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Tanystropheus
Tanystropheus (Ancient Greek: τανυ~ 'long' + στροφευς 'hinged') is an extinct genus of archosauromorph reptile which lived during the Triassic Period in Europe, Asia, and North America. It is recognisable by its extremely elongated neck, longer than the torso and tail combined. The neck was composed of 13 vertebrae strengthened by extensive cervical ribs. Tanystropheus is one of the most well-described non-archosauriform archosauromorphs, known from numerous fossils, including nearly complete skeletons. Some species within the genus may have reached a total length of 6 meters (20 ft), making Tanystropheus the longest non-archosauriform archosauromorph as well. Tanystropheus is the namesake of the family Tanystropheidae, a clade collecting many long-necked Triassic archosauromorphs previously described as "protorosaurs" or "prolacertiforms".
Tanystropheus contains at least two valid species as well as fossils which cannot be referred to a specific species. The type species of Tanystropheus is T. conspicuus, a dubious name applied to particularly large fossils from Germany and Poland. Complete skeletons are common in the Besano Formation at Monte San Giorgio, on the border of Italy and Switzerland. Monte San Giorgio fossils belong to two species: the smaller T. longobardicus and the larger T. hydroides. These two species were formally differentiated in 2020 primarily on the basis of their strongly divergent skull anatomy. When T. longobardicus was first described in 1886, it was initially mistaken for a pterosaur and given the name "Tribelesodon". Starting in the 1920s, systematic excavations at Monte San Giorgio unearthed many more Tanystropheus fossils, revealing that the putative wing bones of "Tribelesodon" were actually neck vertebrae.
Most Tanystropheus fossils hail from marine or coastal deposits of the Middle Triassic epoch (Anisian and Ladinian stages), with some exceptions. For example, a vertebra from Nova Scotia was recovered from primarily freshwater sediments. The youngest unambiguous fossils in the genus are a pair of well-preserved skeletons from the Zhuganpo Formation, a geological unit in China which dates to the earliest part of the Late Triassic (early Carnian stage). The oldest putative fossils belong to "T. antiquus", a European species from the latest part of the Early Triassic (late Olenekian stage). T. antiquus had a proportionally shorter neck than other Tanystropheus species, so some paleontologists consider that T. antiquus deserves a separate genus, Protanystropheus. Some studies have suggested that T. antiquus is not even part of the family Tanystropheidae, instead being closer to another long-necked early archosauromorph, Dinocephalosaurus.
The lifestyle of Tanystropheus has been the subject of much debate. Tanystropheus is unknown from drier environments and its neck is rather stiff and ungainly, suggesting a reliance on water. Conversely, the limbs and tail lack most adaptations for swimming and closely resemble their equivalents in terrestrial reptiles. Recent studies have supported an intermediate position, reconstructing Tanystropheus as an animal equally capable on land and in the water. Despite its length, the neck was lightweight and stabilized by tendons, so it would not have been a fatal hindrance to terrestrial locomotion. The hindlimbs and the base of the tail were large and muscular, capable of short bursts of active swimming in shallow water. Tanystropheus was most likely a piscivorous ambush predator: the narrow subtriangular skull of T. longobardicus is supplied with three-cusped teeth suited for holding onto slippery prey, while the broader skull of T. hydroides bears an interlocking set of large curved fangs similar to the fully aquatic plesiosaurs.
19th century excavations at Monte San Giorgio, a UNESCO world heritage site on the Italy-Switzerland border, revealed a fragmentary fossil of an animal with three-cusped (tricuspid) teeth and elongated bones. Monte San Giorgio preserves the Besano Formation (also known as the Grenzbitumenzone), a late Anisian-early Ladinian lagerstätte recognised for its spectacular fossils. In 1886, Francesco Bassani interpreted the unusual tricuspid fossil as a pterosaur, which he named Tribelesodon longobardicus. The holotype specimen of Tribelesodon longobardicus was stored in the Museo Civico di Storia Naturale di Milano (Natural History Museum of Milan), and was destroyed by allied bombing of Milan in World War II.
