Hauffiopteryx Typicus Half
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''Hauffiopteryx'' 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 ...
of
ichthyosaur Ichthyosauria is an order of large extinct marine reptiles sometimes referred to as "ichthyosaurs", although the term is also used for wider clades in which the order resides. Ichthyosaurians thrived during much of the Mesozoic era; based on fo ...
known from
Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
,
Luxembourg Luxembourg, officially the Grand Duchy of Luxembourg, is a landlocked country in Western Europe. It is bordered by Belgium to the west and north, Germany to the east, and France on the south. Its capital and most populous city, Luxembour ...
,
Switzerland Switzerland, officially the Swiss Confederation, is a landlocked country located in west-central Europe. It is bordered by Italy to the south, France to the west, Germany to the north, and Austria and Liechtenstein to the east. Switzerland ...
and
Somerset Somerset ( , ), Archaism, archaically Somersetshire ( , , ) is a Ceremonial counties of England, ceremonial county in South West England. It is bordered by the Bristol Channel, Gloucestershire, and Bristol to the north, Wiltshire to the east ...
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 Northwestern Europe, off the coast of European mainland, the continental mainland. It comprises England, Scotlan ...
. Two species are known: ''H. typicus'' and ''H. altera''.


History of study

''Hauffiopteryx'' was first described by Michael W. Maisch on the basis of some specimens that previously referred to '' Stenopterygius hauffianus''. Maisch found that the lectotype of ''S. hauffianus'' can be determined as ''Stenopterygius cf. S. quadriscissus'' at best, and therefore this species should be considered a
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium,'' it may be impossible to determine whether a ...
. He also found out that most specimens previously referred to ''S. hauffianus'' can be referred to ''S. quadriscissus'', while the rest belongs to a highly distinctive new taxon that can't be referred to any valid species of ''Stenopterygius''. ''Hauffiopteryx'' is known from the
lectotype In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally associated. In other words, a type is an example that serves to anchor or centralizes ...
GPIT 1491/4, articulated complete
skeleton A skeleton is the structural frame that supports the body of most animals. There are several types of skeletons, including the exoskeleton, which is a rigid outer shell that holds up an organism's shape; the endoskeleton, a rigid internal fra ...
which preserved the
skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
and some soft tissues. The animal is about in length. It was collected from the ''Harpoceras elegantulum-exaratum'' ammonoid subzones (more specifically Lias ε II4), ''Harpoceras falcifer'' zone, of the famous Posidonien-Schiefer
lagerstätte A Fossil-Lagerstätte (, from ''Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that preserves an exceptionally high amount of palaeontological information. ''Konzentrat-Lagerstätten'' preserv ...
(
Posidonia Shale The Posidonia Shale (, also called Schistes Bitumineux in Luxembourg) geologically known as the Sachrang Formation, is an Early Jurassic (Early to Late Toarcian) geological formation in Germany, northern Switzerland, northwestern Austria, souther ...
) of
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 ...
, dating to the early
Toarcian The Toarcian is, in the International Commission on Stratigraphy, ICS' geologic timescale, an age (geology), age and stage (stratigraphy), stage in the Early Jurassic, Early or Lower Jurassic. It spans the time between 184.2 Megaannum, Ma (million ...
stage of the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic ...
, about 182
million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...
. Referred specimens 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 ...
, Germany and
Dudelange Dudelange (; ; ) is a commune with town status in southern Luxembourg. It is the fourth-most populous commune, with 22,043 inhabitants. Dudelange is situated close to the border with France. The commune also includes the smaller town of Bude ...
, Luxembourg include MHH '9', WAT 1, SMNS 51552, SMNS 80225 and probably the poorly preserved SMNS 81965. They were collected from the ''Harpoceras semicelatum-elegantulum-exaratum'' ammonoid subzones (Lias ε II1-5, about 182.7-181.8 mya), ''Harpoceras tenuicostatum-falcifer'' zones, of the Posidonia Shale. Additional materials were described by Hannah Caine and Michael J. Benton in 2011, from the early Toarcian of Strawberry Bank,
Ilminster Ilminster is a market town and civil parish in Somerset, England, with a population of 5,808. Bypassed in 1988, the town now lies just east of the junction of the A303 (London to Exeter) and the A358 (Taunton to Chard and Axminster). The parish ...
of England. The specimens are all juveniles or infants which preserved almost complete skeletons and some skulls. They include BRLSI M1399 (which was described earlier by Maisch), BRLSI M1400, BRLSI M1401, BRLSI M1403, BRLSI M1404 and BRLSI M1406.


