title: Ichthyosaur

text: Ichthyosaurs became even more popular in 1854 by the rebuilding at Sydenham Hill of the Crystal Palace, originally erected at the world exhibition of 1851. In the surrounding park, life-sized, painted, concrete statues of extinct animals were placed, which were designed by Benjamin Waterhouse Hawkins under the direction of Richard Owen. Among them were three models of an ichthyosaur. Although it was known that ichthyosaurs had been animals of the open seas, they were shown basking on the shore, a convention followed by many nineteenth century illustrations with the aim, as Conybeare once explained, of better exposing their build. This led to the misunderstanding that they really had an amphibious lifestyle. The pools in the park were at the time subjected to tidal changes, so that fluctuations in the water level at intervals submerged the ichthyosaur statues, adding a certain realism. Remarkably, internal skeletal structures, such as the scleral rings and the many phalanges of the flippers, were shown at the outside.


title: Ichthyosaur

text: During the nineteenth century, the number of described ichthyosaur genera gradually increased. New finds allowed for a better understanding of their anatomy. Owen had noted that many fossils showed a downward bend in the rear tail. At first, he explained this as a "post mortem" effect, a tendon pulling the tail end downwards after death. However, after an article on the subject by Philip Grey Egerton, Owen considered the possibility that the oblique section could have supported the lower lobe of a tail fin. This hypothesis was confirmed by new finds from Germany. In the Posidonia Shale at Holzmaden, dating from the early Jurassic, already in the early nineteenth century, the first ichthyosaur skeletons had been found. During the latter half of the century, the rate of discovery quickly increased to a few hundred each year. Ultimately, over four thousand were uncovered, forming the bulk of ichthyosaur specimens displayed today. The sites were also a "Konservat-Lagerstätte", meaning not only the quantity, but also the quality was exceptional. The skeletons were very complete and often preserved soft tissues, including tail and dorsal fins. Additionally, female individuals were discovered with embryos.


title: Ichthyosaur

text: In the early twentieth century, ichthyosaur research was dominated by the German paleontologist Friedrich von Huene, who wrote an extensive series of articles, taking advantage of an easy access to the many specimens found in his country. The amount of anatomical data was hereby vastly increased. Von Huene also travelled widely abroad, describing many fossils from locations outside of Europe. During the 20th century, North America became an important source of new fossils. In 1905, the Saurian Expedition led by John Campbell Merriam and financed by Annie Montague Alexander, found twenty-five specimens in central Nevada, which were under a shallow ocean during the Triassic. Several of these are now in the collection of the University of California Museum of Paleontology.


title: Ichthyosaur

text: After a slack during the middle of the century, with no new genera being named between the 1930s and the 1970s, the rate of discoveries picked up towards its end. Other specimens are embedded in the rock and visible at Berlin–Ichthyosaur State Park in Nye County. In 1977 the Triassic ichthyosaur "Shonisaurus" became the state fossil of Nevada. About half of the ichthyosaur genera today seen as valid were described after 1990. In 1992 Canadian paleontologist Elizabeth Nicholls uncovered the largest known specimen, a "Shastasaurus". The new finds have allowed a gradual improvement in knowledge about the anatomy and physiology of what had already been seen as rather advanced "Mesozoic dolphins". Christopher McGowan published a larger number of articles and also brought the group to the attention of the general public. The new method of cladistics provided a means to exactly calculate the relationships between groups of animals, and in 1999, Ryosuke Motani published the first extensive study on ichthyosaur phylogenetics. In 2003, McGowan and Motani published the first modern textbook on the Ichthyosauria and their closest relatives.


