Hauffiosaurus
Encyclopedia
Hauffiosaurus is an extinct genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 of Early Jurassic
Early Jurassic
The Early Jurassic epoch is the earliest of three epochs of the Jurassic period...

 (early Toarcian stage) pliosaurid plesiosaur
Plesiosaur
Plesiosauroidea is an extinct clade of carnivorous plesiosaur marine reptiles. Plesiosauroids, are known from the Jurassic and Cretaceous Periods...

 known from Holzmaden
Holzmaden
Holzmaden is a town in Baden-Württemberg, Germany that lies between Stuttgart and Ulm.The ground in and around the city contains rich layers of well preserved fossils of the Jurassic period. The fossils are found in the 160 million year old Posidonia Shale and displayed in the local Museum Hauff...

 of Germany
Germany
Germany , officially the Federal Republic of Germany , is a federal parliamentary republic in Europe. The country consists of 16 states while the capital and largest city is Berlin. Germany covers an area of 357,021 km2 and has a largely temperate seasonal climate...

 and from Yorkshire
Yorkshire
Yorkshire is a historic county of northern England and the largest in the United Kingdom. Because of its great size in comparison to other English counties, functions have been increasingly undertaken over time by its subdivisions, which have also been subject to periodic reform...

 of the United Kingdom
United Kingdom
The United Kingdom of Great Britain and Northern IrelandIn the United Kingdom and Dependencies, other languages have been officially recognised as legitimate autochthonous languages under the European Charter for Regional or Minority Languages...

. It was first named by Frank Robin O’Keefe in 2001
2001 in paleontology
-Newly named insects:-Newly named bony fishes:-Newly named amphibians:-Newly named turtles:-Ichthyosaurs:-Newly named basal lepidosauromorphs:-Newly named plesiosaurs:-Newly named sphenodonts:-Newly named basal archosauromorphs:...

 and the type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...

 is Hauffiosaurus zanoni. In 2011, two additional species were assigned to this genus: H. longirostris and H. tomistomimus.

Description

Hauffiosaurus is known from the holotype
Holotype
A holotype is a single physical example of an organism, known to have been used when the species was formally described. It is either the single such physical example or one of several such, but explicitly designated as the holotype...

 and only specimen uncataloged Hauff Museum, an almost complete and articulated skeleton
Skeleton
The skeleton is the body part that forms the supporting structure of an organism. There are two different skeletal types: the exoskeleton, which is the stable outer shell of an organism, and the endoskeleton, which forms the support structure inside the body.In a figurative sense, skeleton can...

, found from the Posidonien-Schiefer, dating to early Toarcian
Toarcian
The Toarcian is, in the ICS' geologic timescale, an age or stage in the Early or Lower Jurassic. It spans the time between 183.0 Ma and 175.6 Ma...

 stage of the Early Jurassic
Early Jurassic
The Early Jurassic epoch is the earliest of three epochs of the Jurassic period...

. The holotype
Holotype
A holotype is a single physical example of an organism, known to have been used when the species was formally described. It is either the single such physical example or one of several such, but explicitly designated as the holotype...

 specimen preserved in a single block of the original matrix, exposed in ventral view. The body outline visible around the specimen is an artifact of preparation, not preservation; no remains of soft tissue were preserved. The skeleton was discovered during the early 19th Century, in beds of the famous Posidonien-Schiefer lagerstätte
Lagerstätte
A Lagerstätte is a sedimentary deposit that exhibits extraordinary fossil richness or completeness.Palaeontologists distinguish two kinds....

 at Holzmaden
Holzmaden
Holzmaden is a town in Baden-Württemberg, Germany that lies between Stuttgart and Ulm.The ground in and around the city contains rich layers of well preserved fossils of the Jurassic period. The fossils are found in the 160 million year old Posidonia Shale and displayed in the local Museum Hauff...

, Baden-Württemberg
Württemberg
Württemberg , formerly known as Wirtemberg or Wurtemberg, is an area and a former state in southwestern Germany, including parts of the regions Swabia and Franconia....

, in southeastern Germany. However, it was not recognized as a valid taxon
Taxon
|thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement...

, and no thorough description of the fossil
Fossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...

 was made until 2001.

Hauffiosaurus zanoni is a plesiosaur of medium size, measuring 3.4 metres (11 ft). The skull measures about 430 millimetres (1 ft) along the midline. The holotype is an adult individual, but incomplete fusion of the pectoral
Pectoral
Pectoral may refer to:* Pectoral cross, a cross worn on the chest* a decorative, usually jeweled version of a gorget* Pectoral , often iconographic with hieroglyphs* Pectoralis major muscle, commonly referred to as "pectorals" or "pecs"...

 and pelvic girdle indicate it's not an old adult. The specimen is displayed at the Urwelt-Museum Hauff, Holzmaden. The exact phylogenetic position of Hauffiosaurus within the Plesiosauria has yet to resolved, though Vincent (2011) states that it may "reasonably be placed within the Pliosauroidea".
A second species, H. tomistomimus, was named by Roger B. J. Benson, Hilary F. Ketchum, Leslie F. Noè and Marcela Gómez-Pérez in 2011
2011 in paleontology
-Lobopods:-Arachnids:-Insects:-Newly named jawless vertebrates:-Newly named acanthodian:-Newly named cartilaginous fishes:-Newly named bony fishes:-Newly named lepospondyls:-Newly named temnospondyls:-Newly named lissamphibians:...

