Pterodactylus
Encyclopedia
Pterodactylus is a genus
of pterosaur
s, whose members are popularly known as pterodactyls. It was the first to be named and identified as a flying reptile. Its fossil
remains 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, though more fragmentary remains have been identified from elsewhere in Europe
and in Africa
. It was a carnivore
and probably preyed upon fish and other small animals. Like all pterosaurs, the wings of Pterodactylus were formed by a skin and muscle membrane stretching from its elongated fourth finger to its hind limbs. It was supported internally by collagen
fibres and externally by keratin
ous ridges.
The name derives from the Greek
words pteron (πτερόn, meaning 'wing') and daktylos (δάκτυλος, meaning 'finger') and refers to the way in which the wing is supported by one large finger.
specimens, and though most of those are juveniles, many preserve complete skeletons. The discovery of several specimens with well-preserved soft tissue traces has allowed scientists to faithfully reconstruct the life appearance of Pterodactylus. Pterodactylus was a relatively small pterosaur, with an estimated adult wingspan of about 1.5 meters (5 ft) in P. antiquus. 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 Ctenochasma
, Germanodactylus
and Gnathosaurus
.
The skulls of adult Pterodactylus were long and narrow with about 90 large, conical teeth. The teeth extended back from the tips of both jaws, and became smaller farther away from the jaw tips (unlike some relatives, where teeth were absent in the upper jaw tip and were relatively uniform in size). The teeth extended farther back into the jaw than in close relatives, as some were present below the front of the nasoantorbital fenestra, the largest opening in the skull. Unlike related species, the skull and jaws were straight, not curved upwards. A small, hooked beak was present in the very tips of the jaws, with both upper and lower hook no larger than the teeth that surrounded them.
The neck was long, and covered in long, bristle-like pycnofibres. A throat pouch extended from about the middle of the lower jaw to the upper part of the neck.
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. The back of the crest extended upward into a backward-curving cone-shaped structure. The crest was composed mainly of long, hardened fibres (twisted together in a spiral pattern inside the conical part of the crest), and covered in scales. In at least one specimen of P. longicollum, the crest had a short bony base, also seen in related pterosaurs like Germanodactylus. Crests have only been found on large, fully adult specimens of Pterodactylus, indicating that this was a display structure and only developed when individuals reached maturity.
The wings were long, and the wing membranes appear to have lacked the furry covering of pycnofibres present in some other pterosaurs (such as Pterorhynchus
and Jeholopterus
). The wing membrane extended between the fingers and toes as webbing, and a uropatagium (secondary membrane between the feet and tail) was present, as well as a propatagium (membrane between the wrist and shoulder). Both the finger and toe claws were covered in keratin sheaths that extended and curved into sharp hooks well beyond their bony cores.
Like other pterosaurs (notably Rhamphorhynchus), Pterodactylus specimens can vary considerably based on age or level of maturity. Both the proportions of the limb bones, size and shape of the skull, and size and number of teeth changed as the animals grew. Historically, this has led to various growth stages (including growth stages of related pterosaurs) being mistaken for new species of Pterodactylus. Several detailed studies using various methods to measure growth curves among known specimens have demonstrated that there is actually only one valid Pterodactylus species, P. antiquus.
The youngest immature Pterodactylus specimens have a small number of teeth (as few as 15), and the teeth have a relatively broad base. The teeth of older specimens are both narrower and more numerous (up to 90 teeth are present in some specimens).
Pterodactylus specimens can be divided into two distinct year classes. In the first year class, the skulls are only 15-45mm in length. The second year class is characterized by skulls 55-95mm 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. 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.
ns, rather than the rapid growth of modern bird
s.
s of Pterodactylus antiquus and modern birds and reptiles suggest that it may have been diurnal. This may also indicate niche partitioning with contemporary pterosaurs inferred to be nocturnal, such as Ctenochasma
and Rhamphorhynchus
.
ever to be identified. The first Pterodactylus specimen was described by the Italian scientist Cosimo Alessandro Collini in 1784, based on a fossil
skeleton unearthed from the Solnhofen limestone
of Bavaria
. Collini was the curator of the "Naturalienkabinett", or nature cabinet
(a precursor to the modern concept of the natural history
museum), in the palace of Charles Theodore, Elector of Bavaria
at Mannheim
. The specimen had been given to the collection by Count Friedrich Ferdinand zu Pappenheim
, probably around 1780, having been recovered from a lithographic limestone
quarry in Eichstätt
. Collini, however, did not conclude that the specimen was a flying animal. In fact, Collini could not fathom what kind of animal it might have been, rejecting affinities with the birds or the bats. He speculated that it may have been a sea creature, not for any anatomical reason, but because he thought the ocean depths were more likely to have housed unknown types of animals. The idea that pterosaurs were aquatic animals persisted among a minority of scientists as late as 1830, when the German zoologist Johann Georg Wagler
published a text on "amphibians" which included an illustration of Pterodactylus using its wings as flippers. Wagler went so far as to classify Pterodactylus, along with other aquatic vertebrates (namely plesiosaur
s, ichthyosaur
s, and monotreme
s), in the class Gryphi, between birds and mammals.