Etymology

''Hauffiopteryx'' was originally recognized by
Friedrich von Huene Baron Friedrich Richard von Hoyningen-Huene (22 March 1875 – 4 April 1969) was a German nobleman paleontologist who described a large number of dinosaurs, more than anyone else in 20th-century Europe. He studied a range of Permo-Carbonife ...
in 1931 as a subspecies of ''S. hauffianus'' and named ''Stenopterygius hauffianus typica''. Michael W. Maisch in
2008 2008 was designated as: *International Year of Languages *International Year of Planet Earth *International Year of the Potato *International Year of Sanitation The Great Recession, a worldwide recession which began in 2007, continued throu ...
elevated it to specific rank and reassigned to its own genus, ''Hauffiopteryx''. The
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
is therefore ''Hauffiopteryx typicus''. The generic name honors the Hauff family 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 ...
, for their generation-long work to increase the knowledge on the Posidonia Shale and its fauna, especially the ichthyosaurs, and ''pteryx'' (πτερυξ),
Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...
for "fin" or "wing". The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
means typical. ''H. altera'' comes from the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
word meaning "different from" or "other" due to the differences between it and ''H. typicus''.


Description

''Hauffiopteryx'' is a relatively small ichthyosaur, reaching in length.


Skull

The snout of ''Hauffiopteryx'' is short, but also very thin, tapering to a point. The outer surface of the
tooth crown In dentistry, the crown is the visible part of the tooth above the gingival margin and is an essential component of dental anatomy. Covered by Tooth enamel, enamel, the crown plays a crucial role in cutting, tearing, and grinding food. Its shap ...
s is smooth, and the back teeth are larger than those in front. The upper jaw is slightly longer than the lower jaw, though this overbite is not carried to the extreme seen in ''
Excalibosaurus ''Excalibosaurus'' (meaning "Excalibur's lizard") is a monotypic genus of marine prehistoric reptiles (ichthyosaurs) that lived during the Sinemurian stage (approximately 199.5 ± 0.3 Ma to 192.9 ± 0.3 Ma (million years ago)) of the Early Jura ...
'' and ''
Eurhinosaurus ''Eurhinosaurus'' (Greek for 'well-nosed lizard'- eu meaning 'well or good', rhino meaning 'nose' and sauros meaning 'lizard') is an extinct genus of ichthyosaur from the Early Jurassic (Toarcian), ranging between 183 and 175 million years. Fossi ...
''. The
nasals In phonetics, a nasal, also called a nasal occlusive or nasal stop in contrast with an oral stop or nasalized consonant, is an occlusive consonant produced with a lowered velum, allowing air to escape freely through the nose. The vast majorit ...
form the majority of the snout's midline, rather than 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 ...
e (upper tooth-bearing bones). In ''H. altera'', the tallest points of 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 ...
e (rear upper tooth-bearing bones) are positioned behind the
external nares A nostril (or naris , : nares ) is either of the two orifices of the nose. They enable the entry and exit of air and other gasses through the nasal cavities. In birds and mammals, they contain branched bones or cartilages called turbinates, wh ...