. It is known from the holotype
Holotype
A holotype is a single physical example of an organism, known to have been used when the species was formally described. It is either the single such physical example or one of several such, but explicitly designated as the holotype...

 and only specimen, MMUM LL 8004, an almost complete, three-dimensionally preservedand and articulated skeleton
Skeleton
The skeleton is the body part that forms the supporting structure of an organism. There are two different skeletal types: the exoskeleton, which is the stable outer shell of an organism, and the endoskeleton, which forms the support structure inside the body.In a figurative sense, skeleton can...

, found from the Hildoceras bifrons Zone of the Alum Shale Member, Whitby Limestone Formation, dating to early Toarcian
Toarcian
The Toarcian is, in the ICS' geologic timescale, an age or stage in the Early or Lower Jurassic. It spans the time between 183.0 Ma and 175.6 Ma...

 stage. This skeleton was discovered in Yorkshire
Yorkshire
Yorkshire is a historic county of northern England and the largest in the United Kingdom. Because of its great size in comparison to other English counties, functions have been increasingly undertaken over time by its subdivisions, which have also been subject to periodic reform...

, UK. The holotype H. tomistomimus is about 4.83 metres (16 ft) long. Benson et al. 2011 reassigned Macroplata
Macroplata
Macroplata , meaning, , is an extinct genus of Early Jurassic basal plesiosaur which grew up to in length. Like other plesiosaurs, Macroplata probably lived on a diet of fish, using its sharp needle-like teeth to catch prey. Its shoulder bones were fairly large, indicating a powerful forward...

 longirostris (originally Plesiosaurus
Plesiosaurus
Plesiosaurus was a genus of large marine sauropterygian reptile that lived during the early part of the Jurassic Period, and is known by nearly complete skeletons from the Lias of England...

 longirostris) to Hauffiosaurus. The holotype and only specimen of H. longirostris, MCZ 1033, found from the Harpoceras serpentinum Zone of the Alum Shale Member, Whitby Limestone Formation, early Toarcian of Yorkshire, England.

Few Early Jurassic plesiosaurians have the sort of long, thin snout present in Hauffiosaurus, a shape usually considered to indicate a diet of fish
Fish
Fish are a paraphyletic group of organisms that consist of all gill-bearing aquatic vertebrate 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...

 (ichthyophagy). The teeth are slender and elongated and possess fine longitudinal ridges. This sort of tooth
Tooth
Teeth are small, calcified, whitish structures found in the jaws of many vertebrates that are used to break down food. Some animals, particularly carnivores, also use teeth for hunting or for defensive purposes. The roots of teeth are embedded in the Mandible bone or the Maxillary bone and are...

, coupled with the long rostrum
Rostrum (anatomy)
The term rostrum is used for a number of unrelated structures in different groups of animals:*In crustaceans, the rostrum is the forward extension of the carapace in front of the eyes....

 seen in this genus is effective in piercing soft prey.

Phylogeny

Smith & Dyke, 2008 were the first who found Hauffiosaurus to be basal pliosauroid. Benson, Ketchum, Noè and Gómez-Pérez who assigned two additional species to this genus, confirmed the affinity of this taxon to Pliosauroidea using cladistic analysis which was based on Ketchum & Benson's (2010) analysis. Cladogram
Cladogram
A cladogram is a diagram used in cladistics which shows ancestral relations between organisms, to represent the evolutionary tree of life. Although traditionally such cladograms were generated largely on the basis of morphological characters, DNA and RNA sequencing data and computational...

 after Benson et al., 2011:
With the description of Marmornectes
Marmornectes
Marmornectes is an extinct genus of pliosaurid known from the Bedfordshire of United Kingdom.-Description:Marmornectes is known from the holotype BEDFM 1999.201, articulated partial skeleton which preserved the skull...

 in 2011, Ketchum & Benson suggested for the first time that many basal plesiosaurs and pliosauroids are members of Pliosauridae
Pliosauridae
Pliosauridae is a family of pliosauroid plesiosaurs from the Earliest Jurassic to the early Late Cretaceous of Australia, Europe, North America and South America. It was formally named by Harry G...

 and Rhomaleosauridae
Rhomaleosauridae
Rhomaleosauridae is a family of extinct reptiles in the Plesiosauria order. It includes the Macroplata, Maresaurus, Rhomaleosaurus, Yuzhoupliosaurus and Meyerasaurus genera....

. Both Rhomaleosauridae and Pliosauridae were found to be monophyletic, and the relations between Hauffiosauruss species remained the same. The cladogram below follows Ketchum & Benson, 2011.
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