It was the German/French scientist Johann Hermann
who first stated that Pterodactylus used its long fourth finger to support a wing membrane. In March 1800, Hermann alerted the French scientist George Cuvier to the existence of Collini's fossil, believing that it had been captured by the occupying armies of Napoleon and sent to the French collections in Paris
(and perhaps to Cuvier himself) as war booty; at the time special French political commissar
s systematically seized art treasures and objects of scientific interest. Hermann sent Cuvier a letter containing his own interpretation of the specimen (though he had not examined it personally), which he believed to be a mammal
, including the first known life restoration of a pterosaur. Hermann restored the animal with wing membranes extending from the long fourth finger to the ankle and a covering of fur (neither wing membranes nor fur had been preserved in the specimen). Hermann also added a membrane between the neck and wrist, as is the condition in bat
s. Cuvier agreed with this interpretation, and at Hermann's suggestion, Cuvier became the first to publish these ideas in December 1800 in a very short description. Cuvier remarked, "[It is not possible to doubt that the long finger served to support a membrane that, by lengthening the anterior extremity of this animal, formed a good wing.]" However, contrary to Hermann, Cuvier was convinced the animal was a reptile
.
The specimen had not in fact been seized by the French. Rather, in 1802, following the death of Charles Theodore, it was brought to Munich
, where Baron Johann Paul Carl von Moll had obtained a general exemption of confiscation for the Bavarian collections. Cuvier asked von Moll to study the fossil but was informed it could not be found. In 1809 Cuvier published a somewhat longer description, naming the animal, in which he refuted a hypothesis by Johann Friedrich Blumenbach
that it would have been a shore bird.
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. 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. 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 (for its short snout, now understood to be a juvenile character), and provided a restoration of the skeleton, the first one published for any pterosaur. 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. Von 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. Some of these elements have been confirmed, some refuted by modern research, while others remain disputed.
In 1998, the discovery of one specimen assigned to P. kochi shed light on the life appearance of Pterodactylus, as it preserved unique soft-tissue traits not present in previous fossil skeletons, including long, bristly pycnofibres (a fur-like body covering known only in pterosaurs) on the neck, details of a urpatagium (hind wing membrane between the legs and tail) that also stretched between the toes as webbing
, and a pelican
-like throat pouch. An additional specimen, studied using ultra-violet light, revealed even more information on the soft anatomy of Pterodactylus. This specimen (catelogue number JME
SOS 4784) showed that like many other pterosaurs, Pterodactylus had a striated soft-tissue crest on the skull. Soft tissue impressions also showed unusually long, sharp, and recurved keratin sheaths on its claws. This specimen was also covered in hair-like pycnofibres, with unusually long pycnofibres covering the back of its neck. The remains of a small, hooked beak
were preserved at the tips of the jaws between its upper and lower front teeth.
The genus
now known as Pterodactylus was originally named Petro-Dactyle by Cuvier in 1809, though this was a typographical error, later corrected by him to Ptéro-Dactyle. In 1812, Samuel Thomas von Sömmerring named the same specimen Ornithocephalus antiquus. As the senior synonym, Cuvier's name has precedence, so applying modern rules for the combination of the genus name and the specific epithet, the valid species name became Ptéro-Dactyle antiquus. The genus name was Latinized to the current Pterodactylus by Constantine Samuel Rafinesque in 1815, which by these rules is the valid name as they do not allow for diacritic
s or hyphens. Unaware of Rafinesque's publication Cuvier himself in 1819 again Latinized the genus, 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 family has more recently been used to refer to many similar species from Germany and elsewhere, though recent studies suggest it may be a paraphyletic or polyphyletic unnatural grouping with respect to more advanced members of the Ctenochasmatoidea
(or Archaeopterodactyloidea).
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 "waste-basket 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
. Another species of Pterodactylus based on small, immature specimens is 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 suggested that it probably belongs either to Gnathosaurus subulatus
or one of the Ctenochasma species, but more data and study would be required to determine which one.
The only well-known and well-supported species left were P. antiquus and P. kochi. However, most studies since the 1990s have found little reason to separate even these two, and have treated them as synonymous. In 1996, Bennett suggested that the differences between specimens of P. kochi and P. antiquus could be explained by differences in age. 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.
A special case 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. 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.
("doubtful names"). The following list includes names that are based on German material presently, or until recently, thought to be pertaining to Pterodactylus proper and names based on other material that has as yet not been assigned to other genera.
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
of pterosaur
Pterosaur
Pterosaurs were flying reptiles of the clade or order Pterosauria. They existed from the late Triassic to the end of the Cretaceous Period . Pterosaurs are the earliest vertebrates known to have evolved powered flight...
s, whose members are popularly known as pterodactyls. It was the first to be named and identified as a flying reptile. Its fossil
Fossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...
remains have been found primarily in the Solnhofen limestone
Solnhofen limestone
The Solnhofen Plattenkalk is a Jurassic Konservat-Lagerstätte that preserves a rare assemblage of fossilized organisms, including highly detailed imprints of soft bodied organisms such as sea jellies...
of Bavaria
Bavaria
Bavaria, formally the Free State of Bavaria is a state of Germany, located in the southeast of Germany. With an area of , it is the largest state by area, forming almost 20% of the total land area of Germany...
