Germanodactylus

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Germanodactylus
Temporal range: Late Jurassic, 150.8–148.5  Ma
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Germanodactylus spec 01.JPG
Fossil specimen of G. cristatus
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Order: Pterosauria
Suborder: Pterodactyloidea
Family: Germanodactylidae
Genus: Germanodactylus
Yang, 1964
Type species
Pterodactylus cristatus
Wiman, 1925
Species
Synonyms

Germanodactylus ("German finger") is a genus of germanodactylid pterodactyloid pterosaur from Upper Jurassic-age rocks of Germany, including the Solnhofen Limestone. Its specimens were long thought to pertain to Pterodactylus . The head crest of Germanodactylus is a distinctive feature.

Contents

History

Counter-slab of a fossilized specimen Germanodactylus cristatus.JPG
Counter-slab of a fossilized specimen

G. cristatus is based on specimen BSP 1892.IV.1, from the Solnhofen limestone of Eichstätt, Germany. It was originally described by Plieninger in 1901 as a specimen of Pterodactylus kochi, [1] and was given its current specific name by Carl Wiman in 1925, meaning "crested" in Latin. [2] Yang Zhongjian determined that it deserved its own genus in 1964. [3] A second species called G. ramphastinus (in 1858 accidentally revised to rhamphastinus by Christian Erich Hermann von Meyer) was named as a distinct species long before G. cristatus, described by Johann Andreas Wagner in 1851 as a species of the now deprecated genus Ornithocephalus . The specific name refers to the toucan, ramphastinos in Greek. It is based on specimen BSP AS.I.745, a skeleton from the slightly younger Mörnsheimer Limestone of Daiting, Germany. [4] Peter Wellnhofer added it to Germanodactylus in 1970, [5] although Maisch and his coauthors have suggested that it deserves its own genus, "Daitingopterus". David M. Unwin has also referred miscellaneous limb bones and vertebrae from the somewhat older Kimmeridge Clay of Dorset, England to the genus; these finds at the time marked the earliest appearance of short-tailed pterosaurs in the fossil record. [6] Bennett suggested in 1996 that Germanodactylus represented adults of Pterodactylus, [7] but this has been rejected by further studies, [8] [9] [10] [11] including his own. [12] [13] Bennett's 2006 reappraisal of Germanodactylus found both species to be valid and included within the genus, with G. cristatus known from four specimens, including two juveniles, and G. rhamphastinus from two specimens. The genus differs from other pterosaurs by a combination of characteristics including a sharply pointed jaw tip, four to five premaxillary teeth, and eight to twelve maxillary teeth per side of the upper jaw, robust maxillary teeth that, unlike in Pterodactylus, are not reduced in size farther from the tip of the jaw, a naso-antorbital fenestra twice the length of the eye socket, and various proportional differences. G. cristatus differs from "G. rhampastinus" by having no teeth in the tip of the jaw and fewer teeth (~13 in each side of the upper jaw and ~12 in the lower versus 16 upper and 15 lower on each side for "G. rhamphastinus"). [13]

Description

Germanodactylus is described as being "raven-sized" in weight. [8] G. cristatus had a 0.98 meter wingspan (3.2 ft) and a 13 centimeter (5.1 in) long skull, while "G. rhamphastinus" was somewhat larger, with a 1.08 meter (3.5 ft) wingspan, and a skull measuring 21 centimeters (8.3 in) long. [14]

Head crest

Germanodactylus is known for its head crest, which had a bony portion (a low ridge running up the midline of the skull) and a soft-tissue portion that more than doubled its height. The bony part does not go as far up the head in G. cristatus as in G. rhamphastinus. The soft-tissue portion was not known early on, being first described in 2002 by S. Christopher Bennett. It was probably composed of cornified epidermis. Germanodactylus is the first genus for which a soft-tissue component of the crest is known, but similar structures were probably widespread among pterosaurs. [12] Head crests like these are now known to be far more extensive in Pterosauria. Of all the pterosaurs known to date, the most basal form with such a crest is Austriadactylus and the most derived are Hamipterus and Tapejara . Darwinopterus and Cuspicephalus also possess headcrests made of "fibrous" bone, demonstrating that the character is a homology, [15] and not a homoplasy.