title: Ichthyosaur

text: The origin of the ichthyosaurs is contentious. Until recently, clear transitional forms with land-dwelling vertebrate groups had not yet been found, the earliest known species of the ichthyosaur lineage being already fully aquatic. In 2014, a small basal ichthyosauriform from the upper Lower Triassic was described that had been discovered in China with characteristics suggesting an amphibious lifestyle. In 1937, Friedrich von Huene even hypothesised that ichthyosaurs were not reptiles, but instead represented a lineage separately developed from amphibians. Today, this notion has been discarded and a consensus exists that ichthyosaurs are amniote tetrapods, having descended from terrestrial egg-laying amniotes during the late Permian or the earliest Triassic. However, establishing their position within the amniote evolutionary tree has proven difficult, due to their heavily derived morphology obscuring their ancestry. Several conflicting hypotheses have been posited on the subject. In the second half of the 20th century, ichthyosaurs were usually assumed to be of the Anapsida, seen as an early branch of "primitive" reptiles. This would explain the early appearance of ichthyosaurs in the fossil record, and also their lack of clear affinities with other reptile groups, as anapsids were supposed to be little specialised. This hypothesis has become unpopular for being inherently vague because Anapsida is an unnatural, paraphyletic group. Modern exact quantitative cladistic analyses consistently indicate that ichthyosaurs are members of the clade Diapsida. Some studies showed a basal, or low, position in the diapsid tree. More analyses result in their being Neodiapsida, a derived diapsid subgroup.


title: Ichthyosaur

text: Until the 1980s, a close relationship was assumed between the Ichthyosauria and the Sauropterygia, another marine reptile group, within an overarching Euryapsida. Today, however, this is usually rejected, with the Euryapsida being seen as an unnatural polyphyletic assemblage of reptiles that happen to share some adaptations to a swimming lifestyle. An exception is a study in 1997 by John Merck, which showed monophyletic archosauromorph euryapsids.


title: Ichthyosaur

text: Since 1959, a second enigmatic group of ancient sea reptiles is known, the Hupehsuchia. Like the Ichthyopterygia, the Hupehsuchia have pointed snouts and show polydactyly, the possession of more than five fingers or toes. Their limbs more resemble those of land animals, making them appear as a transitional form between these and ichthyosaurs. Initially, this possibility was largely neglected because the Hupehsuchia have a fundamentally different form of propulsion, with an extremely stiffened trunk. The similarities were explained as a case of convergent evolution. Furthermore, the descent of the Hupehsuchia is no less obscure, meaning a possible close relationship would hardly clarify the general evolutionary position of the ichthyosaurs.


title: Ichthyosaur

text: In 2014, "Cartorhynchus" was announced, a small species with a short snout, large flippers, and a stiff trunk. Its lifestyle might have been amphibious. Motani found it to be more basal than the Ichthyopterygia and named an encompassing clade Ichthyosauriformes. The latter group was combined with the Hupesuchia into the Ichthyosauromorpha. The ichthyosauromorphs were found to be diapsids.


title: Ichthyosaur

text: The earliest ichthyosaurs are known from the Early and Early-Middle (Olenekian and Anisian) Triassic strata of Canada, China, Japan, and Spitsbergen in Norway, being up to 246 million years old. These first forms included the genera "Chaohusaurus", "Grippia", and "Utatsusaurus". This diversity suggests an even earlier origin. They more resembled finned lizards than the fishes or dolphins to which the later, more familiar species were similar. Their bodies were elongated and they probably used an anguilliform locomotion, swimming by undulations of the entire trunk. Like land animals, their pectoral girdles and pelves were robustly built, and their vertebrae still possessed the usual interlocking processes to support the body against the force of gravity. However, they were already rather advanced in having limbs that had been completely transformed into flippers. They also were probably warm-blooded and viviparous.


title: Ichthyosaur

text: These very early "proto-ichthyosaurs" had such a distinctive build compared to "ichthyosaurs proper" that Motani excluded them from the Ichthyosauria and placed them in a basal position in a larger clade, the Ichthyopterygia. However, this solution was not adopted by all researchers.


title: Ichthyosaur

text: The basal forms quickly gave rise to ichthyosaurs in the narrow sense sometime around the boundary between the Early Triassic and Middle Triassic; the earliest Ichthyosauria in the sense Motani gave to the concept, appear about 245 million years ago. These later diversified into a variety of forms, including the still sea serpent-like "Cymbospondylus", a problematic form which reached ten metres in length, and smaller, more typical forms like "Mixosaurus". The Mixosauria were already very fish-like with a pointed skull, a shorter trunk, a more vertical tail fin, a dorsal fin, and short flippers containing many phalanges. The sister group of the Mixosauria were the more advanced Merriamosauria. By the Late Triassic, merriamosaurs consisted of both the large, classic Shastasauria and more advanced, "dolphin-like" Euichthyosauria. Experts disagree over whether these represent an evolutionary continuum, with the less specialised shastosaurs a paraphyletic grade that was evolving into the more advanced forms, or whether the two were separate clades that evolved from a common ancestor earlier on. Euichthyosauria possessed more narrow front flippers, with a reduced number of fingers. Basal euichthyosaurs were "Californosaurus" and "Toretocnemus". A more derived branch were the Parvipelvia, with a reduced pelvis, basal forms of which are "Hudsonelpidia" and "Macgowania".


