Temporal range: Early Tithonian,
150.8–148.5 Ma
Bsp as i 739 modified.png
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Order: Pterosauria
Suborder: Pterodactyloidea
Clade: Pterodactylidae
Genus: Pterodactylus
Cuvier, 1809
Type species
Pterodactylus antiquus

Pterodactylus (meaning "winged finger"[2]) is an extinct genus of pterosaurs, whose members are commonly known as pterodactyls ( /ˌtɛrəˈdæktɪlz/). It is thought to contain only a single species, Pterodactylus antiquus, the first pterosaur species to be named and identified as a flying reptile.

The fossil remains of this species have been found primarily in the Solnhofen limestone of Bavaria, Germany, dated to the late Jurassic Period (early Tithonian), about 150.8–148.5 million years ago,[3] though more fragmentary remains have been tentatively identified from elsewhere in Europe and in Africa.




Pterodactylus is known from over 30 fossil specimens, and though most of those are juveniles, many preserve complete skeletons.[4][5] Pterodactylus antiquus was a relatively small pterosaur, with an estimated adult wingspan of about 1.04 meters (3 feet 5 inches) (the only known adult specimen is represented by an isolated skull).[4] Other "species" were once thought to be smaller. However, these smaller specimens have been shown to represent juveniles of Pterodactylus, as well as its contemporary relatives including Ctenochasma, Germanodactylus, Aurorazhdarcho, Gnathosaurus (and hypothetically Aerodactylus if this genus is truly valid).[6]


Life restoration of BMMS 7, the largest known Pterodactylus specimen. The shape of the soft tissue crest is based on BSP 1929.

Pterodactylus, like related pterosaurs, had a crest on its skull composed mainly of soft tissues. In adult Pterodactylus, this crest extended between the back edge of the antorbital fenestra (the largest opening in the skull) and the back of the skull. In at least one specimen, the crest had a short bony base, also seen in related pterosaurs like Germanodactylus. Solid crests have only been found on large, fully adult specimens of Pterodactylus, indicating that this was a display structure that became larger and more well developed as individuals reached maturity.[4][8] Bennett (2013) noted that other authors claimed that the soft tissue crest of Pterodactylus extended backward behind the skull; Bennett himself, however, didn't find any evidence for the crest extending past the back of the skull.[4] Two specimens of P. antiquus (the holotype specimen BSP AS I 739 and the incomplete skull BMMS 7, the largest known skull of P. antiquus) have a low bony crest on their skulls; in BMMS 7 it is 47.5 mm long (1.87 inches, more or less 24% of the estimated total length of its skull) and has a maximum height of 0.9 mm (0.035 inches) above the orbit.[4] Several specimens previously referred to P. antiquus preserved evidence of the soft tissue extensions of these crests, including an "occipital lappet", a flexible, tab-like structure extending from the back of the skull. Most of these specimens have been reclassified in the related species Aerodactylus scolopaciceps, which may however be nothing more than a junior synonym. Even if Aerodactylus were valid, at least one specimen with these features is still considered to belong to Pterodactylus. This is BSP 1929 I 18, which has an occipital lappet similar to the proposed Aerodactylus definition. This specimen also has a small triangular soft tissue crest with the peak of the crest positioned above the eyes.[4]


Year classes

Juvenile specimen of P. antiquus


The youngest immature Pterodactylus antiquus specimens (alternately interpreted as young specimens of the distinct species P. kochi) have a small number of teeth (as few as 15), and the teeth have a relatively broad base.[5] The teeth of other P. antiquus specimens are both narrower and more numerous (up to 90 teeth are present in some specimens).[7]

Pterodactylus specimens can be divided into two distinct year classes. In the first year class, the skulls are only 15-45mm (0.59-1.77 inches) in length. The second year class is characterized by skulls 55-95mm (2.16-3.74 inches) long, but still immature. These first two size groups were once classified as juveniles and adults of the species P. kochi, until further study showed that even the supposed "adults" were immature, and possibly belong to a distinct genus. A third year class is represented by specimens of the "traditional" P. antiquus, as well as a few isolated, large specimens once assigned to P. kochi that overlap P. antiquus in size. However, all specimens in this third year class also show sign of immaturity. Fully mature Pterodactylus specimens remain unknown, or may have been mistakenly classified as a different genus.[5]