Classification

This genus is unspecialized compared to the pterosaurs of the Cretaceous, and has had varying placements in Pterosauria. Yang Zhongjian, who named the genus, gave it its own family Germanodactylidae. [3] Bennett included the genus in the family Pterodactylidae, [13] and Alexander W.A. Kellner found it to be related to Pterodactylus in his 2003 phylogenetic analysis. [9] David M. Unwin, on the other hand, preferred to consider it a basal dsungaripteroid, [10] [16] a group that evolved into dedicated shellfish-eaters. [17] Maisch and co-authors considered the genus Germanodactylus to be paraphyletic, meaning the two species were not congeneric. For "G. rhamphastinus" the genus name "Daitingopterus" was used in a table, [18] but the rules of the ICZN were not observed, making the new name a nomen nudum. However, Maisch and his co-authors did place both Germanodactylus species in Dsungaripteroidea, like Unwin. [10] Vidovic and Martill not only considered the contents of Germanodactylus to be paraphyletic, but they found the two species to be entirely distinct in their cladistic analysis. G. cristatus was considered the sister taxon of both Dsungaripteroidea and Azhdarchoidea, while "G. rhamphastinus" was sister taxon to a group they called Aurorazhdarchia. Vidovic and Martill did not initially propose a new name for "G. rhamphastinus", but they suggested that it might represent an adult Diopecephalus , if that genus proves to be valid. [19] However, in a subsequent paper published in 2017, Vidovic and Martill created a new genus for "G. rhamphastinus", Altmuehlopterus . [20]

Below is a cladogram showing the results of a phylogenetic analysis presented by Longrich, Martill, and Andres in 2018. In this analysis, they placed the species G. cristatus as the sister taxon of "G. rhamphastinus" within the family Germanodactylidae, and contrary to the concepts mentioned above, both of these species were recovered as closer relatives of more primitive pterosaurs, such as Pterodactylus, within the group Archaeopterodactyloidea. [21]

Archaeopterodactyloidea
Germanodactylidae

Germanodactylus cristatus

Germanodactylus rhamphastinus Altmuehlopterus DB.jpg

Euctenochasmatia

Pterodactylus antiquus Pterodactylus BMMS7 life.png

Ctenochasmatoidea
Gallodactylidae

Cycnorhamphus suevicus

Normannognathus wellnhoferi

Ctenochasmatidae Pterodaustro BW.jpg

See also

Related Research Articles

<i>Pterodactylus</i> Genus of pterodactyloid pterosaur from the Late Jurassic

Pterodactylus is an extinct genus of pterosaurs. It is thought to contain only a single species, Pterodactylus antiquus, which was the first pterosaur to be named and identified as a flying reptile and one of the first prehistoric reptiles to ever be discovered.

<i>Rhamphorhynchus</i> Genus of rhamphorhynchid pterosaur from the Late Jurassic

Rhamphorhynchus is a genus of long-tailed pterosaurs in the Jurassic period. Less specialized than contemporary, short-tailed pterodactyloid pterosaurs such as Pterodactylus, it had a long tail, stiffened with ligaments, which ended in a characteristic soft-tissue tail vane. The mouth of Rhamphorhynchus housed needle-like teeth, which were angled forward, with a curved, sharp, beak-like tip lacking teeth, indicating a diet mainly of fish; indeed, fish and cephalopod remains are frequently found in Rhamphorhynchus abdominal contents, as well as in their coprolites.

<i>Ornithocheirus</i> Genus of ornithocheirid pterosaur from the Early Cretaceous

Ornithocheirus is a pterosaur genus known from fragmentary fossil remains uncovered from sediments in the United Kingdom and possibly Morocco.

Anhanguera is a genus of pterodactyloid pterosaur known from the Early Cretaceous Romualdo Formation of Brazil and the Late Cretaceous Kem Kem Group of Morocco. This pterosaur is closely related to Ornithocheirus, but belongs in the family Anhangueridae. The generic name comes from the Tupi words añanga, meaning "spirit protector of the animals" + wera "bygone".

<i>Ctenochasma</i> Genus of ctenochasmatid pterosaur from the Late Jurassic

Ctenochasma is a genus of Late Jurassic ctenochasmatid pterosaur belonging to the suborder Pterodactyloidea. Three species are currently recognized: C. roemeri, C. taqueti, and C. elegans. Their fossilized remains have been found in the Solnhofen Limestone of Bavaria, Germany, the "Purbeck Group" of northeastern Germany, and the Calcaires tâchetés of eastern France.

<i>Gnathosaurus</i> Genus of ctenochasmatid pterosaur from the Late Jurassic period

Gnathosaurus is a genus of ctenochasmatid pterosaur containing two species: G. subulatus, named in 1833 from the Solnhofen Limestone of Germany, and G. macrurus, known from the Purbeck Limestone of the UK. Its fossil remains dated back to the Late Jurassic period.

<span class="mw-page-title-main">Pterodactyloidea</span> Suborder of monofenestratan pterosaurs

Pterodactyloidea is one of the two traditional suborders of pterosaurs, and contains the most derived members of this group of flying reptiles. They appeared during the middle Jurassic Period, and differ from the basal rhamphorhynchoids by their short tails and long wing metacarpals. The most advanced forms also lack teeth, and by the late Cretaceous, all known pterodactyloids were toothless. Many species had well-developed crests on the skull, a form of display taken to extremes in giant-crested forms like Nyctosaurus and Tupandactylus. Pterodactyloids were the last surviving pterosaurs when the order became extinct at the end of the Cretaceous Period, together with the non-avian dinosaurs and most marine reptiles.