Excavations by University of Zürich paleontologist Bernhard Peyer in the late 1920s and 1930s revealed many more complete fossils of the species from Monte San Giorgio. Peyer's discoveries allowed Tribelesodon longobardicus to be recognised as a non-flying reptile, more than 40 years after its original description. Its supposed elongated finger bones were recognized as neck vertebrae, which compared favorably with those previously described as Tanystropheus from Germany and Poland. Thus, Tribelesodon longobardicus was renamed to Tanystropheus longobardicus and its anatomy was revised into a long-necked, non-pterosaur reptile. Specimen PIMUZ T 2791, which was discovered in 1929, has been designated as the neotype of the species.
Well-preserved T. longobardicus fossils continue to be recovered from Monte San Giorgio up to the present day. Fossils from the mountain are primarily stored at the rebuilt Museo Civico di Storia Naturale di Milano (MSNM), the Paleontological Museum of Zürich (PIMUZ), and the Museo Cantonale di Scienze Naturali di Lugano (MCSN). Rupert Wild reviewed and redescribed all specimens known at the time via several large monographs in 1973/4 and 1980. In 2005, Silvio Renesto described a T. longobardicus specimen from Switzerland which preserved the impressions of skin and other soft tissue. Five new MSNM specimens of T. longobardicus were described by Stefania Nosotti in 2007, allowing for a more comprehensive view of the species' anatomy.
A small but well-preserved skull and neck, specimen PIMUZ T 3901, was found in the slightly younger Meride Limestone at Monte San Giorgio. Wild (1980) gave it a new species, T. meridensis, based on a set of skull and vertebral traits proposed to differ from T. longobardicus. Later reinvestigations failed to confirm the validity of these differences, rendering T. meridensis a junior synonym of T. longobardicus. A 2019 revision of Tanystropheus concluded that T. longobardicus and T. antiquus were the only valid species in the genus, though this conclusion would prove to be short-lived.
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Tanystropheus
Tanystropheus (Ancient Greek: τανυ~ 'long' + στροφευς 'hinged') is an extinct genus of archosauromorph reptile which lived during the Triassic Period in Europe, Asia, and North America. It is recognisable by its extremely elongated neck, longer than the torso and tail combined. The neck was composed of 13 vertebrae strengthened by extensive cervical ribs. Tanystropheus is one of the most well-described non-archosauriform archosauromorphs, known from numerous fossils, including nearly complete skeletons. Some species within the genus may have reached a total length of 6 meters (20 ft), making Tanystropheus the longest non-archosauriform archosauromorph as well. Tanystropheus is the namesake of the family Tanystropheidae, a clade collecting many long-necked Triassic archosauromorphs previously described as "protorosaurs" or "prolacertiforms".
Tanystropheus contains at least two valid species as well as fossils which cannot be referred to a specific species. The type species of Tanystropheus is T. conspicuus, a dubious name applied to particularly large fossils from Germany and Poland. Complete skeletons are common in the Besano Formation at Monte San Giorgio, on the border of Italy and Switzerland. Monte San Giorgio fossils belong to two species: the smaller T. longobardicus and the larger T. hydroides. These two species were formally differentiated in 2020 primarily on the basis of their strongly divergent skull anatomy. When T. longobardicus was first described in 1886, it was initially mistaken for a pterosaur and given the name "Tribelesodon". Starting in the 1920s, systematic excavations at Monte San Giorgio unearthed many more Tanystropheus fossils, revealing that the putative wing bones of "Tribelesodon" were actually neck vertebrae.
