Rhamphorhynchus

Rhamphorhynchus (/ˌræmfəˈrɪŋkəs, -foʊ-/,[1] "beak snout") 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 diamond-shaped vane. The jaws 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.[2]

Although fragmentary fossil remains possibly belonging to Rhamphorhynchus have been found in England, Tanzania, and Spain, the best preserved specimens come from the Solnhofen limestone of Bavaria, Germany. Many of these fossils preserve not only the bones but impressions of soft tissues, such as wing membranes. Scattered teeth believed to belong to Rhamphorhynchus have been found in Portugal as well.[3]

Rhamphorhynchus
Temporal range: Late Jurassic, 150.8–148.5 Ma
Rhamphorhynchus munsteri
Cast of the first specimen found with wing membranes, Musée de sciences naturelles de Bruxelles
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Order: Pterosauria
Family: Rhamphorhynchidae
Subfamily: Rhamphorhynchinae
Genus: Rhamphorhynchus
Meyer, 1846
Type species
Rhamphorhynchus longicaudus
Münster, 1839
Species
  • Rhamphorhynchus muensteri
    (Goldfuss, 1831)
  • Rhamphorhynchus etchesi
    (O'Sullivan & Martill, 2015)
Synonyms
  • Ornithopterus
    Meyer, 1838b
  • Odontorhynchus
    Stolley, 1936
  • Pteromonodactylus
    Teryaev, 1967

Description

Rhamphorhynchus Scale
Size comparison
Rhamphorhynchus DB
Restoration

The largest known specimen of Rhamphorhynchus muensteri (catalog number BMNH 37002) measures 1.26 meters (4.1 ft) long with a wingspan of 1.81 meters (5.9 ft).

Skull

Contrary to a 1927 report by pterosaur researcher Ferdinand Broili, Rhamphorhynchus lacked any bony or soft tissue crest, as seen in several species of contemporary small pterodactyloid pterosaurs. Broili claimed to have found a two-millimeter-tall crest made of thin bone that ran much of the skull's length in one Rhamphorhynchus specimen, evidenced by an impression in the surrounding rock and a few small fragments of the crest itself.[4] However, subsequent examination of this specimen by Wellnhofer in 1975 and Bennett in 2002 using both visible and ultraviolet light found no trace of a crest; both concluded that Broili was mistaken. The supposed crest, they concluded, was simply an artifact of preservation.[5][6]

The teeth of Rhamphorhynchus intermesh when the jaw is closed and are suggestive of a piscivorous diet.[3] There are twenty teeth in the upper jaws and fourteen in the lower jaws.[3]

History and classification

Rhamphorhynchus reconstruction Riou 1863
One of the first ever restorations of Rhamphorhynchus from 1863, shown with tracks now known to belong to Mesolimulus

The classification and taxonomy of Rhamphorhynchus, like many pterosaur species known since the Victorian era, is complex, with a long history of reclassification under a variety of names, often for the same specimens.

The first named specimen of Rhamphorhynchus was brought to the attention of Samuel Thomas von Sömmerring by the collector Georg Graf zu Münster in 1825. Von Sömmerring concluded that it belonged to an ancient bird. When further preparation uncovered teeth, Graf zu Münster sent a cast to Professor Georg August Goldfuss, who recognised it as a pterosaur. Like most pterosaurs described in the mid 19th century, Rhamphorhynchus was originally considered to be a species of Pterodactylus. However, at the time, many scientists incorrectly considered Ornithocephalus to be the valid name for Pterodactylus. This specimen of Rhamphorhynchus was therefore originally named Ornithocephalus Münsteri. This was first mentioned in 1830 by Graf zu Münster himself.[7] However, the description making the name valid was given by Goldfuss in an 1831 follow-up to Münster's short paper.[8] Note that the ICZN later ruled that non-standard Latin characters, such as ü, would not be allowed in scientific names, and the spelling münsteri was emended to muensteri by Richard Lydekker in 1888.

