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''Hauffiosaurus'' 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
Early Jurassic The Early Jurassic Epoch (in chronostratigraphy corresponding to the Lower Jurassic Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-Jurassic extinction event, 201.3 Ma  ...
(early Toarcian stage) pliosaurid plesiosaur known from
Holzmaden Holzmaden is a town in Baden-Württemberg, Germany that lies between Stuttgart and Ulm. Holzmaden is 4 km south-east from Kirchheim unter Teck and 19 km south-east of Esslingen am Neckar. The A 8 runs south from Holzmaden. The town a ...
of
Germany Germany, officially the Federal Republic of Germany (FRG),, is a country in Central Europe. It is the most populous member state of the European Union. Germany lies between the Baltic and North Sea to the north and the Alps to the sou ...
and from
Yorkshire Yorkshire ( ; abbreviated Yorks), formally known as the County of York, is a historic county in northern England and by far the largest in the United Kingdom. Because of its large area in comparison with other English counties, functions have ...
of the
United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Europe, off the north-western coast of the European mainland, continental mainland. It comprises England, Scotlan ...
. It was first named by Frank Robin O’Keefe in
2001 The September 11 attacks against the United States by Al-Qaeda, which killed 2,977 people and instigated the global war on terror, were a defining event of 2001. The United States led a multi-national coalition in an invasion of Afghanistan ...
and the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen( ...
is ''Hauffiosaurus zanoni''. In 2011, two additional species were assigned to this genus: ''H. longirostris'' and ''H. tomistomimus''.


Description

The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
specimen of ''Hauffiosaurus'', housed in the Hauff Museum, is an almost complete and articulated
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
, found from the Posidonien-Schiefer, dating to early
Toarcian The Toarcian is, in the ICS' geologic timescale, an age and stage in the Early or Lower Jurassic. It spans the time between 182.7 Ma (million years ago) and 174.1 Ma. It follows the Pliensbachian and is followed by the Aalenian. The Toarc ...
stage of the
Early Jurassic The Early Jurassic Epoch (in chronostratigraphy corresponding to the Lower Jurassic Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-Jurassic extinction event, 201.3 Ma  ...
. The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
specimen preserved in a single block of the original matrix, exposed in
ventral 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 ...
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 A Lagerstätte (, from ''Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that exhibits extraordinary fossils with exceptional preservation—sometimes including preserved soft tissues. These for ...
at
Holzmaden Holzmaden is a town in Baden-Württemberg, Germany that lies between Stuttgart and Ulm. Holzmaden is 4 km south-east from Kirchheim unter Teck and 19 km south-east of Esslingen am Neckar. The A 8 runs south from Holzmaden. The town a ...
, Baden-
Württemberg Württemberg ( ; ) is a historical German territory roughly corresponding to the cultural and linguistic region of Swabia. The main town of the region is Stuttgart. Together with Baden and Hohenzollern, two other historical territories, Wür ...
, in southeastern Germany. However, it was not recognized as a valid
taxon In biology, a taxon ( back-formation from '' taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular n ...
, and no thorough description of the
fossil 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 ...
was made until 2001. ''Hauffiosaurus zanoni'' is a plesiosaur of medium size, measuring . The skull measures about along the midline. The holotype is an adult individual, but incomplete fusion of the pectoral and
pelvic The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton). The ...
girdle indicate it is not an old adult. The specimen is displayed at the Urwelt-Museum Hauff, Holzmaden. The exact
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
position of ''Hauffiosaurus'' within the Plesiosauria has yet to be 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. It is known from the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
and only specimen, MMUM LL 8004, an almost complete, three-dimensionally preservedand and articulated
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
, found from the '' Hildoceras bifrons'' Zone (181.2–180.7 Ma) of the
Alum Shale Member An alum () is a type of chemical compound, usually a hydrated double sulfate salt of aluminium with the general formula , where is a monovalent cation such as potassium or ammonium. By itself, "alum" often refers to potassium alum, wit ...
,
Whitby Mudstone Formation The Whitby Mudstone is a Toarcian (Early Jurassic; ''Falciferum''-''Bifrons'' in regional chronostratigraphy) geological formation in Yorkshire and Worcestershire, England.Toarcian The Toarcian is, in the ICS' geologic timescale, an age and stage in the Early or Lower Jurassic. It spans the time between 182.7 Ma (million years ago) and 174.1 Ma. It follows the Pliensbachian and is followed by the Aalenian. The Toarc ...
stage. This skeleton was discovered in
Yorkshire Yorkshire ( ; abbreviated Yorks), formally known as the County of York, is a historic county in northern England and by far the largest in the United Kingdom. Because of its large area in comparison with other English counties, functions have ...
, UK. The holotype of ''H. tomistomimus'', missing the snout tip, is about long. Benson ''et al.'' (2011) also reassigned ''
Macroplata ''Macroplata'' (meaning "big plate") is an extinct genus of Early Jurassic rhomaleosaurid plesiosaur which grew up to in length and weighed up to . Like other plesiosaurs, ''Macroplata'' probably lived on a diet of fish, using its sharp needl ...
longirostris'' (originally ''
Plesiosaurus ''Plesiosaurus'' (Greek: ' ('), near to + ' ('), lizard) is a genus of extinct, large marine sauropterygian reptile that lived during the Early Jurassic. It is known by nearly complete skeletons from the Lias of England. It is distinguishable by ...
longirostris'') to ''Hauffiosaurus''. The holotype and only specimen of ''H. longirostris'', MCZ 1033, found from the ''Harpoceras serpentinum'' Zone (182.7–181.2 Ma) of the
Jet Rock Member Jet, Jets, or The Jet(s) may refer to: Aerospace * Jet aircraft, an aircraft propelled by jet engines ** Jet airliner ** Jet engine ** Jet fuel * Jet Airways, an Indian airline * Wind Jet (ICAO: JET), an Italian airline * Journey to Enceladus a ...
, Whitby Mudstone Formation, early Toarcian of Yorkshire, England. The individual to which this specimen belongs is estimated to have measured approximately in total body length and weighed . 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 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% ...
(ichthyophagy). The teeth are slender and elongated and possess fine longitudinal ridges. This sort of
tooth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, t ...
, coupled with the long rostrum 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 Pliosauroidea is an extinct clade of plesiosaurs, known from the earliest Jurassic to early Late Cretaceous. They are best known for the subclade Thalassophonea, which contained crocodile-like short-necked forms with large heads and massive to ...
using cladistic analysis which was based on Ketchum & Benson's (2010) analysis.
Cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
after Benson ''et al.'', 2011: With the description of '' Marmornectes'' in 2011, Ketchum & Benson suggested for the first time that many basal plesiosaurs and pliosauroids are members of
Pliosauridae Pliosauridae is a family of plesiosaurian marine reptiles from the Latest Triassic to the early Late Cretaceous (Rhaetian to Turonian stages) of Australia, Europe, North America and South America. The family is more inclusive than the archetyp ...
and
Rhomaleosauridae Rhomaleosauridae is a family of plesiosaurs from the Earliest Jurassic to the latest Middle Jurassic (Hettangian to Callovian stages) of Europe, North America, South America and possibly Asia. Most rhomaleosaurids are known from England, many ...
. 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.


See also

* List of plesiosaur genera *
Timeline of plesiosaur research This timeline of plesiosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, taxonomic revisions, and cultural portrayals of plesiosaurs, an order of marine reptiles that flourished d ...


References

{{Taxonbar, from=Q3783670 Early Jurassic plesiosaurs of Europe Fossil taxa described in 2001 Pliosaurids Sauropterygian genera