title: Ichthyosaur

text: During the Carnian and Norian, Shastosauria reached huge sizes. "Shonisaurus popularis", known from a number of specimens from the Carnian of Nevada, was long. Norian Shonisauridae are known from both sides of the Pacific. "Himalayasaurus tibetensis" and "Tibetosaurus" (probably a synonym) have been found in Tibet. These large (10- to 15-m-long) ichthyosaurs have by some been placed into the genus "Shonisaurus".


title: Ichthyosaur

text: The gigantic "Shonisaurus sikanniensis" (considered as a shastasaurus between 2011 and 2013) whose remains were found in the Pardonet Formation of British Columbia by Elizabeth Nicholls, has been estimated to be as much as in length—if correct, the largest marine reptile known to date.


title: Ichthyosaur

text: In the Late Triassic, ichthyosaurs attained the peak of their diversity. They occupied many ecological niches. Some were apex predators; others were hunters of small prey. Several species perhaps specialised in suction feeding or were ram feeders; also, durophagous forms are known. Towards the end of the Late Triassic, a decline of variability seems to have occurred. The giant species seemed to have disappeared at the end of the Norian. Rhaetian (latest Triassic) ichthyosaurs are known from England, and these are very similar to those of the Early Jurassic. A possible explanation is an increased competition by sharks, Teleostei, and the first Plesiosauria. Like the dinosaurs, the ichthyosaurs and their contemporaries, the plesiosaurs, survived the Triassic–Jurassic extinction event, and quickly diversified again to fill the vacant ecological niches of the early Jurassic.


title: Ichthyosaur

text: During the Early Jurassic, the ichthyosaurs still showed a large variety of species, ranging from in length. From this epoch, the original British discoveries were made, so that among ichthyosaurs, their names are the ones most familiar to the general public. Genera include "Eurhinosaurus", "Ichthyosaurus", "Leptonectes", "Stenopterygius", and the large predator "Temnodontosaurus", along with the basal parvipelvian "Suevoleviathan", which was little changed from its Norian ancestors. The general morphological variability had been strongly reduced, however. Giant forms, suction feeders and durophagous species were absent. All of these animals were streamlined, dolphin-like forms, although the more basal animals were perhaps more elongated than the advanced and compact "Stenopterygius" and "Ichthyosaurus". The latter belonged to a parvipelvian subgroup, the Thunnosauria. These were fully adapted to an efficient thunniform locomotion, propelling themselves with the end of the tail only, equipped with a vertical tail fin. Their front flippers had more than five fingers. Another parvipelvian branch was the Eurhinosauria such as "Leptonectes" and "Eurhinosaurus", which were specialised forms having very elongated and pointy snouts.


title: Ichthyosaur

text: Few ichthyosaur fossils are known from the Middle Jurassic. This might be a result of the poor fossil record in general of this epoch. The strata of the Late Jurassic seem to indicate that a further decrease in diversity had taken place. From the Middle Jurassic onwards, almost all ichthyosaurs belonged to the thunnosaurian clade Ophthalmosauridae. Represented by the "Ophthalmosaurus" and related genera, they were very similar in general build to "Ichthyosaurus". The eyes of "Ophthalmosaurus" were huge, and these animals likely hunted in dim and deep water. However, new finds from the Cretaceous indicate that ichthyosaur diversity in the Late Jurassic must have been underestimated.