In 1839, Münster described another specimen that he considered to belong to Ornithocephalus (i.e. Pterodactylus), with a distinctive long tail. He named it Ornithocephalus longicaudus, meaning "long tail", to differentiate it from the specimens with short tails (the true specimens of Pterodactylus).[9]

Ornithocephalus Münsteri
Holotype R. munsteri skull as illustrated in 1851

In 1845, Hermann von Meyer officially emended the original species Ornithocephalus münsteri to Pterodactylus münsteri, since the name Pterodactylus had been by that point recognized as having priority over Ornithocephalus.[10] In a subsequent 1846 paper describing a new species of long-tailed 'pterodactyl', von Meyer decided that the long-tailed forms of Pterodactylus were different enough from the short-tailed forms to warrant placement in a subgenus, and he named his new species Pterodactylus (Rhamphorhynchus) gemmingi after a specimen owned by collector Captain Carl Eming von Gemming that was later by von Gemming sold for three hundred guilders to the Teylers Museum in Haarlem.[11] It was not until 1847 that von Meyer elevated Rhamphorhynchus to a full-fledged genus, and officially included in it both long-tailed species of Pterodactylus known at the time, R. longicaudus (the original species preserving a long tail) and R. gemmingi.[12] The type species of Rhamphorhynchus is R. longicaudus; its type specimen or holotype also was sold to the Teylers Museum, where it still resides as TM 6924.

The original species, Pterodactylus muensteri, remained misclassified until a re-evaluation was published by Richard Owen in an 1861 book, in which he renamed it as Rhamphorhynchus münsteri.[13] The type specimen of R. muensteri, described by Münster and Goldfuss, was lost during World War II. If available, a new specimen or neotype is designated the type if the original is lost or deemed too poorly preserved. Peter Wellnhofer declined to designate a neotype in his 1975 review of the genus, because a number of high-quality casts of the original specimen were still available in museum collections.[14] These can serve as plastotypes.

By the 1990s (and following Wellnhofer's consolidation of many previously named species), about five species of Rhamphorhynchus were recognized from the Solnhofen limestone of Germany, with a few others having been named from Africa, Spain, and the UK based on fragmentary remains.[5][14] Most of the Solnhofen species were differentiated based on their relative size, and size-related features, such as the relative length of the skull.[14]

Rhamphorhynchus skull
Restored skull

In 1995, pterosaur researcher Chris Bennett published an extensive review of the currently recognized German species. Bennett concluded that all the supposedly distinct German species were actually different year-classes of a single species, R. muensteri, representing distinct age groups, with the smaller species being juveniles and the larger adults. Bennett's paper did not cover the British and African species, though he suggested that these should be considered indeterminate members of the family Rhamphorhynchidae and not necessarily species of Rhamphorhynchus itself. Despite the reduction of the genus to a single species, the type species remains R. longicaudus.[14]

The cladogram (family tree) of rhamphorhynchids below is the result of a large phylogenetic analysis published by Andres & Myers in 2013.[15]

 Breviquartossa 
 Rhamphorhynchidae 

Scaphognathus crassirostris

 Rhamphorhynchinae 

Dorygnathus banthensis

Cacibupteryx caribensis

Nesodactylus hesperius

Rhamphorhynchus muensteri

Harpactognathus gentryii

Angustinaripterus longicephalus

Sericipterus wucaiwanensis

Paleobiology

Life history

Rhamphorhynchus muensteri, Solnhofen, Germany, Late Jurassic - Royal Ontario Museum - DSC00041
Specimen with tail vane, Royal Ontario Museum

Traditionally, the large size variation between specimens of Rhamphorhynchus has been taken to represent species variation. However, in a 1995 paper, Bennett argued that these "species" actually represent year-classes of a single species, Rhamphorhynchus muensteri, from flaplings to adults. Following from this interpretation, Bennett found several notable changes that occurred in R. muensteri as the animal aged.[14]

Juvenile Rhamphorhynchus had relatively short skulls with large eyes, and the toothless beak-like tips of the jaws were shorter in juveniles than adults, with rounded, blunt lower jaw tips eventually becoming slender and pointed as the animals grew. Adult Rhamphorhynchus also developed a strong upward "hook" at the end of the lower jaw. The number of teeth remained constant from juvenile to adult, though the teeth became relatively shorter and stockier as the animals grew, possibly to accommodate larger and more powerful prey. The pelvic and pectoral girdles fused as the animals aged, with full pectoral fusion attained by one year of age.[14]

The shape of the tail vane also changed across various age classes of Rhamphorhynchus. In juveniles, the vane was shallow relative to the tail and roughly oval, or "lancet-shaped". As growth progressed, the tail vane became diamond-shaped, and finally triangular in the largest individuals.[14]

Rhamphorhynchus intermedius
R. intermedius, a small specimen that might be a juvenile of R. muensteri