<span class="mw-page-title-main">Gallodactylidae</span> Family of ctenochasmatoid pterosaurs

Gallodactylidae is a group of pterosaurs within the suborder Pterodactyloidea. Gallodactylids differed from other related pterosaurs in several distinct features, including fewer than 50 teeth present only in the jaw tips, and rounded crests present on the rear portion of the skull and jaws but not near the ends of their snouts. At least some species possessed jaw flanges, possibly used to bissect hard-shelled prey.

<i>Coloborhynchus</i> Genus of anhanguerid pterosaur from the Cretaceous period

Coloborhynchus is a genus of pterodactyloid pterosaur belonging to the family Anhangueridae, though it has also been recovered as a member of the Ornithocheiridae in some studies. Coloborhynchus is known from the Lower Cretaceous of England, and depending on which species are included, possibly the Albian and Cenomanian ages as well. Coloborhynchus was once thought to be the largest known toothed pterosaur, however, a specimen of the closely related Tropeognathus is now thought to have had a larger wingspan.

<i>Cycnorhamphus</i> Genus of gallodactylid pterosaur Late Jurassic

Cycnorhamphus is a genus of gallodactylid ctenochasmatoid pterosaur from the Late Jurassic period of France and Germany, about 152 million years ago. It is synonymous with the genus Gallodactylus.

<i>Diopecephalus</i> Genus of euctenochasmatian pterosaur from the Late Jurassic

Diopecephalus is a genus of pterodactyloid pterosaur from the Lower Tithonian of the Lithographic Limestone, Bavaria, Germany. The type and only species is D. kochi, although the name has been applied to Pterodactylus longicollum, with longicollum erroneously listed as the type species.

<i>Altmuehlopterus</i> Genus of pterodactyloid pterosaur from the Late Jurassic

Altmuehlopterus is a genus of pterosaur belonging to the Pterodactyloidea. It lived in the Late Jurassic of what is now Germany. It was formerly known as "Daitingopterus", a nomen nudum, informally coined in 2004.

<span class="mw-page-title-main">Euctenochasmatia</span> Clade of archaeopterodactyloid pterosaurs

Euctenochasmatia is an extinct group of pterodactyloid pterosaurs. It was named by David Unwin in 2003 as the group that contains the most recent common ancestor of Pterodactylus and Ctenochasma, and all their descendants.

<i>Aurorazhdarcho</i> Genus of aurorazhdarchid pterosaur from the Late Jurassic

Aurorazhdarcho is an extinct genus of ctenochasmatoid pterosaur known from the Late Jurassic period of what is now Bavaria, southern Germany.

<span class="mw-page-title-main">Archaeopterodactyloidea</span> Infraorder of pterodactyloid pterosaurs

Archaeopterodactyloidea is an extinct clade of pterodactyloid pterosaurs that lived from the middle Late Jurassic to the latest Early Cretaceous periods of Africa, Asia, Europe and North America. It was named by Alexander Wilhelm Armin Kellner in 1996 as the group that contains Germanodactylus, Pterodactylus, the Ctenochasmatidae and the Gallodactylidae. In 2003, Kellner defined the clade as a node-based taxon consisting of the last common ancestor of Pterodactylus, Ctenochasma and Gallodactylus and all its descendants. Although phylogenetic analyses that based on David Unwin's 2003 analysis do not recover monophyletic Archaeopterodactyloidea, phylogenetic analyses that based on Kellner's analyses, or the analyses of Brian Andres recover monophyletic Archaeopterodactyloidea at the base of the Pterodactyloidea.

<i>Ardeadactylus</i> Genus of ctenochasmatoid pterosaur from the Late Jurassic

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<span class="mw-page-title-main">Timeline of pterosaur research</span>

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<span class="mw-page-title-main">Ornithocheiromorpha</span> Clade of pteranodontoid pterosaurs

Ornithocheiromorpha is a group of pterosaurs within the suborder Pterodactyloidea. Fossil remains of this group date back from the Early to Late Cretaceous periods, around 140 to 92.5 million years ago. Ornithocheiromorphs were discovered worldwide except Antarctica, though most genera were recovered in Europe, Asia and South America. They were the most diverse and successful pterosaurs during the Early Cretaceous, but throughout the Late Cretaceous they were replaced by better adapted and more advanced pterosaur species such the pteranodontids and azhdarchoids. The Ornithocheiromorpha was defined in 2014 by Andres and colleagues, and they made Ornithocheiromorpha the most inclusive clade containing Ornithocheirus, but not Pteranodon.

<i>Serradraco</i> Genus of pteranodontoid pterosaur from the Early Cretaceous

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References

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  12. 1 2 Bennett, S. Christopher (2002). "Soft tissue preservation of the cranial crest of the pterosaur Germanodactylus from Solnhofen". Journal of Vertebrate Paleontology. 22 (1): 43–48. doi:10.1671/0272-4634(2002)022[0043:STPOTC]2.0.CO;2. S2CID   86308635.
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