title: Ichthyosaur

text: Traditionally, ichthyosaurs were seen as decreasing in diversity even further with the Cretaceous, though they had a worldwide distribution. All fossils from this period were referred to a single genus: "Platypterygius". This last ichthyosaur genus was thought to have become extinct early in the late Cretaceous, during the Cenomanian about 95 million years ago. The ichthyosaurs thus would have disappeared much earlier than other large Mesozoic reptile groups that generally survived until the end of the Cretaceous. Two major explanations were given for this. Firstly, it could have been a matter of chance. The second explanation had the extinction as caused by competition. Less hydrodynamically efficient animals, such as the Mosasauridae and long-necked plesiosaurs, flourished. The ichthyosaurian overspecialisation could be a contributing factor to their extinction, possibly being unable to 'keep up' with the fast-swimming and highly evasive new teleost fish, which had become dominant at this time, against which the sit-and-wait ambush strategies of the mosasaurids proved superior. This model thus emphasised evolutionary stagnation, the only innovation shown by "Platypterygius" being its ten fingers.


title: Ichthyosaur

text: Recent studies, however, show that ichthyosaurs were actually far more diverse in the Cretaceous than previously thought. Fragments referred to "Platypterygius" in fact represented diverse species. In 2012, at least eight lineages spanned the Jurassic-Cretaceous boundary, among them "Acamptonectes", "Sveltonectes", "Caypullisaurus", and "Maiaspondylus". In 2013, a Cretaceous basal thunnosaurian was revealed: "Malawania". Indeed, likely a radiation during the Early Cretaceous occurred due to an increase of coastlines when the continents further broke up.


title: Ichthyosaur

text: The demise of the ichthyosaurs has recently been described as a two-step process. A first extinction event in the beginning of the Cenomanian eliminated two of the three ichthyosaur feeding guilds then present, the 'soft-prey specialists' and the 'generalists', leaving only an apex predator group, which probably was not particularly specialized. The second major ichthyosaur extinction took place during the Cenomanian-Turonian boundary event, an 'anoxic event', eliminating the apex predators, after which just a single lineage survived, "Platypterygius hercynicus", which disappeared shortly afterwards, about 93 million years ago. Ichthyosaur extinction was thus a pair of abrupt events rather than a long decline, probably related to the environmental upheavals and climatic changes in the Cenomanian and Turonian. If so, it was not caused by mosasaurid competition; large mosasaurs did not appear until 3 million years after the ichthyosaur extinction, likely to fill the resulting ecological void. Plesiosaurian polycoltylids perhaps also filled some of the niches previously occupied by ichthyosaurs, although they had coexisted for 19 million years. The extinction was most likely the result of ecological change and volatility that caused changes in migration, food availability, and birthing grounds. This part of the Cretaceous was one in which many other marine extinctions occurred, including those of some types of microplankton, ammonites, belemnites, and reef-building bivalves.


title: Ichthyosaur

text: This taxonomy was presented by Michael Werner Maisch in his 2010 review of ichthyosaur classification.


title: Ichthyosaur

text: In modern phylogeny, clades are defined that contain all species forming a certain branch of the evolutionary tree. This also allows one to clearly indicate all relationships between the several subgroups in a cladogram. In 1999, a node clade Ichthyopterygia was defined by Motani as the group consisting of the last common ancestor of "Ichthyosaurus communis", "Utatsusaurus hataii" and "Parvinatator wapitiensis"; and all its descendants. Within Motani's phylogeny, the Ichthyopterygia were the larger parent clade of a smaller stem clade Ichthyosauria that was defined as the group consisting of "Ichthyosaurus communis" and all species more closely related to "Ichthyosaurus" than to "Grippia longirostris". Motani's concept of the Ichthyosauria was thus more limited than the traditional one that also contained basal forms, such as "Grippia", "Utatsusaurus", and "Parvinatator".


title: Ichthyosaur

text: An alternative terminology was proposed by Maisch & Matzke in 2000, trying to preserve the traditional, more encompassing content of the concept Ichthyosauria. They defined a node clade Ichthyosauria as the group consisting of the last common ancestor of "Thaisaurus chonglakmanii", "Utatsusaurus hataii", and "Ophthalmosaurus icenicus", and all its descendants. Ichthyosauria "sensu" Motani might materially be identical to a clade that Maisch & Matzke in 2000 called Hueneosauria, depending on the actual relationships.