, Germany
Germany
Germany , officially the Federal Republic of Germany , is a federal parliamentary republic in Europe. The country consists of 16 states while the capital and largest city is Berlin. Germany covers an area of 357,021 km2 and has a largely temperate seasonal climate...
, dated to the late Jurassic
Jurassic
The Jurassic is a geologic period and system that extends from about Mya to Mya, that is, from the end of the Triassic to the beginning of the Cretaceous. The Jurassic constitutes the middle period of the Mesozoic era, also known as the age of reptiles. The start of the period is marked by...
Period (early Tithonian
Tithonian
In the geologic timescale the Tithonian is the latest age of the Late Jurassic epoch or the uppermost stage of the Upper Jurassic series. It spans the time between 150.8 ± 4 Ma and 145.5 ± 4 Ma...
), about 150.8-148.5 million years ago, though more fragmentary remains have been identified from elsewhere in Europe
Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...
and in Africa
Africa
Africa is the world's second largest and second most populous continent, after Asia. At about 30.2 million km² including adjacent islands, it covers 6% of the Earth's total surface area and 20.4% of the total land area...
. It was a carnivore
Carnivore
A carnivore meaning 'meat eater' is an organism that derives its energy and nutrient requirements from a diet consisting mainly or exclusively of animal tissue, whether through predation or scavenging...
and probably preyed upon fish and other small animals. Like all pterosaurs, the wings of Pterodactylus were formed by a skin and muscle membrane stretching from its elongated fourth finger to its hind limbs. It was supported internally by collagen
Collagen
Collagen is a group of naturally occurring proteins found in animals, especially in the flesh and connective tissues of mammals. It is the main component of connective tissue, and is the most abundant protein in mammals, making up about 25% to 35% of the whole-body protein content...
fibres and externally by keratin
Keratin
Keratin refers to a family of fibrous structural proteins. Keratin is the key of structural material making up the outer layer of human skin. It is also the key structural component of hair and nails...
ous ridges.
The name derives from the Greek
Ancient Greek
Ancient Greek is the stage of the Greek language in the periods spanning the times c. 9th–6th centuries BC, , c. 5th–4th centuries BC , and the c. 3rd century BC – 6th century AD of ancient Greece and the ancient world; being predated in the 2nd millennium BC by Mycenaean Greek...
words pteron (πτερόn, meaning 'wing') and daktylos (δάκτυλος, meaning 'finger') and refers to the way in which the wing is supported by one large finger.
Description
Pterodactylus is known from over 27 fossilFossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...
specimens, and though most of those are juveniles, many preserve complete skeletons. The discovery of several specimens with well-preserved soft tissue traces has allowed scientists to faithfully reconstruct the life appearance of Pterodactylus. Pterodactylus was a relatively small pterosaur, with an estimated adult wingspan of about 1.5 meters (5 ft) in P. antiquus. 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 Ctenochasma
Ctenochasma
Ctenochasma is a genus of Late Jurassic pterosaur belonging to the suborder pterodactyloidea. Three species are currently recognized: C. roemeri , C. taqueti, and C. elegans...
, Germanodactylus
Germanodactylus
Germanodactylus is a genus of dsungaripteroid pterodactyloid pterosaur from Late Jurassic-age rocks of Germany, including the Solnhofen limestone. Its specimens were long thought to pertain to Pterodactylus. Its head crest is a distinctive feature.-Description:Germanodactylus is described as...
and Gnathosaurus
Gnathosaurus
Gnathosaurus is a genus of ctenochasmatid pterosaur known from a single species, G. subulatus, described in 1833. This pterosaur had an estimated wingspan of about 1.7 meters. The slender, 28-cm-long skull had up to 130 needle-like teeth arranged laterally around the spoon-shaped tip...
.
The skulls of adult Pterodactylus were long and narrow with about 90 large, conical teeth. The teeth extended back from the tips of both jaws, and became smaller farther away from the jaw tips (unlike some relatives, where teeth were absent in the upper jaw tip and were relatively uniform in size). The teeth extended farther back into the jaw than in close relatives, as some were present below the front of the nasoantorbital fenestra, the largest opening in the skull. Unlike related species, the skull and jaws were straight, not curved upwards. A small, hooked beak was present in the very tips of the jaws, with both upper and lower hook no larger than the teeth that surrounded them.
The neck was long, and covered in long, bristle-like pycnofibres. A throat pouch extended from about the middle of the lower jaw to the upper part of the neck.
The wings were long, and the wing membranes appear to have lacked the furry covering of pycnofibres present in some other pterosaurs (such as Pterorhynchus
Pterorhynchus
Pterorhynchus was a genus of rhamphorhynchid "rhamphorhynchoid" pterosaur from the Late Jurassic-age Daohugou Formation of Inner Mongolia, China....
and Jeholopterus
Jeholopterus
Jeholopterus was a small anurognathid pterosaur from the Daohugou Beds of northeastern China , between 168 and 152 million years ago), preserved with hair and skin remains.-Naming:...
). The wing membrane extended between the fingers and toes as webbing, and a uropatagium (secondary membrane between the feet and tail) was present, as well as a propatagium (membrane between the wrist and shoulder). Both the finger and toe claws were covered in keratin sheaths that extended and curved into sharp hooks well beyond their bony cores.