Growth and breeding seasons



Daily activity patterns



Original copper engraving of the P. antiquus holotype by Egid Verhelst II and published by Italian scientist Cosimo Alessandro Collini, 1784


Wagler's 1830 restoration of an aquatic Pterodactylus


Hermann's original life restoration, the first of any pterosaur, 1800



Von Soemmerring's incorrect 1817 restoration of P. brevirostris

Contrary to von Moll's report, the fossil was not missing; it was being studied by Samuel Thomas von Sömmerring, who gave a public lecture about it on 27 December 1810. In January 1811, von Sömmerring wrote a letter to Cuvier deploring the fact that he had only recently been informed of Cuvier's request for information. His lecture was published in 1812, and in it von Sömmerring named the species Ornithocephalus antiquus.[18] The animal was described as being both a mammal, a bat, and a form in between mammals and birds, i.e. not intermediate in descent but in "affinity" or archetype. Cuvier disagreed, and the same year in his Ossemens fossiles provided a lengthy description in which he restated that the animal was a reptile.[19] It was not until 1817 that a second specimen of Pterodactylus came to light, again from Solnhofen. This tiny specimen was that year described by von Soemmerring as Ornithocephalus brevirostris, named for its short snout, now understood to be a juvenile character (this specimen is now thought to represent a juvenile specimen of a different genus, probably Ctenochasma[4]). He provided a restoration of the skeleton, the first one published for any pterosaur.[15] This restoration was very inaccurate, von Soemmerring mistaking the long metacarpals for the bones of the lower arm, the lower arm for the humerus, this upper arm for the breast bone and this sternum again for the shoulder blades.[20] Soemmerring did not change his opinion that these forms were bats and this "bat model" for interpreting pterosaurs would remain influential long after a consensus had been reached around 1860 that they were reptiles. The standard assumptions were that pterosaurs were quadrupedal, clumsy on the ground, furred, warmblooded and had a wing membrane reaching the ankle.[21] Some of these elements have been confirmed, some refuted by modern research, while others remain disputed.


A P. antiquus specimen (AMNH 1942) showing muscle impressions

The genus now known as Pterodactylus was originally named Petro-Dactyle by Cuvier in 1809,[22] though this was a typographical error, later corrected by him to Ptéro-Dactyle.[15] In 1812, Samuel Thomas von Sömmerring named the same specimen Ornithocephalus antiquus. The genus name was emended to the current Pterodactylus by Constantine Samuel Rafinesque in 1815. Unaware of Rafinesque's publication, Cuvier himself in 1819 again emended the genus name,[23] but the specific name he then gave, longirostris, has to give precedence to von Soemmerring's antiquus. In 1888 Richard Lydekker designated Pterodactylus antiquus the type species. The original specimen is the holotype of the genus, BSP No. AS.I.739.

Hermann von Meyer, in 1830, used the name Pterodactyli to contain Pterodactylus and other pterosaurs known at the time. This was emended to the family Pterodactylidae by Prince Charles Lucien Bonaparte in 1838. This group has more recently been given several competing definitions.[24][25]



Normannognathus wellnhoferi

Germanodactylus cristatus

Germanodactylus rhamphastinus


Pterodactylus antiquus

Pterodactylus kochi

Ardeadactylus longicollum


Former species

Specimen of P. kochi, Bürgermeister Müller Museum
Pterodactylus kochi

Numerous species have been assigned to Pterodactylus in the years since its discovery. In the first half of the nineteenth century any new pterosaur species would be named Pterodactylus, which thus became a typical "wastebasket taxon". Even after clearly different forms had later been given their own generic name, new species would be created from the very productive late Jurassic German sites, often based on only slightly different material.

