Interpretations of the habits of basal groups have changed profoundly.  ''[[Dimorphodon]]'', envisioned as a [[puffin]] analogue in the past, is indicated by its jaw structure, gait, and poor flight capabilities, as a terrestrial/semiarboreal predator of small mammals, [[squamates]], and large insects.{{sfn|Witton|2013|p=103}} Its robust dentition caused ''[[Campylognathoides]]'' to be seen as a generalist or a terrestrial predator of small vertebrates, but the highly robust humerus and high-aspect wing morphology, suggest it may have been capable of grabbing prey on the wing;{{sfn|Witton|2013|p=121}} a later study indicates it was [[teuthophagous]] based on squid findings within its gut.<ref>Cooper, S. L. A.; Smith, R. E.; Martill, D. M. (2024). "Dietary tendencies of the Early Jurassic pterosaurs Campylognathoides Strand, 1928, and Dorygnathus Wagner, 1860, with additional evidence for teuthophagy in Pterosauria". Journal of Vertebrate Paleontology. e2403577. doi:10.1080/02724634.2024.2403577.</ref> The small insectivorous ''[[Carniadactylus]]'' and the larger ''[[Eudimorphodon]]'' were highly aerial animals and fast, agile flyers with long robust wings. ''Eudimorphodon'' has been found with fish remains in its stomach, but its dentition suggests an opportunistic diet. Slender-winged ''[[Austriadactylus]]'' and ''[[Caviramus]]'' were likely terrestrial/semiarboreal generalists. ''Caviramus'' likely had a strong bite force, indicating an adaptation towards hard food items that might have been chewed in view of the tooth wear.{{sfn|Witton|2013|p=122}}
[[File:Haliskia_Life_Restoration.png|thumb|Many [[pteranodontoid]] pterosaurs such as ''[[Haliskia]]'' likely fed on fish at sea]]
Some [[Rhamphorhynchidae]], such as ''[[Rhamphorhynchus]]'' itself or ''[[Dorygnathus]]'', were fish-eaters with long, slender wings, needle-like dentition and long, thin jaws. ''[[Sericipterus]]'', ''[[Scaphognathus]]'' and ''[[Harpactognathus]]'' had more robust jaws and teeth (which were ziphodont, dagger-shaped, in '' Sericipterus''), and shorter, broader wings. These were either terrestrial/aerial predators of vertebrates<ref name=ACX10>{{cite journal |last1=Andres |first1=B. | last2=Clark |first2=J. M. | last3=Xing | first3=X. |year=2010 |title=A new rhamphorhynchid pterosaur from the Upper Jurassic of Xinjiang, China, and the phylogenetic relationships of basal pterosaurs |journal=Journal of Vertebrate Paleontology |volume=30 |issue=1 |pages=163–87 |doi=10.1080/02724630903409220|bibcode=2010JVPal..30..163A |s2cid=53688256 |url=http://doc.rero.ch/record/31614/files/PAL_E956.pdf }}</ref> or [[corvid]]-like generalists.{{sfn|Witton|2013|p=134}} [[Wukongopteridae]] like ''[[Darwinopterus]]'' were first considered aerial predators. Lacking a robust jaw structure or powerful flying muscles, they are now seen as arboreal or semiterrestrial insectivores. ''Darwinopterus robustidens'', in particular, seems to have been a beetle specialist.<ref name=robustidens>{{cite journal |author1=Lü J. |author2=Xu L. |author3=Chang H. |author4=Zhang X. | year = 2011 | title = A new darwinopterid pterosaur from the Middle Jurassic of western Liaoning, northeastern China and its ecological implications | journal = Acta Geologica Sinica - English Edition | volume = 85 | issue = 3| pages = 507–14 | doi = 10.1111/j.1755-6724.2011.00444.x|bibcode=2011AcGlS..85..507L |s2cid=128545851 }}</ref>