Year classes
The youngest immature Pterodactylus specimens have a small number of teeth (as few as 15), and the teeth have a relatively broad base. The teeth of older specimens are both narrower and more numerous (up to 90 teeth are present in some specimens).
Pterodactylus specimens can be divided into two distinct year classes. In the first year class, the skulls are only 15-45mm in length. The second year class is characterized by skulls 55-95mm 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. 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.
Growth and breeding seasons
The distinct year classes of Pterodactylus antiquus specimens show that this species, like the contemporary Rhamphorhynchus muensteri, likely bred seasonally and grew consistently during its lifetime. A new generation of 1st year class P. antiquus would have been produced seasonally, and reached 2nd-year size by the time the next generation hatched, creating distinct 'clumps' of similarly-sized and aged individuals in the fossil record. The smallest size class probably consisted of individuals that had just begun to fly and were less than one year old. The second year class represents individuals one to two years old, and the rare third year class is composed of specimens over two years old. This growth pattern is similar to modern crocodiliaCrocodilia
Crocodilia is an order of large reptiles that appeared about 84 million years ago in the late Cretaceous Period . They are the closest living relatives of birds, as the two groups are the only known survivors of the Archosauria...
ns, rather than the rapid growth of modern bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...
s.
Daily activity patterns
Comparisons between the scleral ringSclerotic ring
Sclerotic rings are rings of bone found in the eyes of several groups of vertebrate animals, except for mammals and crocodilians. They can be made up of single bones or small bones together. They are believed to have a role in supporting the eye, especially in animals whose eyes are not spherical,...
s of Pterodactylus antiquus and modern birds and reptiles suggest that it may have been diurnal. This may also indicate niche partitioning with contemporary pterosaurs inferred to be nocturnal, such as Ctenochasma
Ctenochasma
Ctenochasma is a genus of Late Jurassic pterosaur belonging to the suborder pterodactyloidea. Three species are currently recognized: C. roemeri , C. taqueti, and C. elegans...
and Rhamphorhynchus
Rhamphorhynchus
Rhamphorynchus may refer to:*Rhamphorhynchus, a genus of pterosaur*Rhamphorhynchus, a former monotypic genus of orchid, containing only the species now called Aspidogyne mendoncae...
.
History
The animal now known as Pterodactylus was the first pterosaurPterosaur
Pterosaurs were flying reptiles of the clade or order Pterosauria. They existed from the late Triassic to the end of the Cretaceous Period . Pterosaurs are the earliest vertebrates known to have evolved powered flight...
ever to be identified. The first Pterodactylus specimen was described by the Italian scientist Cosimo Alessandro Collini in 1784, based on a fossil
Fossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...
skeleton unearthed from the Solnhofen limestone
Solnhofen limestone
The Solnhofen Plattenkalk is a Jurassic Konservat-Lagerstätte that preserves a rare assemblage of fossilized organisms, including highly detailed imprints of soft bodied organisms such as sea jellies...
of Bavaria
Bavaria
Bavaria, formally the Free State of Bavaria is a state of Germany, located in the southeast of Germany. With an area of , it is the largest state by area, forming almost 20% of the total land area of Germany...
. Collini was the curator of the "Naturalienkabinett", or nature cabinet
Cabinet of curiosities
A cabinet of curiosities was an encyclopedic collection in Renaissance Europe of types of objects whose categorical boundaries were yet to be defined. They were also known by various names such as Cabinet of Wonder, and in German Kunstkammer or Wunderkammer...
(a precursor to the modern concept of the natural history
Natural history
Natural history is the scientific research of plants or animals, leaning more towards observational rather than experimental methods of study, and encompasses more research published in magazines than in academic journals. Grouped among the natural sciences, natural history is the systematic study...
museum), in the palace of Charles Theodore, Elector of Bavaria
Charles Theodore, Elector of Bavaria
Charles Theodore, Prince-Elector, Count Palatine and Duke of Bavaria reigned as Prince-Elector and Count palatine from 1742, as Duke of Jülich and Berg from 1742 and also as Prince-Elector and Duke of Bavaria from 1777, until his death...
at Mannheim
Mannheim
Mannheim is a city in southwestern Germany. With about 315,000 inhabitants, Mannheim is the second-largest city in the Bundesland of Baden-Württemberg, following the capital city of Stuttgart....
. The specimen had been given to the collection by Count Friedrich Ferdinand zu Pappenheim
Pappenheim (state)
Pappenheim was a German statelet in western Bavaria, Germany, located on the Altmühl river between Treuchtlingen and Solnhofen, and south of Weißenburg. Pappenheim originated as a Lordship around 1030, and was raised to a county in 1628. Pappenheim was partitioned twice: between itself, Aletzheim,...
, probably around 1780, having been recovered from a lithographic limestone
Lithographic Limestone
Lithographic limestone is hard limestone that is sufficiently fine-grained, homogeneous and defect free to be used for lithography. Geologists use the term lithographic texture to refer to a grain size under 1/250 mm...
quarry in Eichstätt
Eichstätt
Eichstätt is a town in the federal state of Bavaria, Germany, and capital of the District of Eichstätt. It is located along the Altmühl River, at , and had a population of 13,078 in 2002. It is home to the Katholische Universität Eichstätt-Ingolstadt, the lone Catholic university in Germany. The...