title: Ichthyosaur

text: Cladogram based on Maisch and Matzke (2000) and Maisch and Matzke (2003) with clade names following Maisch (2010):


title: Ichthyosaur

text: Ichthyosaurs averaged about in length. Some individual specimens were as short as ; some species were much larger: the Triassic "Shonisaurus popularis" was about long and in 2004 "Shonisaurus sikanniensis" (classified as a shastasaurus between 2011 and 2013) was estimated to have been in length. Fragmentary finds suggest the presence of a form in the early Jurassic. In 2018, lower jaw fragments from England were reported indicating a length of between 20 and 25 m (66 to 82 ft). According to weight estimates by Ryosuke Motani a "Stenopterygius" weighed around , whilst a "Ophthalmosaurus icenicus" weighed .


title: Ichthyosaur

text: While the earliest known members of the ichthyosaur lineage were more eel-like in build, later ichthyosaurs resembled more typical fishes or dolphins, having a porpoise-like head with a short neck and a long snout. Ichthyosaur fore and hind limbs had been fully transformed into flippers. Some species had a fin on their backs and a more or less vertical fin at the rear of a rather short tail. Although ichthyosaurs looked like fish, they were not.Evolutionary biologist Stephen Jay Gould said that the ichthyosaur was his favourite example of convergent evolution, where similarities of structure are analogous, not homologous, thus not caused by a common descent, but by a similar adaptation to an identical environment:


title: Ichthyosaur

text: This sea-going reptile with terrestrial ancestors converged so strongly on fishes that it actually evolved a dorsal fin and tail in just the right place and with just the right hydrological design. These structures are all the more remarkable because they evolved from nothing—the ancestral terrestrial reptile had no hump on its back or blade on its tail to serve as a precursor.


title: Ichthyosaur

text: Derived ichthyosaurs in the narrow sense, as defined by Motani in 1999, differ from their closest basal ichthyopterygian relatives in certain traits. Motani listed a number of these. The external nostril is located on the side of the skull, and is hardly visible from above. The upper rim of the eye socket consists of a bone bar formed by the prefrontal and the postfrontal bones. The postorbital in side view is excluded from the supratemporal fenestra. The opening for the parietal eye is located on the border of the parietal and the frontal bone. The lateral wing of the pterygoid is incompletely and variably ossified. The ulna lacks the part behind the original shaft axis. The rear dorsal vertebrae are disc-shaped.


title: Ichthyosaur

text: Basal Ichthyopterygia already had elongated, triangular skulls. With ichthyosaurs in the narrow sense, their snouts became very pointy. The snout is formed by the premaxilla. The maxilla behind it is usually shorter and sometimes excluded from the external nostril by the rear branch of the premaxilla. Accordingly, the number of premaxillary teeth is high, while the maxillary teeth are fewer in number or even completely absent. The rear top of the snout is formed by the nasal bones. Derived species have a foramen internasale, a midline opening separating the rear of the nasal bones. The nasal bone usually forms the top and front rim of the bony nostril, itself often placed just in front of the eye socket. However, with some Triassic species, the premaxilla is so strongly extended at its back that it even excludes the nasal from the nostril.


title: Ichthyosaur

text: The rear of the skull is dominated by a large eye socket, often covering the major part of the rear side surface. In the socket, a large scleral ring is present; this is a circular structure of small, overlapping bone segments protecting the eye against the water pressure. Both in the relative and absolute senses, ichthyosaurs have the largest eye sockets of all known vertebrates. The other rear skull elements are typically so compressed and fused that they are difficult to identify. The top rear element of the skull was usually assumed to be the supratemporal bone, while the squamosal and quadratojugal were sometimes fused. However, in 1968, Alfred Sherwood Romer stated that the presumed supratemporal was in fact the squamosal, an interpretation which was supported by McGowan in 1973. In 1990, though, John Steve Massare convinced most researchers that the original identification had been the correct one after all. The supratemporal forms the rear rim of the supratemporal opening; a lower temporal opening at the side is lacking. The front rim of the supratemporal opening is typically formed by the postfrontal; only with the very basal "Utatsusaurus" the postorbital and the squamosal still reach the edge. Between the paired supratemporal openings, the skull roof is narrow; some species have a longitudinal crest on it as an attachment for the jaw muscles. Basal Ichthyopterygia have a parietal eye opening between the paired parietal bones. With ichthyosaurs proper, this opening moves to the front, first to the border between the parietals and the frontals and ultimately between the frontals, a condition shown by derived species. Postparietal and tabular bones are lacking. Often, the bones of the back of the skull and the palate are incompletely ossified, apparently having partly remained cartilage. The occipital condyle is typically very convex. The stapes, the bone transmitting sound waves from the eardrum to the middle ear, is elongated and not pierced by a foramen. Pterygoid teeth are typically lacking.


