Aegirosaurus
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''Aegirosaurus'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
platypterygiine Platypterygiinae is an extinct subfamily of ophthalmosaurid thunnosaur ichthyosaurs from the early Late Jurassic to the early Late Cretaceous (Kimmeridgian - Cenomanian) of Asia, Australia, Europe, North America and South America. Currentl ...
ophthalmosaurid Ophthalmosauridae is an extinct family of thunnosaur ichthyosaurs from the Middle Jurassic to the early Late Cretaceous (Bajocian - Cenomanian) worldwide. Almost all ichthyosaurs from the Middle Jurassic onwards belong to the family, until the ex ...
ichthyosaur Ichthyosaurs (Ancient Greek for "fish lizard" – and ) are large extinct marine reptiles. Ichthyosaurs belong to the order known as Ichthyosauria or Ichthyopterygia ('fish flippers' – a designation introduced by Sir Richard Owen in 1842, altho ...
s known from the
late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
and
early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
of Europe. It was originally named as a species 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, S ...
''.


Discovery and species

Originally described by Wagner (1853) as a species of the genus ''
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, S ...
'' (''I. leptospondylus''), the species ''Aegirosaurus leptospondylus'' has had an unstable taxonomic history. It has been referred to the species '' Ichthyosaurus trigonus posthumus'' (later reclassified in the dubious genus '' Macropterygius'') in the past, and sometimes identified with ''
Brachypterygius extremus ''Brachypterygius'' (meaning ″short wing/paddle″ in Greek) is an extinct genus of platypterygiine ophthalmosaurid ichthyosaur known from the Late Jurassic of England.McGowan, C. & Motani, R. ''Ichthyopterygia''. In Sues, H.-D. (ed.) Handb ...
''. In 2000, Bardet and Fernández selected a complete skeleton in a private collection as the
neotype 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 ...
for the species ''I. leptospondylus'', as the only other described specimen was destroyed in
World War II World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the vast majority of the world's countries—including all of the great powers—forming two opposin ...
. A second specimen from the Munich collection was referred to the same taxon. Bardet and Fernández concluded that the neotype should be assigned to a new genus, ''Aegirosaurus''. The name means ' Aegir (teutonic god of the ocean) lizard with slender vertebrae'. Within the
Ophthalmosauridae Ophthalmosauridae is an extinct family of thunnosaur ichthyosaurs from the Middle Jurassic to the early Late Cretaceous (Bajocian - Cenomanian) worldwide. Almost all ichthyosaurs from the Middle Jurassic onwards belong to the family, until the ex ...
, scientists once believed ''Aegirosaurus'' was most closely related to ''
Ophthalmosaurus ''Ophthalmosaurus'' (meaning "eye lizard" in Greek) is an ichthyosaur of the Jurassic period (165–150 million years ago). Possible remains from the Cretaceous, around 145 million years ago, are also known. It was a relatively medium-sized ichth ...
''. However, many subsequent
cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...
analyses found it is more closely related to '' Sveltonectes'' (and probably to ''
Undorosaurus ''Undorosaurus'' is an extinct genus of ophthalmosaurid ichthyosaur known from western Russia, Svalbard, and Poland. It was a large ichthyosaur, with the type species measuring long and weighing . Discovery and naming ''Undorosaurus'' was named ...
''). ''Aegirosaurus'' lineage was found to include ''
Brachypterygius ''Brachypterygius'' (meaning ″short wing/paddle″ in Greek) is an extinct genus of platypterygiine ophthalmosaurid ichthyosaur known from the Late Jurassic of England.McGowan, C. & Motani, R. ''Ichthyopterygia''. In Sues, H.-D. (ed.) Handb ...
'' and ''
Maiaspondylus ''Maiaspondylus'' is an extinct genus of platypterygiine ophthalmosaurid ichthyosaurs known from Northwest Territories of Canada, the Cambridge Greensand of England and the Voronezh Region of Russia. Description ''Maiaspondylus'' is known ...
'', too, and to nest within the
Platypterygiinae Platypterygiinae is an extinct subfamily of ophthalmosaurid thunnosaur ichthyosaurs from the early Late Jurassic to the early Late Cretaceous (Kimmeridgian - Cenomanian) of Asia, Australia, Europe, North America and South America. Currently, the ...
, which is the sister taxon of
Ophthalmosaurinae Ophthalmosaurinae is an extinct subfamily of ophthalmosaurid thunnosaur ichthyosaurs from the Middle Jurassic to the late Early Cretaceous (Bajocian - Albian) of Europe, North America and South America. Currently, the oldest and the basalm ...
.