The smallest known Rhamphorhynchus specimen has a wingspan of only 290 millimeters; however, it is likely that even such a small individual was capable of flight. Bennett examined two possibilities for hatchlings: that they were altricial, requiring some period of parental care before leaving the nest, or that they were precocial, hatching with sufficient size and ability for flight. If precocious, Bennett suggested that clutches would be small, with only one or two eggs laid per clutch, to compensate for the relatively large size of the hatchings. Bennett did not speculate on which possibility was more likely, though the discovery of a pterosaur embryo (Avgodectes) with strongly ossified bones suggests that pterosaurs in general were precocial, able to fly soon after hatching with minimal parental care.[16] This theory was contested by a histological study of Rhamphorhynchus that showed the initial rapid growth was followed by a prolonged period of slow growth.[17]

Metabolism

Cast of Rhamphorhynchus muensteri 02 - Pterosaurs Flight in the Age of Dinosaurs
Cast of the "dark wing" specimen

Having determined that Rhamphorhynchus specimens fit into discrete year-classes, Bennett was able to estimate the growth rate during one year by comparing the size of one-year-old specimens with two-year-old specimens. He found that the average growth rate during the first year of life for Rhamphorhynchus was 130% to 173%, slightly faster than the growth rate in alligators. Growth likely slowed considerably after sexual maturity, so it would have taken more than three years to attain maximum adult size.[14]

This growth rate is much slower than the rate seen in large pterodactyloid pterosaurs, such as Pteranodon, which attained near-adult size within the first year of life. Additionally, pterodactyloids had determinate growth, meaning that the animals reached a fixed maximum adult size and stopped growing. Previous assumptions of rapid growth rate in rhamphorhynchoids were based on the assumption that they needed to be warm-blooded to sustain active flight. Warm-blooded animals, like modern birds and bats, normally show rapid growth to adult size and determinate growth. Because there is no evidence for either in Rhamphorhynchus, Bennett considered his findings consistent with an ectothermic metabolism, though he recommended more studies needed to be done. Cold-blooded Rhamphorhynchus, Bennett suggested, may have basked in the sun or worked their muscles to accumulate enough energy for bouts of flight, and cooled to ambient temperature when not active to save energy, like modern reptiles.[14]

Swimming

Rhamphorhynchus life restoration
Restoration showing an individual by the sea

Though Rhamphorhynchus is often depicted as an aerial piscivore, recent evidence suggests that, much like most modern aquatic birds, it probably foraged while swimming. Like several pteranodontians it has hatchet-shaped deltopectoral crests, a short torso and short legs, all features associated with water based launching in pterosaurs. Its feet are broad and large, being useful for propulsion, and the predicted floating position is adequate by pterosaur standards.[18]

The animal's ability to swim may account for the genus' generally excellent fossil record, being in a position where preservation would be much easier.

Sexual dimorphism

Both Koh Ting-Pong and Peter Wellnhofer recognized two distinct groups among adult Rhamphorhynchus muensteri, differentiated by the proportions of the neck, wing, and hind limbs, but particularly in the ratio of skull to humerus length. Both researchers noted that these two groups of specimens were found in roughly a 1:1 ratio, and interpreted them as different sexes.[5][19] Bennett tested for sexual dimorphism in Rhamphorhynchus by using a statistical analysis, and found that the specimens did indeed group together into small-headed and large-headed sets. However, without any known variation in the actual form of the bones or soft tissue (morphological differences), he found the case for sexual dimorphism inconclusive.[14]

Head orientation

Rhamphorynchus gemmingi jconway
Depiction by John Conway

In 2003, a team of researchers led by Lawrence Witmer studied the brain anatomy of several types of pterosaurs, including Rhamphorhynchus muensteri, using endocasts of the brain they retrieved by performing CAT scans of fossil skulls. Using comparisons to modern animals, they were able to estimate various physical attributes of pterosaurs, including relative head orientation during flight and coordination of the wing membrane muscles. Witmer and his team found that Rhamphorhynchus held its head parallel to the ground due to the orientation of the osseous labyrinth of the inner ear, which helps animals detect balance. In contrast, pterodactyloid pterosaurs, such as Anhanguera, appear to have normally held their heads at a downward angle, both in flight and while on the ground.[20]

Daily activity patterns

Comparisons between the scleral rings of Rhamphorhynchus and modern birds and reptiles suggest that it may have been nocturnal, and may have had activity patterns similar to those of modern nocturnal seabirds. This may also indicate niche partitioning with contemporary pterosaurs inferred to be diurnal, such as Scaphognathus and Pterodactylus.[21]

Ecology

Rhamphorhynchus and Aspidorhynchus
Fossil specimen WDC CSG 255, including a Rhamphorhynchus with a Leptolepides fish trapped in the pharynx and caught in the jaws of an Aspidorhynchus