Among pterodactyloids, a greater variation in diet is present. [[Pteranodontia]] contained many piscivorous taxa, such as the [[Ornithocheirae]], [[Boreopteridae]], [[Pteranodontidae]] and Nyctosauridae. [[Niche partitioning]] caused ornithocheirans and the later nyctosaurids to be aerial dip-feeders like today's [[frigatebird]]s (with the exception of the plunge-diving adapted ''[[Alcione elainus]]''), while boreopterids were freshwater diving animals similar to [[cormorants]], and pteranodonts pelagic plunge-divers akin to [[boobies]] and [[gannets]]. An analysis of ''[[Lonchodraco]]'' found clusters of [[Foramen|foramina]] at the tip of its beak; birds with similarly numerous foramina have sensitive beaks used to feel for food, so ''Lonchodraco'' may have used its beak to feel for fish or invertebrates in shallow water.<ref>{{Cite journal |last1=Martill |first1=David M. |last2=Smith |first2=Roy E. |last3=Longrich |first3=Nicholas |last4=Brown |first4=James |date=2021-01-01 |title=Evidence for tactile foraging in pterosaurs: a sensitive tip to the beak of Lonchodraco giganteus (Pterosauria, Lonchodectidae) from the Upper Cretaceous of southern England |url=https://www.sciencedirect.com/science/article/pii/S0195667120303232 |journal=Cretaceous Research |language=en |volume=117 |pages=104637 |doi=10.1016/j.cretres.2020.104637 |bibcode=2021CrRes.11704637M |s2cid=225130037 |issn=0195-6671}}</ref> The [[Istiodactylidae|istiodactylids]] were likely primarily scavengers.{{sfn|Witton|2013|pp=150–51}} [[Archaeopterodactyloidea]] obtained food in coastal or freshwater habitats. ''[[Germanodactylus]]'' and ''[[Pterodactylus]]'' were piscivores, while the [[Ctenochasmatidae]] were suspension feeders, using their numerous fine teeth to filter small organisms from shallow water. ''[[Pterodaustro]]'' was adapted for [[flamingo]]-like filter-feeding.{{sfn|Witton|2013|p=199}}
[[File:Kariridraco_by_Júlia_d’Oliveira.jpg|thumb|left|[[Azhdarchoid]] pterosaurs such as ''[[Kariridraco]]'' fed on terrestrial prey]]
In contrast, [[Azhdarchoidea]] mostly were terrestrial pterosaurs. [[Tapejaridae]] were arboreal omnivores, supplementing seeds and fruits with small insects and vertebrates.<ref name="witton2013"/><ref>{{cite journal | last1 = Wu | first1 = Wen-Hao | last2 = Zhou | first2 = Chang-Fu | last3 = Andres | first3 = Brian | year = 2017 | title = The toothless pterosaur ''Jidapterus edentus'' (Pterodactyloidea: Azhdarchoidea) from the Early Cretaceous Jehol Biota and its paleoecological implications | journal = PLOS ONE | volume = 12 | issue = 9| page = e0185486 | doi = 10.1371/journal.pone.0185486 | pmid = 28950013 | pmc = 5614613 | bibcode = 2017PLoSO..1285486W | doi-access = free }}</ref> [[Dsungaripteridae]] were specialist molluscivores, using their powerful jaws to crush the shells of molluscs and crustaceans. [[Thalassodromidae]] were likely terrestrial carnivores. ''[[Thalassodromeus]]'' itself was named after a fishing method known as "skim-feeding", later understood to be biomechanically impossible. Perhaps it pursued relatively large prey, in view of its reinforced jaw joints and relatively high bite force.<ref>Pêgas, R. V., & Kellner, A. W. (2015). Preliminary mandibular myological reconstruction of ''Thalassodromeus sethi'' (Pterodactyloidea: Tapejaridae). Flugsaurier 2015 Portsmouth, abstracts, 47–48</ref> [[Azhdarchidae]] are now understood to be terrestrial predators akin to ground [[hornbills]] or some [[storks]], eating any prey item they could swallow whole.