Hauffiopteryx
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''Hauffiopteryx'' 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 nom ...
of ichthyosaur known from
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
,
Luxembourg Luxembourg ( ; lb, Lëtzebuerg ; french: link=no, Luxembourg; german: link=no, Luxemburg), officially the Grand Duchy of Luxembourg, ; french: link=no, Grand-Duché de Luxembourg ; german: link=no, Großherzogtum Luxemburg is a small lan ...
and
Somerset ( en, All The People of Somerset) , locator_map = , coordinates = , region = South West England , established_date = Ancient , established_by = , preceded_by = , origin = , lord_lieutenant_office =Lord Lieutenant of Somerset , lord_ ...
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 ...
. 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. 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 attached. In other words, a type is an example that serves to anchor or centralizes the ...
GPIT 1491/4, articulated complete skeleton which preserved the
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
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 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 f ...
(
Posidonia Shale The Posidonia Shale (german: Posidonienschiefer, also called Schistes Bitumineux in Luxembourg) geologically known as the Sachrang Formation, is an Early Jurassic (Toarcian) geological formation of southwestern and northeast Germany, northern Swit ...
) 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 ...
, dating to the 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 Toarcian ...
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-J ...
, about 182
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago) ...
. 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 ...
, Germany and
Dudelange Dudelange (; lb, Diddeleng , german: Düdelingen) is a commune with town status in southern Luxembourg. It is the fourth-most populous commune, with 19,734 inhabitants. Dudelange is situated close to the border with France. , the town of Dudela ...
, 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 minster town and civil parish in the South Somerset district of 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 C ...
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 in 1931 as a subspecies of ''S. hauffianus'' and named ''Stenopterygius hauffianus typica''. Michael W. Maisch in
2008 File:2008 Events Collage.png, From left, clockwise: Lehman Brothers went bankrupt following the Subprime mortgage crisis; Cyclone Nargis killed more than 138,000 in Myanmar; A scene from the opening ceremony of the 2008 Summer Olympics in Beijing; ...
elevated it to specific rank and reassigned to its own genus, ''Hauffiopteryx''. 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 specime ...
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 ...
, 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: Greece 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 ...
for "fin" or "wing". The specific name means typical. ''H. altera'' comes from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
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 and in body mass.


Skull

The snout of ''Hauffiopteryx'' is short, but also very thin, tapering to a point. The outer surface of the
tooth crown In dentistry, crown refers to the anatomical area of teeth, usually covered by enamel. The crown is usually visible in the mouth after developing below the gingiva The gums or gingiva (plural: ''gingivae'') consist of the mucosal tiss ...
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'' 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. Foss ...
''. 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 mammal has ...
e (upper tooth-bearing bones). In ''H. altera'', the tallest points of 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. T ...
e (rear upper tooth-bearing bones) are positioned behind the external nares (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 bone is a small and fragile bone of the facial skeleton; it is roughly the size of the little fingernail. It is situated at the front part of the medial wall of the Orbit (anatomy), orbit. It has two surfaces and four borders. Several ...
(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'', the are triradiate, while those of ''H. altera'' are massive and triangular. The
orbits In celestial mechanics, an orbit is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an object or position in space such as a p ...
(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 skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, t ...
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, also known as a third eye or pineal eye, is a part of the epithalamus present in some vertebrates. The eye is located at the top of the head, is photoreceptive and is associated with the pineal gland, regulating circadian rhyth ...
, a small opening on the midline of the skull. Instead of being in line with the front edges of the supratemporal fenestrae (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 t ...
(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 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 characteristi ...
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 ribs (those in the trunk region) are strongly forked, making them double-headed.
Gastralia Gastralia (singular 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 thes ...
(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 (plural 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 eith ...
e (shoulder blades) of ''Hauffiopteryx'' have wide lower ends due to their front margins extending forwards. The front edge of each
coracoid A coracoid (from Greek κόραξ, ''koraks'', raven) 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 prese ...
(shoulder bones located below the scapulae) is concave. No space is enclosed between the
humeri The humerus (; ) 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 a round ...
(upper arm bones) and the
radii In classical geometry, a radius ( : radii) 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 name comes from the latin ''radius'', meaning ray but also the ...
and
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
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, and covering a surface. Mosaics are often used as floor and wall decoration, and were particularly pop ...
-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 (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 p ...
bones (the
pubic bones In vertebrates, the pubic region ( la, pubis) is the most forward-facing (ventral and anterior) of the three main regions making up the coxal bone. The left and right pubic regions are each made up of three sections, a superior ramus, inferior r ...
and
ischia Ischia ( , , ) is a volcanic island in the Tyrrhenian Sea. It lies at the northern end of the Gulf of Naples, about from Naples. It is the largest of the Phlegrean Islands. Roughly trapezoidal in shape, it measures approximately east to west ...
) are narrow and pillar-shaped. Unusually, the ends of these bones which form the
hip socket The acetabulum (), also called the cotyloid cavity, is a concave surface of the pelvis. The head of the femur meets with the pelvis at the acetabulum, forming the hip joint. Structure There are three bones of the ''os coxae'' (hip bone) that ...
are fused together, but the bones markedly diverge away from the socket. The thin
femora 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 with t ...
(thighbones) each bear a strong site for the articulation of the
fibula The fibula 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. Its upper extremity i ...
(rear shin bone). The fibulae are also much larger than 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 ...
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 member of Eurhinosauria or a basalmost member of Thunnosauria (which is an equivalent position to a basalmost member of Stenopterygiidae sensu Maisch
008 008, OO8, O08, or 0O8 may refer to: * The Streetwear Brand @008us , inspired by Ian Fleming & Virgil Abloh *"030", the fictional 030 Agent of MI6 * '' 038: Operation Exterminate'', a 1965 Italian action film * '' Explosivo 030'' a 1940 Argentine c ...
with exclusion of ''
Ichthyosaurus ''Ichthyosaurus'' (derived from Greek ' () meaning 'fish' and ' () meaning 'lizard') is a genus of ichthyosaurs from the Early Jurassic (Hettangian - Pliensbachian), with possible Late Triassic record, from Europe ( Belgium, England, Germany, ...
'').


References

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