Several limestone slabs have been discovered in which fossils of Rhamphorhynchus are found in close association with the ganoid fish Aspidorhynchus. In one of these specimens, the jaws of an Aspidorhynchus pass through the wings of the Rhamphorhynchus specimen. The Rhamphorhynchus also has the remains of a small fish, possibly Leptolepides, in its throat. This slab, cataloged as WDC CSG 255, may represent two levels of predation; one by Rhamphorhynchus and one by Aspidorhynchus. In a 2012 description of WDC CSG 255, researchers proposed that the Rhamphorhynchus individual had just caught a Leptolepides while it was swimming. As the Leptolepides was travelling down its pharynx, a large Aspidorhynchus would have attacked from below the water, accidentally puncturing the left wing membrane of the Rhamphorhynchus with its sharp rostrum in the process. The teeth in its snout were ensnared in the fibrous tissue of the wing membrane, and as the fish thrashed to release itself the left wing of Rhamphorhynchus was pulled backward into the distorted position seen in the fossil. The encounter resulted in the death of both individuals, most likely because the two animals sank into an anoxic layer in the water body, depriving the fish of oxygen. The two may have been preserved together as the weight of the head of Aspidorhynchus held down the much lighter body of Rhamphorhynchus.[22]

Species

Ramphorhynchus sp., view 1, Late Late Jurassic, Tithonian Age, Solnhofen Lithographic Limestone, Eichstatt, Bavaria, Germany - Houston Museum of Natural Science - DSC01862
Rhamphorhynchus sp., Houston Museum of Natural Science
Rhamphorhynchus gemmingi holotype
Holotype of R. gemmingi, Teylers Museum, Haarlem
Rhamphorhynchus Lauer
R. muensteri, Lauer Foundation for Paleontology
Rhamphorhynchus gemmingi pterosaur
Rhamphorhynchus gemmingi from the travelling exhibit "Paleomania"[23]

A large number of Rhamphorhynchus species have been named, but are currently considered year-classes of R. muensteri by most researchers. Currently recognized specimens of Rhamphorhynchus have previously been published on under the following junior synonyms:[24]

Note that Rhamphorhynchus is also a genus of orchid, named in 1977 by botanist L.A. Garay. In biological nomenclature, the same name may be used for an animal that has already been used for a plant or vice versa.

Synonyms

Synonyms of Rhamphorhynchus muensteri:

  • R. longicaudus (Münster, 1839) von Meyer, 1846
    • Pterodactylus longicaudus Münster, 1839
  • R. gemmingi (von Meyer, 1846) von Meyer, 1855
    • Pterodactylus lavateri von Meyer, 1838a
    • Ornithopterus lavateri (von Meyer, 1838a) von Meyer, 1838b
    • Pterodactylus gemmingi von Meyer, 1846
    • Pterodactylus (Rhamphorhynchus) gemmingi von Meyer, 1846
    • Rhamphorhynchus (Pterodactylus) gemmingi (von Meyer, 1846) von Meyer, 1855
    • R. suevicus O. Fraas, 1855
    • Pterodactylus hirundinaceus Wagner, 1857
    • R. curtimanus Wagner, 1858
    • R. longimanus Wagner, 1858
    • R. meyeri Owen, 1870
    • R. phyllurus Marsh, 1882
    • Pteromonodactylus phyllurus (Marsh, 1882) Teryaev, 1967
  • R. longiceps Woodward, 1902
    • R. kokeni F. Plieninger, 1907
    • R. megadactylus von Koenigswald, 1931
    • R. carnegiei Koh, 1937
  • Rhamphorhynchus intermedius Koh, 1937
    • R. intermedius var. brevialata Koh, 1937

Dubious species

Dubious species of Rhamphorhynchus:

  • R. jessoni (Lydekker, 1890)
  • R. tendagurensis (Reck, 1931)

"Odontorhynchus"

"Odontorhynchus" aculeatus was based on a skull with lower jaws that is now lost. This set of jaws supposedly differed in having two teeth united at the tip of the lower jaw, and none at the tip of the upper jaw. The skull was 6.5–7.0 cm (2.6–2.8 in), making it a small form.[25] Stolley, who described the specimen in 1936, argued that R. longicaudus also should be reclassified in the genus "Odontorhynchus". Both Koh and Wellnhofer rejected this idea, arguing instead that "Odontorhynchus" was a junior synonym of R. longicaudus.[5][19] Bennett agreed with their assessments, and included both "Odontorhynchus" and R. longicaudus as synonyms of R. muensteri.[14]