Stratigraphic range

''Aegirosaurus'' is known from the lower
Tithonian In the geological timescale, the Tithonian is the latest age of the Late Jurassic Epoch and the uppermost stage of the Upper Jurassic Series. It spans the time between 152.1 ± 4 Ma and 145.0 ± 4 Ma (million years ago). It is preceded by the K ...
(
Upper Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
) of
Bavaria Bavaria ( ; ), officially the Free State of Bavaria (german: Freistaat Bayern, link=no ), is a state in the south-east of Germany. With an area of , Bavaria is the largest German state by land area, comprising roughly a fifth of the total lan ...
,
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 ...
. Its remains were discovered in the Solnhofen limestone formations, which have yielded numerous well-known fossils, such as ''
Archaeopteryx ''Archaeopteryx'' (; ), sometimes referred to by its German name, "" ( ''Primeval Bird''), is a genus of bird-like dinosaurs. The name derives from the ancient Greek (''archaīos''), meaning "ancient", and (''ptéryx''), meaning "feather" ...
'', ''
Compsognathus ''Compsognathus'' (; Greek ''kompsos''/κομψός; "elegant", "refined" or "dainty", and ''gnathos''/γνάθος; "jaw") is a genus of small, bipedal, carnivorous theropod dinosaur. Members of its single species ''Compsognathus longipes'' c ...
'', and ''
Pterodactylus ''Pterodactylus'' (from Greek () meaning 'winged finger') is an extinct genus of pterosaurs. It is thought to contain only a single species, ''Pterodactylus antiquus'', which was the first pterosaur to be named and identified as a flying rept ...
''. In addition to its late Jurassic occurrence, ''Aegirosaurus'' has been discovered from the late
Valanginian In the geologic timescale, the Valanginian is an age or stage of the Early or Lower Cretaceous. It spans between 139.8 ± 3.0 Ma and 132.9 ± 2.0 Ma (million years ago). The Valanginian Stage succeeds the Berriasian Stage of the Lower Cretaceou ...
(early Cretaceous) of southeastern France ( Laux-Montaux, department of
Drôme Drôme (; Occitan: ''Droma''; Arpitan: ''Drôma'') is the southernmost department in the Auvergne-Rhône-Alpes region of Southeastern France. Named after the river Drôme, it had a population of 516,762 as of 2019.
; Vocontian Basin), the first diagnostic ichthyosaur recorded from the Valanginian. This shows that most types of late Jurassic ichthyosaurs crossed the Jurassic-Cretaceous boundary.


Description

Ichthyosaurs are marine reptiles showing extreme adaptations to life in the water, superficially resembling
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% of li ...
and
dolphins A dolphin is an aquatic mammal within the infraorder Cetacea. Dolphin species belong to the families Delphinidae (the oceanic dolphins), Platanistidae (the Indian river dolphins), Iniidae (the New World river dolphins), Pontoporiidae (t ...
in overall shape. ''Aegirosaurus'' is a medium-sized ichthyosaur. The neotype specimen is long, while the destroyed holotype was estimated to have been about long when complete. The skull of the neotype is long, while that of the smaller specimen BSPHGM 1954 I 608 is approximately in length.