title: Ichthyosaur

text: Like the snout, the lower jaws are elongated. However, in some species, such as "Eurhinosaurus" and "Excalibosaurus", the front of the snout far protrudes beyond the lower jaws. While the front of the lower jaw is typically low, its rear depth is very variable. The greater part of the lower jaw is formed by the front dentary, the tooth-bearing bone. At its inner side the dentary is covered by a splenial that extends forwards until the symphysis, the common contact surface where both lower jaws are grown together. The jaw joints do not allow a horizontal chewing movement: they function as simple hinges to vertically open or close the jaws.


title: Ichthyosaur

text: Ichthyosaur teeth are typically conical. Fish-eating species have long and slender tooth crowns that are slightly recurved. Forms specialised in catching larger prey have shorter, broader, and straighter teeth; sometimes, cutting edges are present. "Thalattoarchon", an apex predator, had larger teeth formed like flattened blades. Durophagous species that ate shellfish have low, convex teeth that are closely packed. Many ichthyosaur dentitions are heterodont, combining several tooth shapes, e.g. small teeth in the front and larger teeth at the rear. The teeth are usually placed in tooth sockets; derived species possess a common tooth groove. In the latter case, adult individuals sometimes become toothless. Teeth in tooth sockets sometimes fuse with the jawbone. With ichthyosaur teeth, the dentine shows prominent vertical wrinkles. Durophagous forms have teeth with deep vertical grooves and wrinkles in the enamel.


title: Ichthyosaur

text: Basal Ichthyopterygia, like their land-dwelling ancestors, still had vertebrae that possessed a full set of processes that allowed them to interlock and articulate, forming a vertebral column supporting the weight of the body. As ichthyosaurs were fully aquatic, their bodies were supported by the Archimedes force exerted by the water; in other words, they were buoyant. Therefore, the vertebral processes had lost much of their function. Early ichthyosaurs proper had rear dorsal vertebrae that had become disc-shaped, like those of typical fishes. With more derived species, the front dorsals also became discs. Gradually, most processes were lost, including those for rib attachment. The vertebral bodies became much shorter. The front and rear sides of the discs were hollowed out, resulting in a so-called amphicoelous condition. A transverse cross-section of such a vertebra has an hourglass shape. This morphology is unique within the Amniota and makes discerning ichthyosaur vertebrae from those of other marine reptiles easy. The only process that kept its function was the spine at the top, serving as an attachment for the dorsal muscles. However, even the spine became a simple structure. The neural arch, of which it was an outgrowth, typically no longer fused to the vertebral centre.


title: Ichthyosaur

text: The neck is short, and derived species show a reduction in the number of cervical vertebrae. The short neck positions the skull close to the trunk, usually in a slight oblique elevation to it. Derived species usually also have a reduced number of dorsals, the total of presacral vertebrae totalling about forty to fifty. The vertebral column is little differentiated. Basal Ichthyopterygia still have two sacral vertebrae, but these are not fused. Early Triassic forms have a transversely flattened tail base with high spines for an undulating tail movement. Derived forms have a shorter tail with the characteristic kink at the end; a section of wedge-shaped vertebrae, itself supporting the fleshy upper tail fin lobe, forced the tail end into the lower fin lobe.


title: Ichthyosaur

text: As derived species no longer have transversal processes on their vertebrae—again a condition unique in the Amniota—the parapophyseal and diapophysael rib joints have been reduced to flat facets, at least one of which is located on the vertebral body. The number of facets can be one or two; their profile can be circular or oval. Their shape often differs according to the position of the vertebra within the column. The presence of two facets per side does not imply that the rib itself is double-headed: often, even in that case, it has a single head. The ribs typically are very thin and possess a longitudinal groove on both the inner and the outer sides. The lower side of the chest is formed by gastralia. These belly ribs have a single centre segment and one or two outer segments per side. They are not fused into a real plastron. Usually two gastralia are present per dorsal rib.


