''Langobardisaurus'' (, meaning Reptile of
Langobardi, in reference to the Long Bearded People, an ancient Italian civilization)
is an
extinct genus
Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial n ...
of
tanystropheid
Tanystropheidae is an extinct family of mostly marine archosauromorph reptiles that lived throughout the Triassic Period. They are characterized by their long, stiff necks formed from elongated cervical vertebrae with very long cervical ribs. Som ...
archosauromorph
Archosauromorpha ( Greek for "ruling lizard forms") is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs (such as crocodilians and dinosaurs, including birds) rather than lepidosaurs (such as tuataras, li ...
reptile, with one known species, ''L. pandolfii''. Its fossils have been found in
Italy
Italy ( it, Italia ), officially the Italian Republic, ) or the Republic of Italy, is a country in Southern Europe. It is located in the middle of the Mediterranean Sea, and its territory largely coincides with the homonymous geographical ...
and
Austria
Austria, , bar, Östareich officially the Republic of Austria, is a country in the southern part of Central Europe, lying in the Eastern Alps. It is a federation of nine states, one of which is the capital, Vienna, the most populous ...
, and it lived during the
Late Triassic
The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch ...
period, roughly 228 to 201 million years ago.
The ''Langobardisaurus'' was first discovered by Italian
paleontologist
Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of foss ...
Silvio Renesto in 1994 from the
Calcare di Zorzino Formation in
Northern Italy
Northern Italy ( it, Italia settentrionale, it, Nord Italia, label=none, it, Alta Italia, label=none or just it, Nord, label=none) is a geographical and cultural region in the northern part of Italy. It consists of eight administrative Regions ...
.
Description
Neck and skull
The ''Langobardisaurus'' was a small
reptile with a body size shorter than 50 cm.
[ Despite its small size, the ''Langobardisaurus'' featured a long neck with elongate ]cervical vertebrae
In tetrapods, cervical vertebrae (singular: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In ...
featuring low neural spines
The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic i ...
.[ Atop its long neck, the ''Langobardisaurus'' had a large yet short triangular skull that featured a small rostrum and large orbits.] Its large orbits are evidence of reliance on visual perception – this suggests that the ''Langobardisaurus'' likely had good eyesight. The ''Langobardisaurus'' skull morphology reflects its unique pattern of dentition. The front part of the upper jaw is toothless, although some grooves on the premaxilla
The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has ...
have been mistaken to be teeth in the past despite their lack of enamel. Past this toothless region of the snout, there were larger tricuspid (three-pronged) cheek teeth on the maxilla
The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The ...
and a large molariform tooth which is elongated in anteroposterior
Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
direction. This molariform tooth was flattened, with its occluding surface bent inwards and covered with tiny denticles. The lower jaw featured a similar molariform tooth which occluded with the aforementioned upper counterpart. Additionally, the lower jaw was robust and had a high coronoid process which suggests that the capability of a powerful bite.[ Given this and its distinct tooth pattern, these traits suggest that the ''Langobardisaurus'' performed excessive grinding of its food. However, none of the discovered specimens included the jaw articulation, so the conclusions that can be drawn are limited.] A dentition pattern as described is certainly unique, and not found in any other 'protorosaur
Protorosauria is an extinct polyphyletic group of archosauromorph reptiles from the latest Middle Permian (Capitanian stage) to the end of the Late Triassic ( Rhaetian stage) of Asia, Europe and North America. It was named by the English anato ...
'. In an analysis of the ''Langobardisaurus'' jaw and teeth morphology, Renesto and Dalla Vecchia speculated that the ''Langobardisaurus'' survived on a diet of large insects
Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs ...
, crustaceans
Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean gr ...
, and small fish
Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% ...
with tough scales. Additionally, it has been hypothesized that the ''Langobardisaurus'' used its long neck to pluck insects out of the air, in addition to burying its head deep into burrows to capture fleeing crustacean prey.
Tail
The long neck was opposed by an even longer tail, which featured 45 caudal vertebrae
The spinal column, a defining synapomorphy shared by nearly all vertebrates, Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
[ – making it twice the length of the trunk.] Paleontologists hypothesize that the long tail of ''Langobardisaurus'' was a key adaptation for the genus that had significant impacts on its daily activity. A long tail allowed the ''Langobardisaurus'' to balance its body in a bipedal stance, despite its long neck. Able to stand tall on its hind legs, the ''Langobardisaurus'' could have utilized its keen eyesight and long neck (extended vertically) to survey nearby terrain for both predators and prey as shown in the image to the left.
Limbs
The fossil record
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
shows that the ''Langobardisaurus'' featured short forelimbs dwarfed by much longer, hollow hind limbs. The tibia
The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
and fibula
The fibula or calf bone is a human leg, leg bone on the Lateral (anatomy), lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long ...
elements were slightly shorter than the femur
The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates wit ...
. Moving distally, the tarsi were small and compact.[
These facts suggest that the ''Langobardisaurus'' was capable of ]bipedal locomotion
Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' ...
.[ Bipedal locomotion was undoubtedly a large advantage for the ''Langobardisaurus'' - such an adaptation would have allowed the animal to both chase after prey and run from predators. Based on the hypothesis that the ''Langobardisaurus'' fed on ]insects
Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs ...
, crustaceans
Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean gr ...
, and fish
Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% ...
, the ability to run after prey afforded the ''Langobardisaurus'' a significant increase in its hunting capabilities, even if it was only able to run in short bursts. Based on the available morphological and geological information on the genus, paleontologists
Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
hypothesize that the ''Langobardisaurus'' had adaptation to aquatic life. Rather, the genus likely lived near marine environments - consistent with the proposition that it survived off of crustaceans
Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean gr ...
found in tidal flats
Mudflats or mud flats, also known as tidal flats or, in Ireland, slob or slobs, are coastal wetlands that form in intertidal areas where sediments have been deposited by tides or rivers. A global analysis published in 2019 suggested that tidal f ...