<ref name="wittonnaish2015">{{cite journal | first1 = M.P. | last1 = Witton | first2 = D. | last2 = Naish | title = Azhdarchid pterosaurs: water-trawling pelican mimics or "terrestrial stalkers"? | journal = Acta Palaeontologica Polonica | date = 2015 | volume = 60 | issue = 3 | doi = 10.4202/app.00005.2013| doi-access = free }}</ref> ''[[Hatzegopteryx]]'' was a robustly built predator of relatively large prey, including medium-sized dinosaurs.<ref name="witton2017">{{cite journal | first2 = M.P. | last2 = Witton | first1 = D. | last1 = Naish | title = Neck biomechanics indicate that giant Transylvanian azhdarchid pterosaurs were short-necked arch predators | volume = 5 | doi = 10.7717/peerj.2908 | pmid = 28133577 | pmc = 5248582 | journal = PeerJ | date = 2017 | page=e2908 | doi-access = free }}</ref><ref>{{cite conference | last1 = Witton | first1 = M. | last2 = Brusatte | first2 = S. | last3 = Dyke | first3 = G. | last4 = Naish | first4 = D. | last5 = Norell | first5 = M. | last6 = Vremir | first6 = M. | title = Pterosaur overlords of Transylvania: short-necked giant azhdarchids in Late Cretaceous Romania | conference = The Annual Symposium of Vertebrate Paleontology and Comparative Anatomy | date = 2013 | location = Edinburgh | url = http://svpca.org/abstracts/abstract.php?abstID=00000001864&prog=on | url-status = dead | archive-url = https://web.archive.org/web/20160406020702/http://svpca.org/abstracts/abstract.php?abstID=00000001864&prog=on | archive-date = 2016-04-06}}</ref> ''[[Alanqa]]'' may have been a specialist molluscivore.<ref name="martillandibrahim2015">{{cite journal |last1=Martill |first1=David M. |last2=Ibrahim |first2=Nizar |title=An unusual modification of the jaws in cf. Alanqa, a mid-Cretaceous azhdarchid pterosaur from the Kem Kem beds of Morocco |journal=Cretaceous Research |date=March 2015 |volume=53 |pages=59–67 |doi=10.1016/j.cretres.2014.11.001 |bibcode=2015CrRes..53...59M |url=https://researchportal.port.ac.uk/portal/en/publications/an-unusual-modification-of-the-jaws-in-cf-alanqa-a-midcretaceous-azhdarchid-pterosaur-from-the-kem-kem-beds-of-morocco(ce004df9-c86a-4cf9-9fb8-1172211774cc).html }}</ref>

A 2021 study reconstructed the adductor musculature of skulls from [[Pterodactyloidea|pterodactyloids]], estimating the bite force and potential dietary habits of nine selected species.<ref name="pegas">{{cite journal |last1=Pêgas |first1=Rodrigo V |last2=Costa |first2=Fabiana R |last3=Kellner |first3=Alexander W A |title=Reconstruction of the adductor chamber and predicted bite force in pterodactyloids (Pterosauria) |journal=Zoological Journal of the Linnean Society |date=24 September 2021 |volume=193 |issue=2 |pages=602–635 |doi=10.1093/zoolinnean/zlaa163 }}</ref> The study corroborated the view of [[pteranodontids]], [[nyctosaurids]] and [[Anhanguera (pterosaur)|anhanuerids]] as [[piscivores]] based on them being relatively weak but fast biters, and suggest that ''[[Tropeognathus mesembrinus]]'' was specialised in consuming relatively large prey compared to ''[[Anhanguera (pterosaur)|Anhanguera]]''. ''[[Dsungaripterus]]'' was corroborated as a [[durophagy|durophage]], with ''[[Thalassodromeus]]'' proposed to share this feeding habit based on high estimated [[bite force quotient]]s (BFQ) and absolute bite force values.<ref name="pegas"/> ''[[Tapejara wellnhoferi]]'' was corroborated as a specialised consumer of hard plant material with a relatively high BFQ and high mechanical advantage, and ''[[Caupedactylus ybaka]]'' and ''[[Tupuxuara leonardii]]'' were proposed to be ground-feeding generalists with intermediate bite force values and less specialised jaws.<ref name="pegas"/>