Skull

The skull of ''Aegirosaurus'' gently tapers into an elongate, thin snout that makes up 62-73% of the length of the lower jaw. Its jaws are densely packed with small teeth lacking or at most bearing very small enamel ridges. The teeth are firmly rooted in a groove. The external nares (nostril openings) are long, thin, and two-lobed in shape, with one lobe protruding upwards. 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 socket) of Aegirosaurus are of medium size for an ichthyosaur, and each contains a ring of 14 bony plates (known as the
sclerotic ring Sclerotic rings are rings of bone found in the eyes of many animals in several groups of vertebrates, except for mammals and crocodilians. They can be made up of single bones or multiple segments and take their name from the sclera. They are bel ...
, which supported the eyeball) that occupies most of the space within. A flange projects outwards from above the orbit. The
jugal The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...
s (bones beneath the eye sockets) is short and extends in front of the orbit. The region of the skull behind the orbits is not very extensive, the length of this region being only about 6-8% of the
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
's (lower jaw) length. The
postorbital The ''postorbital'' is one of the bones in vertebrate skulls which forms a portion of the dermal skull roof and, sometimes, a ring about the orbit. Generally, it is located behind the postfrontal and posteriorly to the orbital fenestra. In some ve ...
s lie on top of the
quadratojugal The quadratojugal is a skull bone present in many vertebrates, including some living reptiles and amphibians. Anatomy and function In animals with a quadratojugal bone, it is typically found connected to the jugal (cheek) bone from the front and ...
s (two pairs of bones in this region), almost completely hiding them when seen from the side. ''Aegirosaurus'' also bears a pair of bones known as
squamosal The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone. In most tetrapods, the squamosal and quadratojugal The quadratojugal is a skull bone present in many vertebrates, including ...
s in its cheek region. These bones are small, triangular, and can be seen when the skull is viewed from the side. However, two pairs of bones of the skull roof, 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, the ...
s and
supratemporal The supratemporal bone is a paired cranial bone present in many tetrapods and tetrapodomorph fish. It is part of the temporal region (the portion of the skull roof behind the eyes), usually lying medial (inwards) relative to the squamosal and latera ...
s, block them from reaching the
temporal fenestra An infratemporal fenestra, also called the lateral temporal fenestra or simply temporal fenestra, is an opening in the skull behind the orbit in some animals. It is ventrally bordered by a zygomatic arch. An opening in front of the eye sockets, ...
e (openings located on top of the skull). The mandible is strongly built. The angulars, one pair rear mandibular bones, can be seen on the mandible's outer surface and extend as far forwards as the
surangular The suprangular or surangular is a jaw bone found in most land vertebrates, except mammals. Usually in the back of the jaw, on the upper edge, it is connected to all other jaw bones: dentary, angular, splenial and articular The articular bone i ...
s, another pair of bones in the same region.