. The late Triassic topographical features of the regions in which the specimens were found further support this claim [
]
Discovery
To date, five specimens of ''Langobardisaurus'' have been discovered. In addition to the Calcare di Zorzino Formation previously described - which produced two specimens, an additional two Italian specimens were discovered in the Forni Dolostone Formation of Northern Italy
Northern Italy ( it, Italia settentrionale, it, Nord Italia, label=none, it, Alta Italia, label=none or just it, Nord, label=none) is a geographical and cultural region in the northern part of Italy. It consists of eight administrative Regions ...
.[
The most recent specimen of ''Langobardisaurus'' was found in 2013 in the Seefeld-Formation in the Northern Calcareous Alps of ]Tyrol
Tyrol (; historically the Tyrole; de-AT, Tirol ; it, Tirolo) is a historical region in the Alps - in Northern Italy and western Austria. The area was historically the core of the County of Tyrol, part of the Holy Roman Empire, Austrian Emp ...
Austria
Austria, , bar, Östareich officially the Republic of Austria, is a country in the southern part of Central Europe, lying in the Eastern Alps. It is a federation of nine states, one of which is the capital, Vienna, the most populous ...
.[ This finding is significant as it expands the paleontological range of the genus, which was previously confined to ]Northern Italy
Northern Italy ( it, Italia settentrionale, it, Nord Italia, label=none, it, Alta Italia, label=none or just it, Nord, label=none) is a geographical and cultural region in the northern part of Italy. It consists of eight administrative Regions ...
.
Saller et al. described the geological setting: “All specimens were found in dark limestone
Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms wh ...
and dolomite that formed in relatively small and deep marine basins surrounded by a shallow-water carbonate platform
A carbonate platform is a sedimentary body which possesses topographic relief, and is composed of autochthonic calcareous deposits. Platform growth is mediated by sessile organisms whose skeletons build up the reef or by organisms (usually mic ...
on which the peritidal sediments forming the Dolomia Principale/Hauptdolomit Formation were deposited”. The aforementioned basins lacked water circulation and ranged from insignificant oxygen levels to totally anoxic
The term anoxia means a total depletion in the level of oxygen, an extreme form of hypoxia or "low oxygen". The terms anoxia and hypoxia are used in various contexts:
* Anoxic waters, sea water, fresh water or groundwater that are depleted of diss ...
- making this environment suitable for preservation of vertebrate
Vertebrates () comprise all animal taxon, taxa within the subphylum Vertebrata () (chordates with vertebral column, backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the ...
bones.
Classification and other proposed species
There have been three proposed species of ''Langobardisaurus'', though only the first discovered, ''Langobardisaurus pandolfi'', is considered taxonomically significant. The two other proposed species: ''Langobardisaurus rossii'' and ''Langobardisaurus Tonelloi'' are no longer considered to be unique species of ''Langobardisaurus''. ''Langobardisaurus'' is considered to be either the sister-taxon of ''Tanystropheus
''Tanystropheus'' (Greek ~ 'long' + 'hinged') is an extinct archosauromorph reptile from the Middle and Late Triassic epochs. It is recognisable by its extremely elongated neck, which measured long—longer than its body and tail combined. T ...
'' or a close relative to ''Tanystropheus
''Tanystropheus'' (Greek ~ 'long' + 'hinged') is an extinct archosauromorph reptile from the Middle and Late Triassic epochs. It is recognisable by its extremely elongated neck, which measured long—longer than its body and tail combined. T ...
'' and ''Macrocnemus
''Macrocnemus'' is an extinct genus of archosauromorph reptile known from the Middle Triassic (Late Anisian to Ladinian) of Europe and China. ''Macrocnemus'' is a member of the Tanystropheidae family and includes three species''. Macrocnemus ...
''. These genera are classified to be a clade of archosauromorph
Archosauromorpha ( Greek for "ruling lizard forms") is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs (such as crocodilians and dinosaurs, including birds) rather than lepidosaurs (such as tuataras, li ...
diapsids called Protorosauria,[ although the clade may be an unnatural grouping. If this is the case, ''Langobardisaurus'', ''Tanystropheus'', ''Macrocnemus,'' and other close relatives (tanystropheids) may not be closely related to '' Protorosaurus''.
]
''Langobardisaurus tonelloi''
''Langobardisaurus tonelloi'' was first described by Muscio as a new species of ''Langobardisaurus'' in 1997 due to observed differences in phalangeal formula
The phalanges (singular: ''phalanx'' ) are digital bones in the hands and feet of most vertebrates. In primates, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as long bones ...
and limb bone proportion, though reinvestigation of these features has rendered them to be taxonomically insignificant.[
]
''Langobardisaurus rossii''
''Langobardisaurus rossii'' was first described by Bizzarini and Muscio in 1995. However, a detailed review of the specimen by Renesto and Dalla Vecchia led them to conclude that the ''Langobardisaurus rossii'' was a lepidosauromorph, and likely a rhynchocephalian
Rhynchocephalia (; ) is an order of lizard-like reptiles that includes only one living species, the tuatara (''Sphenodon punctatus'') of New Zealand. Despite its current lack of diversity, during the Mesozoic rhynchocephalians were a diverse gro ...
.[
]
References
{{Taxonbar, from=Q6486394
Tanystropheids
Late Triassic reptiles of Europe
Fossil taxa described in 1994
Prehistoric reptile genera