title: Ichthyosaur

text: The shoulder girdle of ichthyosaurs is not much modified from its original condition. Some basal forms show a hatchet- or crescent-shaped shoulder blade or scapula; derived forms have an elongated blade positioned on a broader base. The scapula is not fused with the coracoid into a scapulocoracoid, indicating that the forces exerted on the shoulder girdle were moderate. The shoulder joint is positioned on the border between the scapula and the coracoid. Both coracoids are fused on their common midline. The coracoid shape is very variable, but usually it is rather low. The upper part of the shoulder girdle is formed by two long and slender clavicles, crowned by a central interclavicular bone that is large and triangular with basal forms, small and T-shaped in Jurassic species. Breast bones or sterna are absent.


title: Ichthyosaur

text: Basal forms have a forelimb that is still functionally differentiated, in some details resembling the arm of their land-dwelling forebears; the ulna and radius are elongated and somewhat separated; the carpals are rounded, allowing the wrist to rotate; the number of phalanges is within the range shown by land animals. Ichthyosaurs proper, to the contrary, have a forelimb that is fully adapted to its function as a flipper. However, the adaptations are very variable. Triassic species typically have a very derived humerus, changed into a disc. Jurassic species tend to have a more elongated humeral form with a rounded head, narrow shaft, and expanded lower end. The radius and ulna are always strongly flattened, but can be circular, with or without notch, or have a waist. Notches can be homologous to the original shafts, but also be newly formed. Jurassic forms no longer have a space, the "spatium interosseum", between the radius and ulna. Often, the latter bones gradually merge into lower, disc-shaped elements - the up to four carpals which again differ little in form from the up to five metacarpals.


title: Ichthyosaur

text: A strongly derived condition show the phalanges, small, disc-shaped elements positioned in long rows. Sometimes, the number of fingers is reduced, to as low as two. This is a rather common phenomenon within the Tetrapoda. Unique, however, for derived tetrapods, is the fact that some species show nonpathological polydactyly, the number of fingers being higher than five. Some species have ten fingers per hand. These fingers, again, can have an increased number of phalanges, up to thirty, a phenomenon called hyperphalangy, also known from the Plesiosauria, mosasaurs, and the Cetacea. The high number of elements allows the flipper to be shaped as a hydrofoil. When a high number of fingers is present, their identity is difficult to determine. It is usually assumed that fingers were added at both the front and at the rear, perhaps to a core of four original fingers. If fingers are added, often the number of metacarpals and carpals is also increased; sometimes even an extra lower arm element is present. Earlier, ichthyosaurs were commonly divided into "longipinnate" and "latipinnate" forms, according to the long or wide shape of the front flippers, but recent research has shown that these are not natural groups; ichthyosaur clades often contain species with and without elongated forelimbs.


title: Ichthyosaur

text: The ichthyosaur pelvis is typically rather reduced. The three pelvic bones: the ilium, the ischium, and the pubic bone, are not fused and often do not even touch each other. Also, the left and right pelvic sides no longer touch; only basal forms still have sacral ribs connecting the ilia to the vertebral column. The hip joint is not closed on the inside. The pubic bone typically does not connect to the ischium behind it; the space in between is by some workers identified as the "fenestra thyreoidea"; other researchers deny that the term is applicable given the general loose structure of the pelvis. Some later species have a connected pubic bone and ischium, but in this case, the femoral head no longer articulates with the hip joint. Triassic species have plate-like pubic bones and ischia; in later species these elements become elongated with a narrow shaft and can form a single rod.


title: Ichthyosaur

text: Typically, the hindlimbs are shorter than the forelimbs, possessing a lesser number of elements. Often, the rear flipper is only half the length of the front flipper. The thighbone is short and broad, often with a narrow waist and an expanded lower end. The tibia, fibula and metatarsals are merged into a mosaic of bone discs supporting the hydrofoil. Three to six toes are present. The toe phalanges also show hyperphalangy; exceptionally, "Ophthalmosaurus" shows a reduced number of phalanges.