Most Tanystropheus fossils hail from marine or coastal deposits of the Middle Triassic epoch (Anisian and Ladinian stages), with some exceptions. For example, a vertebra from Nova Scotia was recovered from primarily freshwater sediments. The youngest unambiguous fossils in the genus are a pair of well-preserved skeletons from the Zhuganpo Formation, a geological unit in China which dates to the earliest part of the Late Triassic (early Carnian stage). The oldest putative fossils belong to "T. antiquus", a European species from the latest part of the Early Triassic (late Olenekian stage). T. antiquus had a proportionally shorter neck than other Tanystropheus species, so some paleontologists consider that T. antiquus deserves a separate genus, Protanystropheus. Some studies have suggested that T. antiquus is not even part of the family Tanystropheidae, instead being closer to another long-necked early archosauromorph, Dinocephalosaurus.
The lifestyle of Tanystropheus has been the subject of much debate. Tanystropheus is unknown from drier environments and its neck is rather stiff and ungainly, suggesting a reliance on water. Conversely, the limbs and tail lack most adaptations for swimming and closely resemble their equivalents in terrestrial reptiles. Recent studies have supported an intermediate position, reconstructing Tanystropheus as an animal equally capable on land and in the water. Despite its length, the neck was lightweight and stabilized by tendons, so it would not have been a fatal hindrance to terrestrial locomotion. The hindlimbs and the base of the tail were large and muscular, capable of short bursts of active swimming in shallow water. Tanystropheus was most likely a piscivorous ambush predator: the narrow subtriangular skull of T. longobardicus is supplied with three-cusped teeth suited for holding onto slippery prey, while the broader skull of T. hydroides bears an interlocking set of large curved fangs similar to the fully aquatic plesiosaurs.
19th century excavations at Monte San Giorgio, a UNESCO world heritage site on the Italy-Switzerland border, revealed a fragmentary fossil of an animal with three-cusped (tricuspid) teeth and elongated bones. Monte San Giorgio preserves the Besano Formation (also known as the Grenzbitumenzone), a late Anisian-early Ladinian lagerstätte recognised for its spectacular fossils. In 1886, Francesco Bassani interpreted the unusual tricuspid fossil as a pterosaur, which he named Tribelesodon longobardicus. The holotype specimen of Tribelesodon longobardicus was stored in the Museo Civico di Storia Naturale di Milano (Natural History Museum of Milan), and was destroyed by allied bombing of Milan in World War II.
Excavations by University of Zürich paleontologist Bernhard Peyer in the late 1920s and 1930s revealed many more complete fossils of the species from Monte San Giorgio. Peyer's discoveries allowed Tribelesodon longobardicus to be recognised as a non-flying reptile, more than 40 years after its original description. Its supposed elongated finger bones were recognized as neck vertebrae, which compared favorably with those previously described as Tanystropheus from Germany and Poland. Thus, Tribelesodon longobardicus was renamed to Tanystropheus longobardicus and its anatomy was revised into a long-necked, non-pterosaur reptile. Specimen PIMUZ T 2791, which was discovered in 1929, has been designated as the neotype of the species.
Well-preserved T. longobardicus fossils continue to be recovered from Monte San Giorgio up to the present day. Fossils from the mountain are primarily stored at the rebuilt Museo Civico di Storia Naturale di Milano (MSNM), the Paleontological Museum of Zürich (PIMUZ), and the Museo Cantonale di Scienze Naturali di Lugano (MCSN). Rupert Wild reviewed and redescribed all specimens known at the time via several large monographs in 1973/4 and 1980. In 2005, Silvio Renesto described a T. longobardicus specimen from Switzerland which preserved the impressions of skin and other soft tissue. Five new MSNM specimens of T. longobardicus were described by Stefania Nosotti in 2007, allowing for a more comprehensive view of the species' anatomy.
A small but well-preserved skull and neck, specimen PIMUZ T 3901, was found in the slightly younger Meride Limestone at Monte San Giorgio. Wild (1980) gave it a new species, T. meridensis, based on a set of skull and vertebral traits proposed to differ from T. longobardicus. Later reinvestigations failed to confirm the validity of these differences, rendering T. meridensis a junior synonym of T. longobardicus. A 2019 revision of Tanystropheus concluded that T. longobardicus and T. antiquus were the only valid species in the genus, though this conclusion would prove to be short-lived.
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