. Collini, however, did not conclude that the specimen was a flying animal. In fact, Collini could not fathom what kind of animal it might have been, rejecting affinities with the birds or the bats. He speculated that it may have been a sea creature, not for any anatomical reason, but because he thought the ocean depths were more likely to have housed unknown types of animals. The idea that pterosaurs were aquatic animals persisted among a minority of scientists as late as 1830, when the German zoologist Johann Georg Wagler
Johann Georg Wagler
Johann Georg Wagler was a German herpetologist.Wagler was assistant to Johann Baptist von Spix, and became Director of the Zoological Museum at the University of Munich after Spix's death in 1826...
published a text on "amphibians" which included an illustration of Pterodactylus using its wings as flippers. Wagler went so far as to classify Pterodactylus, along with other aquatic vertebrates (namely plesiosaur
Plesiosaur
Plesiosauroidea is an extinct clade of carnivorous plesiosaur marine reptiles. Plesiosauroids, are known from the Jurassic and Cretaceous Periods...
s, ichthyosaur
Ichthyosaur
Ichthyosaurs were giant marine reptiles that resembled fish and dolphins...
s, and monotreme
Monotreme
Monotremes are mammals that lay eggs instead of giving birth to live young like marsupials and placental mammals...
s), in the class Gryphi, between birds and mammals.
It was the German/French scientist Johann Hermann
Johann Hermann
Johann, or Jean, Hermann was a French physician and naturalist. He was professor of medicine at the University of Strasbourg. He was the author of Tabula affinitatum animalium and Observationes zoologicae quibus novae complures, published posthumously in 1804...
who first stated that Pterodactylus used its long fourth finger to support a wing membrane. In March 1800, Hermann alerted the French scientist George Cuvier to the existence of Collini's fossil, believing that it had been captured by the occupying armies of Napoleon and sent to the French collections in Paris
Paris
Paris is the capital and largest city in France, situated on the river Seine, in northern France, at the heart of the Île-de-France region...
(and perhaps to Cuvier himself) as war booty; at the time special French political commissar
Political commissar
The political commissar is the supervisory political officer responsible for the political education and organisation, and loyalty to the government of the military...
s systematically seized art treasures and objects of scientific interest. Hermann sent Cuvier a letter containing his own interpretation of the specimen (though he had not examined it personally), which he believed to be a mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...
, including the first known life restoration of a pterosaur. Hermann restored the animal with wing membranes extending from the long fourth finger to the ankle and a covering of fur (neither wing membranes nor fur had been preserved in the specimen). Hermann also added a membrane between the neck and wrist, as is the condition in bat
Bat
Bats are mammals of the order Chiroptera "hand" and pteron "wing") whose forelimbs form webbed wings, making them the only mammals naturally capable of true and sustained flight. By contrast, other mammals said to fly, such as flying squirrels, gliding possums, and colugos, glide rather than fly,...
s. Cuvier agreed with this interpretation, and at Hermann's suggestion, Cuvier became the first to publish these ideas in December 1800 in a very short description. Cuvier remarked, "[It is not possible to doubt that the long finger served to support a membrane that, by lengthening the anterior extremity of this animal, formed a good wing.]" However, contrary to Hermann, Cuvier was convinced the animal was a reptile
Reptile
Reptiles are members of a class of air-breathing, ectothermic vertebrates which are characterized by laying shelled eggs , and having skin covered in scales and/or scutes. They are tetrapods, either having four limbs or being descended from four-limbed ancestors...
.
The specimen had not in fact been seized by the French. Rather, in 1802, following the death of Charles Theodore, it was brought to Munich
Munich
Munich The city's motto is "" . Before 2006, it was "Weltstadt mit Herz" . Its native name, , is derived from the Old High German Munichen, meaning "by the monks' place". The city's name derives from the monks of the Benedictine order who founded the city; hence the monk depicted on the city's coat...
, where Baron Johann Paul Carl von Moll had obtained a general exemption of confiscation for the Bavarian collections. Cuvier asked von Moll to study the fossil but was informed it could not be found. In 1809 Cuvier published a somewhat longer description, naming the animal, in which he refuted a hypothesis by Johann Friedrich Blumenbach
Johann Friedrich Blumenbach
Johann Friedrich Blumenbach was a German physician, physiologist and anthropologist, one of the first to explore the study of mankind as an aspect of natural history, whose teachings in comparative anatomy were applied to classification of what he called human races, of which he determined...
that it would have been a shore bird.
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. 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
Archetype
An archetype is a universally understood symbol or term or pattern of behavior, a prototype upon which others are copied, patterned, or emulated...
. Cuvier disagreed, and the same year in his Ossemens fossiles provided a lengthy description in which he restated that the animal was a reptile. 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 (for its short snout, now understood to be a juvenile character), and provided a restoration of the skeleton, the first one published for any pterosaur. This restoration was very inaccurate, von Soemmerring mistaking the long metacarpals for the bones of the lower arm, the lower arm for the humerus
Humerus
The humerus is a long bone in the arm or forelimb that runs from the shoulder to the elbow....