See also

References

  1. ^ "Rhamphorhynchus". Oxford Dictionaries. Oxford University Press. Retrieved 2016-01-22.
  2. ^ Frey, E.; Tischlinger, H. (2012). "The Late Jurassic pterosaur Rhamphorhynchus, a frequent victim of the ganoid fish Aspidorhynchus?". PLOS ONE. 7 (3): e31945. doi:10.1371/journal.pone.0031945. PMC 3296705. PMID 22412850.
  3. ^ a b c "Rhamphorhynchus." In: Cranfield, Ingrid (ed.). The Illustrated Directory of Dinosaurs and Other Prehistoric Creatures. London: Salamander Books, Ltd. Pp. 302-305.
  4. ^ Broili, F. (1927). "Ein Exemplar von Rhamphorhynchus mit Resten von Schwimmhaut". Sitzungs-Berichte der bayerischen Akademie der Wissenschaften mathematisch naturwissenschaftlichen Abteilung. 1927: 29–48.
  5. ^ a b c d Wellnhofer, P. (1975). "Die Rhamphorhynchoidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands". Palaeontographica A. 148: 1–33., 148: 132-186, 149: 1-30.
  6. ^ Bennett, S.C. (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.
  7. ^ Münster, G. Graf zu. (1830). "Nachtrag zu der Abhandlung des Professor Goldfuss über den Ornithocephalus Münsteri (Goldf.)." Bayreuth, 8 p.
  8. ^ Goldfuss, G.A. (1831). "Beiträge zur Kenntnis verschiedener Reptilien der Vorwelt". Nova Acta Academiae Caesareae Leopoldino-Carolinae Germanicae Naturae Curiosorum. 15: 61–128.
  9. ^ Münster, G.G. (1839). "Ueber einige neue Versteinerungen in der lithographischen Schiefer von Baiern". Neues Jahrbuch für Mineralogie, Geologie, und Palaeontologie. 1839: 676–682.
  10. ^ Meyer, H. von. (1845). "System der fossilen Saurier [Taxonomy of fossil saurians]". Neues Jahrbuch für Mineralogie, Geognosie, Geologie und Petrefakten-Kunde. 1845: 278–285.
  11. ^ Meyer, H. von. (1846). "Pterodactylus (Rhamphorhynchus) gemmingi aus dem Kalkschiefer von Solenhofen". Palaeontographica. 1: 1–20.
  12. ^ Meyer, H. von. (1847). "Homeosaurus maximiliani und Rhamphorhynchus (Pterodactylus) longicaudus, zwei fossile Reptilien aus der Kalkschiefer von Solenhofen." 4X, Frankfurt, 22 p.
  13. ^ Owen, R. (1861). Palaeontology, or a Systematic Summary of Extinct Animals and their Geological Relations. Adam and Charles Black, Edinburgh, 1-463.
  14. ^ a b c d e f g h i j k Bennett, S. C. (1995). "A statistical study of Rhamphorhynchus from the Solnhofen Limestone of Germany: Year-classes of a single large species". Journal of Paleontology. 69: 569–580.
  15. ^ Andres, B.; Myers, T. S. (2013). "Lone Star Pterosaurs". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 103 (3–4): 1. doi:10.1017/S1755691013000303.
  16. ^ Wang, X.; Zhou, Z. (2004). "Pterosaur embryo from the Early Cretaceous". Nature. 429 (6992): 623. doi:10.1038/429621a. PMID 15190343.
  17. ^ Prondvai, E.; Stein, K.; Ősi, A.; Sander, M. P. (2012). Soares, Daphne (ed.). "Life history of Rhamphorhynchus inferred from bone histology and the diversity of pterosaurian growth strategies". PLoS ONE. 7 (2): e31392. doi:10.1371/journal.pone.0031392. PMC 3280310. PMID 22355361.
  18. ^ Witton, M. P. (2015). "Were early pterosaurs inept terrestrial locomotors?". PeerJ. 3: e1018. doi:10.7717/peerj.1018. PMC 4476129. PMID 26157605.
  19. ^ a b Koh (1937). "Untersuchungen über die Gattung Rhamphorhynchus". Neues Jahrbuch für Mineralogie, Geologie und Palaeontologie, Beilage-Band. 77: 455–506.
  20. ^ Witmer, L.M., S. Chatterjee, J. Franzosa, T. Rowe, and R. C. Ridgely. (2004). "Neuroanatomy and vestibular apparatus of pterosaurs: Implications for flight, posture, and behavior." Annual Meeting of the Society of Integrative and Comparative Biology, New Orleans, LA. Integrative and Comparative Biology, 43(6): 832. [1]
  21. ^ Schmitz, L.; Motani, R. (2011). "Nocturnality in Dinosaurs Inferred from Scleral Ring and Orbit Morphology". Science. 332 (6030): 705–8. doi:10.1126/science.1200043. PMID 21493820.
  22. ^ Frey, E.; Tischlinger, H. (2012). "The Late Jurassic pterosaur Rhamphorhynchus, a frequent victim of the ganoid fish Aspidorhynchus?". PLoS ONE. 7 (3): e31945. doi:10.1371/journal.pone.0031945. PMC 3296705. PMID 22412850.
  23. ^ Paleomania , International Museum Institute
  24. ^ Olshevsky, G. (2001), "Re: Pterosaur Help", discussion group, The Dinosaur Mailing List, viewed January 10, 2009. http://dml.cmnh.org/.
  25. ^ Stolley, E. (1936). "Odontorhynchus aculeatus novo. gen. novo. sp., Ein neuer Rhamphorhynchide von Solnhofen. Neues Jahrbuch für Mineralogie, Geololgie, und Paläontologie". Beilage-Band. 75: 543–564.
Aerodactylus