(openings that housed the nostrils), in contrast to ''H. typicus'', where the tallest points are instead located beneath these openings. Part of the borders of the external nares are formed by the prefrontals (paired bones situated on the upper edges of the eye sockets). In ''H. altera'', this configuration blocks the
lacrimals The lacrimal bones are two small and fragile bones of the facial skeleton; they are roughly the size of the little fingernail and situated at the front part of the medial wall of the orbit. They each have two surfaces and four borders. Several bon ...
(paired bones in front of the eye sockets) from forming part of the external nares, whereas the lacrimals do reach the external nares in ''H. typicus''. A further difference between the two species lies in the shape of their lacrimals; in ''H. typicus'', they are triradiate, while those of ''H. altera'' are massive and triangular. The
orbits In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an physical body, object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an satellite, artificia ...
(eye sockets) of ''H. typicus'' are especially large and circular. In front of the orbits, the nasals of ''H. typicus'' curve upwards. The nasals extend further back in ''H. altera'' than in ''H. typicus'', though the nasals of ''Hauffiopteryx'' do not touch the parietals (a pair of skull roof bones). Externally, the prefrontals have greater surface area than the
postfrontal The postfrontal is a paired cranial bone found in many tetrapods. It occupies an area of the skull roof between and behind the orbits (eye sockets), lateral to the frontal and parietal bones, and anterior to the postorbital bone. The postfrontal ...
s, especially so in ''H. altera''. The frontals (a pair of skull roof bones) form most of the border of the
pineal foramen A parietal eye (third eye, pineal eye) is a part of the epithalamus in some vertebrates. The eye is at the top of the head; is photoreceptive; and is associated with the pineal gland, which regulates circadian rhythmicity and hormone production ...
, a small opening on the midline of the skull. Instead of being in line with the front edges of the
supratemporal fenestrae Temporal fenestrae are openings in the Temple (anatomy), temporal region of the skull of some Amniote, amniotes, behind the Orbit (anatomy), orbit (eye socket). These openings have historically been used to track the evolution and affinities of re ...
(paired openings on the top of the skull's rear), the foramen is instead located in front of them. The supratemporal fenestrae themselves are rounded and small. The supratemporals (paired skull roof bones) have wavy edges. Behind the orbits, the skull is short from front to back, with the cheek region deflected so that it faces backwards. The rear face
basioccipital The basilar part of the occipital bone (also basioccipital) extends forward and upward from the foramen magnum, and presents in front an area more or less quadrilateral in outline. In the young skull, this area is rough and uneven, and is joined ...
(braincase bone to which the vertebral column attaches) bears a considerable amount of surface which is not involved in the joint between the skull and the spine.