, this upper arm for the breast bone and this sternum again for the shoulder blades. Von 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. Some of these elements have been confirmed, some refuted by modern research, while others remain disputed.
In 1998, the discovery of one specimen assigned to P. kochi shed light on the life appearance of Pterodactylus, as it preserved unique soft-tissue traits not present in previous fossil skeletons, including long, bristly pycnofibres (a fur-like body covering known only in pterosaurs) on the neck, details of a urpatagium (hind wing membrane between the legs and tail) that also stretched between the toes as webbing
Webbed toes
Webbed toes is the common name for syndactyly affecting the feet. It is characterised by the fusion of two or more digits of the feet. This is normal in many birds, such as ducks; amphibians, such as frogs; and mammals, such as kangaroos...
, and a pelican
Pelican
A pelican, derived from the Greek word πελεκυς pelekys is a large water bird with a large throat pouch, belonging to the bird family Pelecanidae....
-like throat pouch. An additional specimen, studied using ultra-violet light, revealed even more information on the soft anatomy of Pterodactylus. This specimen (catelogue number JME
Jura Museum
The Jura Museum situated in Eichstätt, Germany is a Natural History Museum that has an extensive exhibit of Jurassic fossils from the quarry of Solnhofen, including marine reptiles, pterosaurs, and one specimen of the early bird Archaeopteryx...
SOS 4784) showed that like many other pterosaurs, Pterodactylus had a striated soft-tissue crest on the skull. Soft tissue impressions also showed unusually long, sharp, and recurved keratin sheaths on its claws. This specimen was also covered in hair-like pycnofibres, with unusually long pycnofibres covering the back of its neck. The remains of a small, hooked beak
Beak
The beak, bill or rostrum is an external anatomical structure of birds which is used for eating and for grooming, manipulating objects, killing prey, fighting, probing for food, courtship and feeding young...
were preserved at the tips of the jaws between its upper and lower front teeth.
Classification
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
now known as Pterodactylus was originally named Petro-Dactyle by Cuvier in 1809, though this was a typographical error, later corrected by him to Ptéro-Dactyle. In 1812, Samuel Thomas von Sömmerring named the same specimen Ornithocephalus antiquus. As the senior synonym, Cuvier's name has precedence, so applying modern rules for the combination of the genus name and the specific epithet, the valid species name became Ptéro-Dactyle antiquus. The genus name was Latinized to the current Pterodactylus by Constantine Samuel Rafinesque in 1815, which by these rules is the valid name as they do not allow for diacritic
Diacritic
A diacritic is a glyph added to a letter, or basic glyph. The term derives from the Greek διακριτικός . Diacritic is both an adjective and a noun, whereas diacritical is only an adjective. Some diacritical marks, such as the acute and grave are often called accents...
s or hyphens. Unaware of Rafinesque's publication Cuvier himself in 1819 again Latinized the genus, but the specific name he then gave, longirostris, has to give precedence to von Soemmerring's antiquus. In 1888 Richard Lydekker
Richard Lydekker
Richard Lydekker was an English naturalist, geologist and writer of numerous books on natural history.-Biography:...
designated Pterodactylus antiquus the type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...
. The original specimen is the holotype
Holotype
A holotype is a single physical example of an organism, known to have been used when the species was formally described. It is either the single such physical example or one of several such, but explicitly designated as the holotype...
of the genus, BSP No. AS.I.739.
Hermann von Meyer
Christian Erich Hermann von Meyer
Christian Erich Hermann von Meyer was a German palaeontologist.He was born at Frankfurt am Main.In 1832 von Meyer issued a work entitled Palaeologica, and in course of time he published a series of memoirs on various fossil organic remains: molluscs, crustaceans, fishes and higher vertebrata,...
, in 1830, used the name Pterodactyli to contain Pterodactylus and other pterosaurs known at the time. This was emended to the family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
Pterodactylidae by Prince Charles Lucien Bonaparte
Charles Lucien Bonaparte
Charles Lucien Jules Laurent Bonaparte, 2nd Prince of Canino and Musignano was a French naturalist and ornithologist.-Biography:...
in 1838. This family has more recently been used to refer to many similar species from Germany and elsewhere, though recent studies suggest it may be a paraphyletic or polyphyletic unnatural grouping with respect to more advanced members of the Ctenochasmatoidea
Ctenochasmatoidea
Ctenochasmatoidea is a group of pterosaurs within the suborder Pterodactyloidea.-Classification:Cladogram after Unwin ....
(or Archaeopterodactyloidea).
Species
Around 1980, subsequent revisions by Peter Wellnhofer
Peter Wellnhofer
Peter Wellnhofer is a German paleontologist at the "Bayerische Staatssammlung fur Paläontologie" in Munich. He is best known for his work on the various fossil specimens of Archaeopteryx or "Urvogel", the first known bird...
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
Ctenochasma
Ctenochasma is a genus of Late Jurassic pterosaur belonging to the suborder pterodactyloidea. Three species are currently recognized: C. roemeri , C. taqueti, and C. elegans...