Aerodactylus (meaning "wind finger") is a dubious pterosaur genus containing a single species, Aerodactylus scolopaciceps, previously regarded as a species of Pterodactylus.

The fossil remains of this species have been found only in the Solnhofen limestone of Bavaria, Germany, dated to the late Jurassic Period (early Tithonian), about 150.8–148.5 million years ago. Like all pterosaurs, the wings of Aerodactylus 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 keratinous ridges. Several well preserved fossils have shown that Aerodactylus was covered in a short, dense coat of bristly pycnofibres, and that it had a rounded triangular crest on its head, as well as a backward-pointing lappet. It is named after the Pokémon Aerodactyl. The validity of Aerodactylus has been disputed, with some pterosaur experts suggesting that none of the specimens referred to this genus are distinguishable from Pterodactylus.

Allkaruen

Allkaruen (meaning "ancient brain") is a genus of rhamphorhynchoid pterosaur from the Early-to-Middle Jurassic Cañadon Asfalto Formation in Argentina. It contains a single species, A. koi.

Aspidogyne mendoncae

Aspidogyne mendoncae is a species of orchid that grows in Brazil.

Aspidorhynchus

Aspidorhynchus (meaning "shield snout") is an extinct genus of ray-finned fish from the Jurassic and Cretaceous periods. Fossils have been found in Europe and Antarctica.

Aspidorhynchus was a slender, fast-swimming fish, 60 centimetres (2.0 ft) long, with tooth-lined, elongated jaws. It also had heavy scales and a symmetrical tail. The upper jaw was longer than the lower jaw, ending in a toothless spike. Although it would have looked superficially similar to the present day gar, its closest living relative is actually the bowfin.

Bellubrunnus

Bellubrunnus (meaning "the beautiful one of Brunn" in Latin) is an extinct genus of rhamphorhynchid pterosaur from the Late Jurassic (Kimmeridgian stage) of southern Germany. It contains a single species, Bellubrunnus rothgaengeri. Bellubrunnus is distinguished from other rhamphorhynchids by its lack of long projections on the vertebrae of the tail, fewer teeth in the jaws, and wingtips that curve forward rather than sweep backward as in other pterosaurs.

Cacibupteryx

Cacibupteryx is a genus of rhamphorhynchid "rhamphorhynchoid" pterosaur from the middle-late Oxfordian-age Upper Jurassic Jagua Formation of Pinar del Río, Cuba.

The genus was named in 2004 by Zulma Gasparini, Marta Fernández and Marcelo de la Fuente. The type species is Cacibupteryx caribensis. The genus name is derived from Cacibu, the "Lord of the Sky" in Taíno and Greek pteryx, "wing". The specific name refers to the Caribbean, Caribe in Spanish.

The genus is based on holotype IGO-V 208, a partial but well-preserved uncrushed skull — missing the tip of the snout, teeth, and lower jaws — and fragmentary left wing: the distal end of the ulna, fragments of the radius, and the first and fourth phalanx of the wing finger. The partial skull is seventeen centimeters long (6.7 inches), is preserved in three dimensions, and has a broad roof. The wingspan and length were not estimated by the describers, but it was indicated to be a relatively large form.The describers assigned Cacibupteryx to the Rhamphorhynchidae. Although there is little overlapping material with contemporaneous Nesodactylus from the same location, the two are clearly different as proven by details of the elbow and quadrate. Cacibupteryx is one of the most complete Oxfordian pterosaurs, and demonstrates additional Oxfordian pterosaur diversity. Phylogenetic analyses have found it to be a member of the subfamily Rhamphorhynchinae, closely related to Rhamphorhynchus and Nesodactylus.

