''Langobardisaurus'' (, meaning Reptile of

Langobardi The Lombards () or Longobards () were a Germanic people who conquered most of the Italian Peninsula between 568 and 774. The medieval Lombard historian Paul the Deacon wrote in the ''History of the Lombards'' (written between 787 and 796) th ...

, in reference to the Long Bearded People, an ancient Central-European civilisation of North Germanic origin) is an

extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

tanystropheid Tanystropheidae is an extinct family (biology), family of archosauromorph reptiles that lived throughout the Triassic Period, often considered to be "protorosaurs". They are characterized by their long, stiff necks formed from elongated cervical ...

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) than to lepidosaurs (such as tuataras, lizards, a ...

reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...

, with one valid species, ''L. pandolfii''. Its fossils have been found in

Italy Italy, officially the Italian Republic, is a country in Southern Europe, Southern and Western Europe, Western Europe. It consists of Italian Peninsula, a peninsula that extends into the Mediterranean Sea, with the Alps on its northern land b ...

and

Austria Austria, formally the Republic of Austria, is a landlocked country in Central Europe, lying in the Eastern Alps. It is a federation of nine Federal states of Austria, states, of which the capital Vienna is the List of largest cities in Aust ...

, 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 a ...

period, roughly 228 to 201 million years ago. ''Langobardisaurus'' was initially described in 1994, based on fossils from the Calcare di Zorzino Formation in

Northern Italy Northern Italy (, , ) is a geographical and cultural region in the northern part of Italy. The Italian National Institute of Statistics defines the region as encompassing the four Northwest Italy, northwestern Regions of Italy, regions of Piedmo ...

. Fossils of the genus are also known from the Forni Dolostone of Northern Italy and the Seefeld Formation of Austria.

Discovery

To date, five specimens of ''Langobardisaurus'' have been found.

Calcare di Zorzino specimens

The first fossils of the genus were discovered in 1974 at a quarry in

Cene, Lombardy Cene (Bergamasque: ) is a ''comune'' (municipality) in the Province of Bergamo in the Italian region of Lombardy, located about northeast of Milan and about northeast of Bergamo. Cene borders the following municipalities: Albino, Bianzano, Casn ...

exposing the

Calcare di Zorzino The Calcare di Zorzino, Italian for Zorzino Limestone is a Late Triassic (Norian) geological formation in Italy (Cene, Lombardy, Cene and ).described in 1994 by Italian

paleontologist Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...

Silvio Renesto. The

holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...

specimen of ''L. pandolfii'', MCSNB 2883, is an incomplete skeleton missing portions of the tail and forelimbs, while the neck and skull are crushed and displaced to the side. The smaller second specimen, MCSNB 4860, is a probable juvenile, with its skull and neck bent fully backwards under the

rib cage The rib cage or thoracic cage is an endoskeletal enclosure in the thorax of most vertebrates that comprises the ribs, vertebral column and sternum, which protect the vital organs of the thoracic cavity, such as the heart, lungs and great ve ...

. Both specimens are stored at the Museo civico di scienze naturali di Bergamo "E. Caffi" (MCSNB), a natural history museum in

Bergamo Bergamo ( , ; ) is a city in the Alps, alpine Lombardy region of northern Italy, approximately northeast of Milan, and about from the alpine lakes Lake Como, Como and Lake Iseo, Iseo and 70 km (43 mi) from Lake Garda, Garda and Lake ...

Dolomia di Forni specimens

Two additional Italian specimens were later discovered in the Dolomia di Forni (Forni Dolostone) in the

Friuli-Venezia Giulia Friuli-Venezia Giulia () is one of the 20 regions of Italy and one of five autonomous regions with special statute. The regional capital is Trieste on the Gulf of Trieste, a bay of the Adriatic Sea. Friuli-Venezia Giulia has an area of and a ...

region of

. One of the Forni specimens, MFSN 1921, is the most complete and well-preserved skeleton in the genus. It was originally described as a new species'', Langobardisaurus tonelloi'', by Muscio (1997). This was justified by apparent differences in

phalangeal formula The phalanges (: 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. Structure ...

and limb bone proportion, though reinvestigation of these features has rendered them to be taxonomically insignificant. ''L. tonelloi'' is thus considered synonymous with ''L. pandolfii''. The second Forni specimen, MFSN 26829, is much less complete, consisting of partial hindlimbs with associated tail and hip material. The Forni specimens are stored at the Museo Friulano di Storia Naturale (MFSN) in

Udine Udine ( ; ; ; ; ) is a city and (municipality) in northeastern Italy, in the middle of the Friuli-Venezia Giulia region, between the Adriatic Sea and the Carnic Alps. It is the capital of the Province of Udine, Regional decentralization entity ...

. Bizzarini and Muscio (1995) proposed a third species'', Langobardisaurus rossii'', based on a skeleton from the Dolomia di Forni, MCSN 19235. However, a detailed review of the specimen by Renesto and Dalla Vecchia in 2007 led them to conclude that the ''Langobardisaurus rossii'' was not actually referrable to the genus. Instead it was a fossil of an indeterminate

lepidosauromorph Lepidosauromorpha (in PhyloCode known as Pan-Lepidosauria) is a group of reptiles comprising all diapsids closer to lizards than to archosaurs (which include crocodiles and birds). The only living sub-group is the Lepidosauria, which contains two ...

, 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 speciose g ...

Austrian specimen

The most recent specimen of ''Langobardisaurus'' was described in 2013 from a fossil found in the Seefeld Formation in the

Northern Calcareous Alps The Northern Limestone Alps (), also called the Northern Calcareous Alps, are the ranges of the Eastern Alps north of the Central Eastern Alps located in Austria and the adjacent Bavarian lands of southeastern Germany. The distinction from the l ...