Around 1980, subsequent revisions by Peter Wellnhofer had reduced the number of recognized species to about half a dozen. Many species assigned to Pterodactylus had been based on juvenile specimens, and subsequently been recognized as immature individuals of other species or genera. By the 1990s it was understood that this was even true for part of the remaining species. P. elegans, for example, was found by numerous studies to be an immature Ctenochasma.[7] Another species of Pterodactylus originally based on small, immature specimens was P. micronyx. However, it has been difficult to determine exactly of what genus and species P. micronyx might be the juvenile form. Stéphane Jouve, Christopher Bennett and others had once suggested that it probably belonged either to Gnathosaurus subulatus or one of the Ctenochasma species,[6][7] though after additional research Bennett assigned it to the genus Aurorazhdarcho.[4] Another species with a complex history is P. longicollum, named by von Meyer in 1854, based on a large specimen with a long neck and fewer teeth. Many researchers, including David Unwin, have found P. longicollum to be distinct from P. kochi and P. antiquus. Unwin found P. longicollum to be closer to Germanodactylus and therefore requiring a new genus name.[24] It has sometimes been placed in the genus Diopecephalus because Harry Govier Seeley based this genus partly on the P. longicollum material. However, it was shown by Bennett that the type specimen later designated for Diopecephalus was a fossil belonging to P. kochi, and no longer thought to be separate from Pterodactylus. Diopecephalus is therefore a synonym of Pterodactylus, and as such is unavailable for use as a new genus for "P." longicollum.[27] "P." longicollum was eventually made the type species of a separate genus Ardeadactylus.[4]

Juvenile type specimen of P. kochi

The only well-known and well-supported species left by the first decades of the 21st century were P. antiquus and P. kochi. However, most studies between 1995 and 2010 found little reason to separate even these two species, and treated them as synonymous.[24][27] In 1996, Bennett suggested that the differences between specimens of P. kochi and P. antiquus could be explained by differences in age, with P. kochi (including specimens alternately classified in the species P. scolopaciceps) representing an immature growth stage of P. antiquus. In a 2004 paper, Jouve used a different method of analysis and recovered the same result, showing that the "distinctive" features of P. kochi were age-related, and using mathematical comparison to show that the two forms are different growth stages of the same species.[7] An additional review of the specimens published in 2013 demonstrated that some of the supposed differences between P. kochi and P. antiquus were due to measurement errors, further supporting their synonymy.[4]

One species or three?

By the 2010s, a large body of research had been developed based on the idea that P. kochi and P. scolopaciceps were early growth stages of P. antiquus. However, in 2014, two scientists began publishing research that challenged this paradigm. Steven Vidovic and David Martill concluded that differences between specimens of P. kochi, P. scolopaciceps, and P. antiquus, such as different lengths of neck vertebrae, thinner or thicker teeth, more rounded skulls, and how far the teeth extended back in the jaws, were significant enough to separate them into three distinct species. Vidovic and Martill also performed a phylogenetic analysis which treated all relevant specimens as distinct units, and found that the P. kochi type specimen did not form a natural group with that of P. antiquus. They concluded that the genus Diopecephalus could be returned to use to distinguish "P". kochi from P. antiquus. They named the new genus Aerodactylus for P. scolopaciceps as well. So, what Bennett considered early growth stages of one species, Vidovic and Martill considered representatives of new species.[28][29]

In 2017, Bennett challenged this hypothesis. He claimed that while Vidovic and Martill had identified real differences between the these three groups of specimens, they had not provided any rationale that the differences were enough to distinguish them as species, rather than just individual variation, growth changes, or simply due to crushing and distortion during the fossilization process. Bennett pointed in particular to the data used to distinguish "Aerodactylus", which was so different from the data for related species, it might be due to an unnatural assemblage of specimens. As a result, Bennett continued to consider Diopecephalus and Aerodactylus simply as year-classes of immature Pterodactylus antiquus.[30]

List of species and synonyms


Name Author Year Status Notes
Ornithocephalus brevirostris von Sömmerring 1816–17 Synonym of Ctenochasma elegans[4] Based on a juvenile specimen
Pterodactylus longirostris Cuvier 1819 Synonym of Pterodactylus antiquus