Postcranial skeleton

In 2011, Caine and Benton stated that ''Hauffiopteryx'' has fewer than 46
vertebrae 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 ...
in front of its hips (presacral vertebrae) and typically over 34 but under 39 between its hips and the bend in its tail. Maxwell and Cortés in 2020 described the lectotype as having 45 or 46 presacral vertebrae and a total of 81 vertebrae before the bend in the tail (preflexural vertebrae), with this bend composed of three vertebrae followed by more than 55 postflexural vertebrae. The upper ends of the dorsal
rib In vertebrate anatomy, ribs () are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the thoracic cavity, enabling the lungs to expand and thus facilitate breathing by expanding the ...
s (those in the trunk region) are strongly forked, making them double-headed.
Gastralia Gastralia (: gastralium) are dermal bones found in the ventral body wall of modern crocodilians and tuatara, and many prehistoric tetrapods. They are found between the sternum and pelvis, and do not articulate with the vertebrae. In these reptil ...
(belly ribs) are present along the underside of the trunk. This series of elements reaches far back, all the way to the level of the 35th vertebra. The
scapula The scapula (: scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on either side ...
e (shoulder blades) of ''Hauffiopteryx'' have wide lower ends due to their front margins extending forwards. The front edge of each
coracoid A coracoid is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is present as part of the scapula, but this is n ...
(shoulder bones located below the scapulae) is concave. No space is enclosed between the
humeri The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of ...
(upper arm bones) and the
radii In classical geometry, a radius (: radii or radiuses) of a circle or sphere is any of the line segments from its center to its perimeter, and in more modern usage, it is also their length. The radius of a regular polygon is the line segment or ...
and
ulna The ulna or ulnar bone (: ulnae or ulnas) is a long bone in the forearm stretching from the elbow to the wrist. It is on the same side of the forearm as the little finger, running parallel to the Radius (bone), radius, the forearm's other long ...
e (lower arm bones), being tightly packed together. This is also the case for the other bones in the upper parts of the limbs, forming a
mosaic A mosaic () is a pattern or image made of small regular or irregular pieces of colored stone, glass or ceramic, held in place by plaster/Mortar (masonry), mortar, and covering a surface. Mosaics are often used as floor and wall decoration, and ...
-like pattern. Each forelimb contains four main digits, and the front edge of the limb consists of some elements with indentations on their front edges. The lower
pelvic The pelvis (: pelves or pelvises) is the lower part of an anatomical 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). ...
bones (the
pubic bones In vertebrates, the pubis or pubic bone () forms the lower and anterior part of each side of the hip bone. The pubis is the most forward-facing (ventral and anterior) of the three bones that make up the hip bone. The left and right pubic bones ar ...
and
ischia Ischia ( , , ) is a volcanic island in the Tyrrhenian Sea. It lies at the northern end of the Gulf of Naples, about from the city of Naples. It is the largest of the Phlegrean Islands. Although inhabited since the Bronze Age, as a Ancient G ...
) are narrow and pillar-shaped. Unusually, the ends of these bones which form the hip socket are fused together, but the bones markedly diverge away from the socket. The thin
femora The femur (; : femurs or femora ), or thigh bone is the only bone in the thigh — the region of the lower limb between the hip and the knee. In many four-legged animals the femur is the upper bone of the hindleg. The top of the femur fits in ...
(thighbones) each bear a strong site for the articulation of the
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. ...
(rear shin bone). The fibulae are also much larger than 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 ...
e (front shin bones). While the front edges of the hindlimbs contain some notched elements like the forelimbs, there are only three main digits in each rear limb. These digits closely approach each other towards the tip of the limb.


Classification

Both the original description by Maisch and the redescription of the English specimens found that ''Hauffiopteryx'' might be either a
basalmost In phylogenetics, basal is the direction of the ''base'' (or root) of a rooted phylogenetic tree or cladogram. The term may be more strictly applied only to nodes adjacent to the root, or more loosely applied to nodes regarded as being close to th ...
member of
Eurhinosauria Leptonectidae is a family of ichthyosaurs Ichthyosauria is an taxonomy (biology), order of large extinction, extinct marine reptiles sometimes referred to as "ichthyosaurs", although the term is also used for wider clades in which the order res ...
or a basalmost member of
Thunnosauria Thunnosauria (Ancient Greek, Greek for "tuna lizard" – ''thunnos'' meaning "tuna" and ''sauros'' meaning "lizard") is an extinct clade of parvipelvian ichthyosaurs from the Early Jurassic to the early Late Cretaceous (Hettangian–Cenomanian) o ...
(which is an equivalent position to a basalmost member of
Stenopterygiidae Stenopterygiidae are a family of the Ichthyosauria Ichthyosauria is an order of large extinct marine reptiles sometimes referred to as "ichthyosaurs", although the term is also used for wider clades in which the order resides. Ichthyosaurians ...
sensu Maisch
008 008, OO8, O08, or 0O8 may refer to: * "008", a fictional 00 Agent In Ian Fleming's James Bond novels and the derived films, the 00 Section of MI6 is considered the secret service's elite. A 00 (pronounced "Double O") is a field agent who ho ...
with exclusion of ''
Ichthyosaurus ''Ichthyosaurus'' (derived from Greek () meaning 'fish' and () meaning 'lizard') is a genus of ichthyosaurs from the Early Jurassic (Hettangian - Pliensbachian) of Europe (Belgium, England, Germany and Portugal). Some specimens of the ichthy ...
'').


References

{{Taxonbar, from=Q5682360 Early Jurassic ichthyosaurs Fossil taxa described in 2008 Ichthyosaurs of Europe Ichthyosauromorph genera