Campylognathoides

Campylognathoides ("curved jaw", Strand 1928) is a genus of pterosaur, discovered in the Württemberg Lias deposits (dated to the early Toarcian age) of Germany; this first specimen consisted however only of wing fragments. Further better preserved specimens were found in the Holzmaden shale: basing on these specimens Felix Plieninger erected a new genus.

Dorygnathus

Dorygnathus ("spear jaw") was a genus of pterosaur that lived in Europe during the Early Jurassic period, 180 million years ago when shallow seas flooded much of the continent. It had a short 1.5 meters (4.9 feet) wingspan, and a relatively small triangular sternum, which is where its flight muscles attached. Its skull was long and its eye sockets were the largest opening therein. Large curved fangs that "intermeshed" when the jaws closed featured prominently at the front of the snout while smaller, straighter teeth lined the back. Having variable teeth, a condition called heterodonty, is rare in modern reptiles but more common in primitive pterosaurs. The heterodont dentition in Dorygnathus is consistent with a piscivorous (fish-eating) diet. The fifth digit on the hindlimbs of Dorygnathus was unusually long and oriented to the side. Its function is not certain, but the toe may have supported a membrane like those supported by its wing-fingers and pteroids. Dorygnathus was according to David Unwin related to the Late Jurassic pterosaur, Rhamphorhynchus and was a contemporary of Campylognathoides in Holzmaden and Ohmden.

Eudimorphodon

Eudimorphodon was a pterosaur that was discovered in 1973 by Mario Pandolfi in the town of Cene, Italy and described the same year by Rocco Zambelli. The nearly complete skeleton was retrieved from shale deposited during the Late Triassic (mid to late Norian stage), making Eudimorphodon one of the oldest pterosaurs known. It had a wingspan of about 100 centimetres (3.3 ft) and at the end of its long bony tail may have been a diamond-shaped flap like in the later Rhamphorhynchus. If so, the flap may have helped it steer while maneuvering in the air. Eudimorphodon is known from several skeletons, including juvenile specimens.

Klobiodon

Klobiodon is a genus of rhamphorhynchid pterosaur from the Middle Jurassic Taynton Limestone Formation of Oxfordshire, England.

Kongamato

The kongamato ("breaker of boats") is a reported pterosaur-like creature said to have been seen by the people of and explorers in the Mwinilunga district's Jiundu swamps of Western Zambia, Angola and Congo. Suggested identities include a modern-day Rhamphorhynchus, a misidentified bird (such as the very large and peculiar saddle-billed stork), or a giant bat. No film has ever been taken, nor have any bodies been examined, leaving all of the stories to rely on large wounds and eyewitness accounts.

Nesodactylus

Nesodactylus was a genus of "rhamphorhynchoid" pterosaur from the middle-late Oxfordian age Upper Jurassic Jagua Formation of Pinar del Río, western Cuba.

Its remains were collected but not prepared by Barnum Brown in 1918, from rocks better known for their fossils of marine life. When seven black chalkstone blocks were prepared from 1966 by Richard Lund by dissolving the substrate in acid, this revealed the remains of a pterosaur.

Ned Colbert described and named the genus in 1969. The type species is Nesodactylus hesperius. The genus name is derived from Greek nesos, "island" and daktylos, "finger", a reference to the island of Cuba and the typical wing finger of pterosaurs. The specific name means "western", from Greek hesperios.

The genus is based on holotype AMNH 2000, a partial skeleton including a skull fragment, numerous vertebrae from all parts of the spine and tail, zygapophyses (interpreted by Colbert as ossified tendons) on the tail, the pectoral girdle and a very deeply keeled sternum, arms and partial hands, part of the pelvis, parts of both femorae, partial metatarsals, and ribs. The specimen was disarticulated but associated and not very compressed; during the preparation from the limestone with acid, the bones were not completely removed.

Colbert found Nesodactylus to have had longer wings and more robust limbs and longer legs than related Rhamphorhynchus, although of a similar size and overall anatomy. He classified it as a rhamphorhynchid and more precisely as a member of the Rhamphorhynchinae.In 1977 James A. Jensen and John Ostrom by mistake referred to it as Nesodon (1977). Although there is little overlapping material with contemporaneous Cacibupteryx, the two are clearly different based on details of the elbow and quadrate. At least one recent review suggests it was a rhamphorhynchine, while another does not classify it.