Tyrol Tyrol ( ; historically the Tyrole; ; ) is a historical region in the Alps of Northern Italy and western Austria. The area was historically the core of the County of Tyrol, part of the Holy Roman Empire, Austrian Empire and Austria-Hungary, f ...

. The Austrian specimen, P 10121, is an impression of a small nearly complete skeleton, close in size to MCSNB 4860. This finding is significant as it expands the paleontological range of the genus, which was previously confined to

. Saller et al. (2013) described the geological setting as “dark

limestone Limestone is a type of carbonate rock, carbonate sedimentary rock which is the main source of the material Lime (material), lime. It is composed mostly of the minerals calcite and aragonite, which are different Polymorphism (materials science) ...

and dolomite that formed in relatively small and deep marine basins surrounded by a shallow-water

carbonate platform A carbonate platform is a Sedimentary rock, sedimentary body which possesses topographic relief, and is composed of Autochthon (geology), autochthonic calcareous deposits. Platform growth is mediated by Sessility (zoology), sessile organisms whose ...

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 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 dissolved ox ...

- making this environment suitable for preservation of

vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...

bones.

Description

Neck and skull

''Langobardisaurus'' was a small

with a body size shorter than 50 cm. Despite its small size, ''Langobardisaurus'' featured a long neck with elongate

cervical vertebrae In tetrapods, cervical vertebrae (: 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 saurop ...

featuring low

neural spines Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...

. Atop its long neck, ''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 ''Langobardisaurus'' likely had good eyesight. The skull morphology of ''Langobardisaurus'' 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 mammals h ...

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 In vertebrates, the maxilla (: maxillae ) 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 two maxil ...

and a large molariform tooth which is elongated in an

anteroposterior Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position provi ...

(front-to-back) direction. This molariform tooth is flattened, with its occluding surface bent inwards and covered with tiny denticles. Langobardisaurus Skull Diagram

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 ''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, likely paraphyletic group of basal 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 t ...

'. In an analysis of ''Langobardisaurus'' jaw and teeth morphology, Renesto and Dalla Vecchia speculated thatthe ''Langobardisaurus'' survived on a diet of large

insects Insects (from Latin ') are 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 of jointed ...

crustaceans Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of Arthropod, arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquat ...

, and small

fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...

with tough scales. Additionally, it has been hypothesized that ''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 Caudal vertebrae are the vertebrae of the tail in many vertebrates. In birds, the last few caudal vertebrae fuse into the pygostyle, and in apes, including humans, the caudal vertebrae are fused into the coccyx. In many reptiles, some of the caud ...

– 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, ''Langobardisaurus'' could have utilized its keen eyesight and long neck (extended vertically) to survey nearby terrain for both predators and prey.

Limbs

''Langobardisaurus'' featured short forelimbs dwarfed by much longer, hollow hind limbs. The

tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...

and

fibula The fibula (: fibulae or fibulas) or calf bone is a leg bone on the 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 bones. ...

elements were slightly shorter than the

femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...

. Moving distally, the tarsi were small and compact. These facts suggest that ''Langobardisaurus'' was capable of

bipedal locomotion Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) 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'' ' ...

. Bipedal locomotion was undoubtedly a large advantage for ''Langobardisaurus'' - such an adaptation would have allowed the animal to both chase after prey and run from predators. Based on the hypothesis that ''Langobardisaurus'' fed on

, and

, the ability to run after prey afforded ''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 as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...

hypothesize that the ''Langobardisaurus'' likely lived near marine environments - consistent with the proposition that it survived off of

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 ...

. The Late Triassic topographical features of the regions in which the specimens were found further support this claim

Classification

''Langobardisaurus'' is a member of the

Tanystropheidae Tanystropheidae is an extinct family (biology), family of archosauromorph reptiles that lived throughout the Triassic Period, often considered to be "protorosaurs". They are characterized by their long, stiff necks formed from elongated cervical ...

, and is considered to be a close relative of ''

Tanystropheus ''Tanystropheus'' (~ 'long' + 'hinged') is an extinct genus of archosauromorph reptile which lived during the Triassic period, Triassic Period in Europe, Asia, and North America. It is recognisable by its extremely elongated neck, longer than 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 bas ...

'', which are also known from Triassic deposits of Italy (albeit from the Middle Triassic). Tanystropheids are classified as

diapsids Diapsids ("two arches") are a clade of sauropsids, distinguished from more primitive eureptiles by the presence of two holes, known as temporal fenestrae, in each side of their skulls. The earliest traditionally identified diapsids, the araeosc ...

. They have often been grouped with ''

Protorosaurus ''Protorosaurus'' (from , 'earlier' and , 'lizard') is an extinct genus of reptile. Members of the genus lived during the late Permian period in what is now Germany and Great Britain. Once believed to have been an ancestor to lizards, ''Protoro ...

'' and other long-necked early archosauromorphs in a group called

Protorosauria Protorosauria is an extinct, likely paraphyletic group of basal 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 t ...

. Some authors have suggested that "protorosaurs" are an unnatural grouping rather than a clade. If this is the case, tanystropheids may not be closely related to ''Protorosaurus''. Spiekman, Fraser & Schayer (2021) analyzed the systematics of "protorosaur" groups in their

phylogenetic analyses In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as Computational phylogenetics, phylogenetic inference. It infers the relationship among organ ...

. The following

cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...

shows the structure of Tanystropheidae according to one analysis, with ratio and ordered characters treated as such and pruning 5 out of 40 OTUs ''a posteriori'' to offer maximum resolution/minimum polytomies:

References

{{Taxonbar, from=Q6486394 Tanystropheidae Late Triassic reptiles of Europe Fossil taxa described in 1994 Prehistoric reptile genera