. Another species of Pterodactylus based on small, immature specimens is 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 suggested that it probably belongs either to Gnathosaurus subulatus
Gnathosaurus
Gnathosaurus is a genus of ctenochasmatid pterosaur known from a single species, G. subulatus, described in 1833. This pterosaur had an estimated wingspan of about 1.7 meters. The slender, 28-cm-long skull had up to 130 needle-like teeth arranged laterally around the spoon-shaped tip...
or one of the Ctenochasma species, but more data and study would be required to determine which one.
The only well-known and well-supported species left were P. antiquus and P. kochi. However, most studies since the 1990s have found little reason to separate even these two, and have treated them as synonymous. In 1996, Bennett suggested that the differences between specimens of P. kochi and P. antiquus could be explained by differences in age. 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.
A special case 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. 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.
List of species and synonyms
During its over 200 year history, the various species of Pterodactylus have gone through a number of changes in classification, and thus have acquired a large number of synonyms. Additionally, a number of species assigned to Pterodactylus are based on poor remains that have proven difficult to assign to one species or another, and are therefore considered nomina dubiaNomen dubium
In zoological nomenclature, a nomen dubium is a scientific name that is of unknown or doubtful application...
("doubtful names"). The following list includes names that are based on German material presently, or until recently, thought to be pertaining to Pterodactylus proper and names based on other material that has as yet not been assigned to other genera.
| Name | Author | Year | Status | Notes |
|---|---|---|---|---|
| Pterodactylus antiquus | (von Sömmerring) Rafinesque | (1812 1812 in paleontology -Newly named plesiosaurs:Refer to article on these carnivorous aquatic reptiles.Plesiosaurs appeared at the start of the Jurassic Period, and thrived until the K-T extinction, at the end of the Cretaceous Period. While they were Mesozoic diapsid reptiles that lived at the same time as dinosaurs,... ) 1815 1815 in paleontology -New taxa:... |
Valid, type species | Originally Ptéro-dactyle, Cuvier 1809 1809 in paleontology -New taxa:... |
| Ornithocephalus antiquus | von Sömmerring | 1812 1812 in paleontology -Newly named plesiosaurs:Refer to article on these carnivorous aquatic reptiles.Plesiosaurs appeared at the start of the Jurassic Period, and thrived until the K-T extinction, at the end of the Cretaceous Period. While they were Mesozoic diapsid reptiles that lived at the same time as dinosaurs,... |
Synonym of Pterodactylus antiquus | Objective junior synonym of P. antiquus |
| Ornithocephalus brevirostris | von Sömmerring | 1816-17 1816 in Paleontology -Fossils:* Englefield publishes a general work on the geology of the Isle of Wight, reporting the discovery of large, almost certainly dinosaurian bones being discovered by geologist Thomas Webster in what would later be identified as Cretaceous strata.... |
Synonym of Pterodactylus antiquus | Based on a juvenile specimen |
| Ptenodracon brevirostris | (von Sömmerring) Lydekker | (1816-17 1816 in Paleontology -Fossils:* Englefield publishes a general work on the geology of the Isle of Wight, reporting the discovery of large, almost certainly dinosaurian bones being discovered by geologist Thomas Webster in what would later be identified as Cretaceous strata.... ) 1888 1888 in paleontology -Dinosaurs:-Newly named plesiosaurs:-New taxa:-Non-mammalian:... |
Synonym of Pterodactylus antiquus | Reclassified from Pterodactylus brevirostris, based on a juvenile specimen |
| Pterodactylus brevirostris | (von Sömmerring) Oken | (1816-17 1816 in Paleontology -Fossils:* Englefield publishes a general work on the geology of the Isle of Wight, reporting the discovery of large, almost certainly dinosaurian bones being discovered by geologist Thomas Webster in what would later be identified as Cretaceous strata.... ) 1819 |
Synonym of Pterodactylus antiquus | Name correction from Ornithocephalus brevirostris, based on a juvenile specimen |
| Pterodactylus longirostris | Cuvier | 1819 | Synonym of Pterodactylus antiquus | - |
| Macrotrachelus longirostris | (Cuvier) Giebel | (1819) 1852 1852 in paleontology -Newly named dinosaurs:-New taxa:-Paleontologists:* Gideon Algernon Mantell, the scientist who described the first known herbivorous dinosaur, dies.... |
Synonym of Pterodactylus antiquus | Reclassified from Pterodactylus longirostris |
| Ornithocephalus longirostris | (Cuvier) Ritgen | (1819) 1826 | Synonym of Pterodactylus antiquus | Incorrect name correction from Pterodactylus longirostris |