Novialoidea

Novialoidea (meaning "new wings") is an extinct clade of macronychopteran pterosaurs that lived from the latest Early Jurassic to the latest Late Cretaceous (early Toarcian to late Maastrichtian age), their fossils having been found on all continents except Antarctica. It was named by Alexander Wilhelm Armin Kellner in 2003 as a node-based taxon consisting of the last common ancestor of Campylognathoides, Quetzalcoatlus and all its descendants. This name was derived from Latin novus "new", and ala, "wing", in reference to the wing synapomorphies that the members of the clade possess. Unwin (2003) named Lonchognatha in the same issue of the journal that published Novialoidea (Geological Society of London, Special Publications 217) and defined it as Eudimorphodon ranzii, Rhamphorhynchus muensteri, their most recent common ancestor and all its descendants (as a node-based taxon). Under Unwin's and Kellner's phylogenetic analyses (where Eudimorphodon and Campylognathoides form a family that basal to both Rhamphorhynchus and Quetzalcoatlus), and because Novialoidea was named first (in pages 105-137, while Lonchognatha was named in pages 139-190), Lonchognatha is an objective junior synonym of the former. However, other analyses find Lonchognatha to be valid (Andres et al., 2010) or synonymous with the Pterosauria (Andres, 2010 and Andres, in press).

Ostromia

Ostromia is a genus of anchiornithid theropod dinosaur from the Late Jurassic Painten Formation of Germany. The genus contains a single species, O. crassipes, named by Christian Foth and Oliver Rauhut in 2017.

Pterodactylus giganteus

Pterodactylus giganteus is a scientific name which has been used for at least three distinct species of pterosaurs previously classified in the genus Pterodactylus. It may refer to:

"Pterodactylus" giganteus (Oken, 1819): Originally Ornithocephalus giganteus, currently considered a synonym of Rhamphorhynchus muensteri

"Pterodactylus" giganteus (Morris, 1854): A synonym of Cimoliornis diomedeus

"Pterodactylus" giganteus (Bowerbank, 1846): Now Lonchodraco giganteus

Pterosaur

Pterosaurs (; from Greek, meaning "winged lizard") were flying reptiles of the extinct clade or order Pterosauria. They existed during most of the Mesozoic: from the late Triassic to the end of the Cretaceous (228 to 66 million years ago). Pterosaurs are the earliest vertebrates known to have evolved powered flight. Their wings were formed by a membrane of skin, muscle, and other tissues stretching from the ankles to a dramatically lengthened fourth finger.Early species had long, fully toothed jaws and long tails, while later forms had a highly reduced tail, and some lacked teeth. Many sported furry coats made up of hair-like filaments known as pycnofibers, which covered their bodies and parts of their wings. Pterosaurs spanned a wide range of adult sizes, from the very small anurognathids to the largest known flying creatures of all time, including Quetzalcoatlus and Hatzegopteryx.Pterosaurs are often referred to in the popular media and by the general public as "flying dinosaurs", but the term "dinosaur" is restricted to just those reptiles descended from the last common ancestor of the groups Saurischia and Ornithischia (clade Dinosauria, which includes birds), and current scientific consensus is that this group excludes the pterosaurs, as well as the various groups of extinct marine reptiles, such as ichthyosaurs, plesiosaurs, and mosasaurs.Unlike these other reptiles, pterosaurs are nonetheless more closely related to birds and dinosaurs than to crocodiles or any other living reptile. Pterosaurs are also colloquially referred to as pterodactyls, particularly in fiction and by journalists. However, technically, pterodactyl only refers to members of the genus Pterodactylus, and more broadly to members of the suborder Pterodactyloidea of the pterosaurs.

Rhamphorhynchidae

Rhamphorhynchidae is a group of early "rhamphorhynchoid" pterosaurs named after Rhamphorhynchus, that lived in the Late Jurassic. The family Rhamphorhynchidae was named in 1870 by Harry Govier Seeley.

Rhamphorhynchus (disambiguation)

Rhamphorhynchus may refer to:

Rhamphorhynchus, a genus of pterosaur

Rhamphorhynchus, a former monotypic genus of orchid, containing only the species now called Aspidogyne mendoncae

Scaphognathus

Scaphognathus was a pterosaur that lived around Germany during the Late Jurassic. It had a wingspan of 0.9 m (3 ft).

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