Postcranial skeleton

''Aegirosaurus'' is estimated to have had around 157 vertebrae, with 52 of them being presacral (neck and trunk) vertebrae while the other 105 caudal (tail) vertebrae. The tail was bent downards at a roughly 45 degree angle. This bend was formed by four vertebrae, with 40 caudal vertebrae in front of them and 61 behind. The vertebrae of ''Aegirosaurus'' are small and both ends of their
centra Centra is a convenience shop chain that operates throughout Ireland. The chain operates as a symbol group owned by Musgrave Group, the food wholesaler, meaning the stores are all owned by individual franchisees. The chain has three different f ...
(vertebral bodies) are concave. The centra are about twice as wide, and, in the case of the presacral vertebrae, tall as they are long. The diapophyses (projections from the top part of the vertebrae) flow smoothly into the
neural arches 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 characteristic ...
(which form the top part of the neural canal). Another set of projections, the parapophyses, which are located lower than the diapophyses, are positioned towards the front of the vertebrae. The slender
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 eithe ...
e (shoulder blades) of ''Aegirosaurus'' are expanded at both ends and constricted at their middles. The front edges of the
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 ...
s (another paired shoulder bone) lack notches. The
pelvis 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 ...
of ''Aegirosaurus'' is composed of two bones on each side, each
pubic bone 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 ra ...
and
ischium The ischium () form ...
(lower hip bones) having been fused into a single unit known as the puboischiatic complex. The lower end of the puboischiatic complex is over twice as wide as its upper, and the bone is not pierced by a
foramen In anatomy and osteology, a foramen (;Entry "foramen"
in
. The short, massively built
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) of ''Aegirosaurus'' have a lower ends that are wider than their upper ends. Each humerus bears facets for three bones on its lower end, two larger ones for the
radius 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 ...
(lower arm bones) and a smaller one in the middle for the wedge-shaped intermedium (one of the
carpals The carpal bones are the eight small bones that make up the wrist (or carpus) that connects the hand to the forearm. The term "carpus" is derived from the Latin carpus and the Greek καρπός (karpós), meaning "wrist". In human anatomy, th ...
). The radius is significantly smaller than the ulna, and the intermedium is longer from its lower to upper end than it is wide. An additional bone is present in each front flipper, located in front of and slightly lower than the radius. This bone, known as an extrazeugopodial element, has an accessory digit attached to it. In total, ''Aegirosaurus'' has six digits in each foreflipper, five of them primary digits (digits that contact the wrist). The fourth digit, which contains 23 individual bones, is the longest in the foreflipper. The hindflippers are short, with the
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 ...
(thigh bones) only half the length of the humeri and smaller than the lower hip bones. There are two bones, 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 ...
and
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 is ...
(shin bones) that contact the lower end of each femur, and the hind flippers each bear four digits, three of them primary digits (contacting the ankle). Both the front and hind flippers have a similar bone configuration, with the bones angular and tightly packed together further up while those further towards the flipper's tip being less tightly packed and rounder. Except for the fibulae, all of the bones in the flippers lack notches.


Soft tissue

The soft tissue of the front flippers of ''Aegirosaurus'' closely follows the shape of the bones. The front flippers are elongate but narrow, with their strongly concave rear edges. The hind flippers, however, have extensive soft tissue envelopes, making them wide. A caudal fin in the shape of a crescent moon is present on the tail. Bardet and Fernández reported three types of skin impressions in the neotype: a wavy texture running parallel to the body; straight, fibrous tissue at a right angle to the ripples; and what appeared to be very small
scales Scale or scales may refer to: Mathematics * Scale (descriptive set theory), an object defined on a set of points * Scale (ratio), the ratio of a linear dimension of a model to the corresponding dimension of the original * Scale factor, a number w ...
. This last texture was found on the upper lobe of the caudal fin, and would contradict the idea that ichthyosaurs lacked scales. However, the authors cautioned that the specimen would have to be studied using
microscope A microscope () is a laboratory instrument used to examine objects that are too small to be seen by the naked eye. Microscopy is the science of investigating small objects and structures using a microscope. Microscopic means being invisibl ...
s before the presence of scales on ''Aegirosaurus'' could be confirmed. A study of the ''Aegirosaurus'' sp. specimen JME-SOS-08369 by Lene Delsett and colleagues in 2022, however, found it to have smooth, scaleless skin, as typical for ichthyosaurs.


Palaeobiology

Tooth morphology and wear pattern suggest that ''Aegirosaurus'' belonged to the "Pierce II/ Generalist" feeding guild.


See also

* List of ichthyosaurs *
Timeline of ichthyosaur research This timeline of ichthyosaur research is a chronological listing of events in the history of paleontology focused on the ichthyosauromorphs, a group of secondarily aquatic marine reptiles whose later members superficially resembled dolphins, shar ...


References

{{Taxonbar, from=Q2640204 Late Jurassic ichthyosaurs Early Cretaceous ichthyosaurs Ichthyosaurs of Europe Jurassic reptiles of Europe Early Cretaceous reptiles of Europe Solnhofen fauna Fossil taxa described in 2000 Ophthalmosauridae Ichthyosauromorph genera