Pterodactylus crocodilocephaloides Ritgen 1826 Synonym of Pterodactylus antiquus
Pterodactylus spectabilis von Meyer 1861 Synonym of Pterodactylus antiquus
Pterodactylus grandis Cuvier 1825 Nomen dubium, possible synonym of Rhamphorhynchus muensteri
Ornithocephalus kochi Wagner 1837 Synonym of Pterodactylus kochi
Pterodactylus meyeri Muenster 1842 Synonym of Pterodactylus kochi
Pterodactylus grandipelvis von Meyer 1860 Nomen dubium
Pterodactylus cerinensis von Meyer 1860 Nomen dubium
Pterodactylus suprajurensis Sauvage 1873 Nomen dubium
Pterodactylus manseli Owen 1874 Nomen dubium
Pterodactylus pleydelli Owen 1874 Nomen dubium
Pterodactylus arningi Reck 1931 Nomen dubium
Pterodactylus maximus Reck 1931 Nomen dubium

See also


  2. ^ Gudger, E.W. (Aug 1944). "The Earliest Winged Fish-Catchers". The Scientific Monthly. 59 (2): 120–129. Bibcode:1944SciMo..59..120G. JSTOR 18398.
  3. ^ Schweigert, G. (2007). "Ammonite biostratigraphy as a tool for dating Upper Jurassic lithographic limestones from South Germany – first results and open questions". Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 245 (1): 117–125. doi:10.1127/0077-7749/2007/0245-0117.
  4. ^ a b c d e f g h i j k l Bennett, S. Christopher (2013). "New information on body size and cranial display structures of Pterodactylus antiquus, with a revision of the genus". Paläontologische Zeitschrift. in press (2): 269–289. doi:10.1007/s12542-012-0159-8.
  5. ^ a b c d e Bennett, S.C. (1996). "Year-classes of pterosaurs from the Solnhofen Limestone of Germany: Taxonomic and Systematic Implications". Journal of Vertebrate Paleontology. 16 (3): 432–444. doi:10.1080/02724634.1996.10011332.
  7. ^ a b c d e f Jouve, S. (2004). "Description of the skull of a Ctenochasma (Pterosauria) from the latest Jurassic of eastern France, with a taxonomic revision of European Tithonian Pterodactyloidea". Journal of Vertebrate Paleontology. 24 (3): 542–554. doi:10.1671/0272-4634(2004)024[0542:DOTSOA]2.0.CO;2.
  8. ^ Frey, E. & Martill, D.M. (1998). "Soft tissue preservation in a specimen of Pterodactylus kochi (Wagner) from the Upper Jurassic of Germany". Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 210 (3): 421–441. doi:10.1127/njgpa/210/1998/421.
  15. ^ a b c d Taquet, P. & Padian, K. (2004). "The earliest known restoration of a pterosaur and the philosophical origins of Cuvier's Ossemens Fossiles". Comptes Rendus Palevol. 3 (2): 157–175. doi:10.1016/j.crpv.2004.02.002.
  20. ^ Sömmering, T. v. (1817). "Über einen Ornithocephalus brevirostris der Vorwelt". Denkschr. KGL. Bayer Akad. Wiss., Math.phys. Cl. 6: 89–104.
  27. ^ a b Bennett, S. Christopher (2006). "Juvenile specimens of the pterosaur Germanodactylus cristatus, with a review of the genus". Journal of Vertebrate Paleontology. 26 (4): 872–878. doi:10.1671/0272-4634(2006)26[872:JSOTPG]2.0.CO;2.
  28. ^ Vidovic, S. U.; Martill, D. M. (2014). "Pterodactylus scolopaciceps Meyer, 1860 (Pterosauria, Pterodactyloidea) from the Upper Jurassic of Bavaria, Germany: The Problem of Cryptic Pterosaur Taxa in Early Ontogeny". PLoS ONE. 9 (10): e110646. doi:10.1371/journal.pone.0110646. PMC 4206445. PMID 25337830.
  29. ^ Vidovic, Steven U.; Martill, David M. (2017). "The taxonomy and phylogeny of Diopecephalus kochi (Wagner, 1837) and Germanodactylus rhamphastinus (Wagner, 1851)" (PDF). Geological Society, London, Special Publications. 455: 125–147. doi:10.1144/SP455.12.
  30. ^ Bennett, S.C. (2017). New smallest specimen of the pterosaur Pteranodon and ontogenetic niches in pterosaurs. Journal of Paleontology, 1-18. doi:10.1017/jpa.2017.84