| Pterodactylus "suevicus" | Oken | 1825 1825 in paleontology -Newly named dinosaurs:Data courtesy of George Olshevsky's dinosaur genera list and Dr. Jeremy Montague's dinosaur genus database.... |
Synonym of Pterodactylus antiquus | Nomen nudum |
| Pterodactylus crocodilocephaloides | Ritgen | 1826 | Synonym of Pterodactylus antiquus | |
| Pterodactylus spectabilis | von Meyer | 1861 1861 in paleontology -Dinosaurs:-New taxa:... |
Synonym of Pterodactylus antiquus | |
| Pterodactylus grandis | Cuvier | 1824 1824 in paleontology -Dinosaurs:* William Buckland realizes that fossils he previously believed to be of cetacean origin were actually Iguanodon fossils. This mistake cost him the chance to describe the genus himself.-Newly named dinosaurs:... |
Nomen dubium, possible synonym of Rhamphorhynchus muensteri | |
| Pterodactylus longicollum | von Meyer | 1854 1854 in paleontology -Newly named dinosaurs:-Newly named plesiosaurs:-Non-mammalian:... |
Valid, may belong to a separate genus | |
| Pterodactylus micronyx | von Meyer | 1852 1852 in paleontology -Newly named dinosaurs:-New taxa:-Paleontologists:* Gideon Algernon Mantell, the scientist who described the first known herbivorous dinosaur, dies.... |
Possible synonym of Gnathosaurus subulatus Gnathosaurus Gnathosaurus is a genus of ctenochasmatid pterosaur known from a single species, G. subulatus, described in 1833. This pterosaur had an estimated wingspan of about 1.7 meters. The slender, 28-cm-long skull had up to 130 needle-like teeth arranged laterally around the spoon-shaped tip... |
|
| Pterodactylus nettecephaloides | Ritgen | 1826 | Synonym of Pterodactylus micronyx | |
| Ornithocephalus redenbacheri | Wagner | 1851 1851 in paleontology -Newly named dinosaurs:-Newly named plesiosaurs:... |
Synonym of Pterodactylus micronyx | |
| Pterodactylus redenbacheri | (Wagner) Wagner | (1851 1851 in paleontology -Newly named dinosaurs:-Newly named plesiosaurs:... ) 1861 1861 in paleontology -Dinosaurs:-New taxa:... |
Synonym of Pterodactylus micronyx | Name correction from Ornithocephalus redenbacheri |
| Pterodactylus pulchellus | von Meyer | 1861 1861 in paleontology -Dinosaurs:-New taxa:... |
Synonym of Pterodactylus micronyx | |
| Pterodactylus kochi | (Wagner) | (1837 1837 in paleontology -Newly named dinosaurs:Data courtesy of George Olshevsky's dinosaur genera list and Dr. Jeremy Montague's dinosaur genus database.... ) |
Synonym of Pterodactylus antiquus | Name correction from Ornithocephalus kochi |
| Ornithocephalus kochi | Wagner | 1837 1837 in paleontology -Newly named dinosaurs:Data courtesy of George Olshevsky's dinosaur genera list and Dr. Jeremy Montague's dinosaur genus database.... |
Synonym of Pterodactylus kochi | |
| Diopecephalus kochi | (Wagner) Seeley | (1837 1837 in paleontology -Newly named dinosaurs:Data courtesy of George Olshevsky's dinosaur genera list and Dr. Jeremy Montague's dinosaur genus database.... ) 1871 1871 in paleontology -Newly named dinosaurs:-New taxa:-New taxa:... |
Synonym of Pterodactylus kochi | Reclassification of Pterodactylus kochi |
| Pterodactylus meyeri | Muenster | 1842 1842 in paleontology -Newly named dinosaurs:... |
Synonym of Pterodactylus kochi | |
| Ornithocephalus meyeri | (Muenster) Wagner | (1842 1842 in paleontology -Newly named dinosaurs:... ) 1851 1851 in paleontology -Newly named dinosaurs:-Newly named plesiosaurs:... |
Synonym of Pterodactylus kochi | Incorrect name correction from Pterodactylus meyeri |
| Pterodactylus scolopaciceps | von Meyer | 1850 1850 in paleontology -Newly named dinosaurs:... |
Synonym of Pterodactylus kochi | |
| Rhamphorhynchus scolopaciceps | (von Meyer) | (1850 1850 in paleontology -Newly named dinosaurs:... ) |
Synonym of Pterodactylus kochi | Reclassification of Pterodactylus scolopaciceps |
| Pterodactylus cerinensis | von Meyer | 1860 1860 in paleontology -New taxa:-New taxa:-Non-mammalian:... |
Nomen dubium | |
| Pterodactylus grandipelvis | von Meyer | 1860 1860 in paleontology -New taxa:-New taxa:-Non-mammalian:... |
Nomen dubium | |
| Pterodactylus manseli | Owen | 1874 1874 in paleontology -Newly named dinosaurs:-New taxa:-Newly named pterosaurs:-new taxa:... |
Nomen dubium | |
| Pterodactylus pleydelli | Owen | 1874 1874 in paleontology -Newly named dinosaurs:-New taxa:-Newly named pterosaurs:-new taxa:... |
Nomen dubium | |
| Pterodactylus suprajurensis | Sauvage | 1873 1873 in paleontology -New taxa:... |
Nomen dubium | |
| Pterodactylus arningi | Reck | 1931 1931 in paleontology -Newly named dinosaurs:Data courtesy of George Olshevsky's dinosaur genera list and Dr. Jeremy Montague's dinosaur genus database.-Non-mammalian:... |
Nomen dubium | |
| Pterodactylus maximus | Reck | 1931 1931 in paleontology -Newly named dinosaurs:Data courtesy of George Olshevsky's dinosaur genera list and Dr. Jeremy Montague's dinosaur genus database.-Non-mammalian:... |
Nomen dubium | |
External links
- The Pterosaur Database - Upper Jurassic Several species are discussed following individual links here.