title: Ichthyosaur

text: The earliest reconstructions of ichthyosaurs all omitted dorsal fins and caudal (tail) flukes, which were not supported by any hard skeletal structure, so were not preserved in many fossils. Only the lower tail lobe is supported by the vertebral column. In the early 1880s, the first body outlines of ichthyosaurs were discovered. In 1881, Richard Owen reported ichthyosaur body outlines showing tail flukes from Lower Jurassic rocks in Barrow-upon-Soar, England. Other well-preserved specimens have since shown that in some more primitive ichthyosaurs, like a specimen of "Chaohusaurus geishanensis", the tail fluke was weakly developed and only had a dorsal tail lobe, making the tail more paddle-like. Over the years, the visibility of the tail lobe has faded away in this specimen.


title: Ichthyosaur

text: The presence of dorsal fins in ichthyosaurs has been controversial. Finely preserved specimens from the Holzmaden "Lagerstätten" in Germany found in the late 19th century revealed additional traces, usually preserved in black, of the outline of the entire body, including the first evidence of dorsal fins in ichthyosaurs. Unique conditions permitted the preservation of these outlines, which probably consist of bacterial mats, not the remains of the original tissues themselves. In 1987, David Martill argued that, given the indirect method of conservation by bacteria, these outlines were unlikely to have been reliably preserved in any fine detail. He concluded that no authentic dorsal fins had been discovered. After displaced skins flaps from the body would have initially been misinterpreted as fins, fossil preparators later came to expect such fins to be present, and would have identified any discolouration in the appropriate position as a dorsal fin or even have falsified such structures. The lack of a dorsal fin would also explain why ichthyosaurs, contrary to porpoises, retained hind flippers, as these were needed for stability. Other researchers noted that, while the outlines might have been sharpened and smoothed by preparators because fossil bacterial mats usually have indistinct edges, many of the preserved dorsal fins were probably authentic and at least somewhat close to the true body outline. At least one specimen, R158 (in the collections of the Paleontologiska Museet, Uppsala University), shows the expected faded edges of a bacterial mat, so it has not been altered by preparators, yet still preserves a generally tuna-like body outline including a dorsal fin. In 1993, Martill admitted that at least some dorsal fin specimens are authentic.


title: Ichthyosaur

text: The fossil specimens that preserved dorsal fins also showed that the flippers were pointy and often far wider than the underlying bones would suggest. The fins were supported by fibrous tissue. In some specimens, four layers of collagen are visible, the fibres of the covering layers crossing those of the collagen below.


title: Ichthyosaur

text: In 2017, from the German Posidonia Shale the discovery was reported of 182.7-million-year-old vertebrae of "Stenopterygius" in a carbonate nodule, still containing collagen fibers, cholesterol, platelets, and red and white blood cells. The structures would not have been petrified, but represent the original organic tissues of which the biomolecules could be identified. The exceptional preservation was explained by the protective environment offered by the nodule. The red blood cells found, were one-fourth to one fifth the size of those of modern mammals. This would have been an adaptation for an improved oxygen absorption, also in view of the low oxygen levels during the Toarcian. The cholesterol had a high-carbon-13 isotope component which might indicate a higher position in the food chain and a diet of fish and cephalopods.


title: Ichthyosaur

text: Typically, fossils that preserve it suggest that the skin of ichthyosaurs was smooth and elastic, lacking scales. However, these remains are not impressions "per se", but outlines formed from bacterial growth. In one case, a true impression of the skin was reported from a specimen of "Aegirosaurus" found in the Solnhofen Plattenkalk, rocks which were capable of preserving even the finest detail. Minuscule scales seemed to be visible in this specimen.


title: Ichthyosaur

text: The colouration of ichthyosaurs is difficult to determine. In 1956, Mary Whitear reported finding melanocytes, pigment cells in which reddish-brown pigment granules would still be present, in a skin specimen of a British fossil, R 509. Ichthyosaurs are traditionally assumed to have employed countershading (dark on top, light at the bottom) like sharks, penguins, and other modern animals, serving as camouflage during hunting. This was contradicted in 2014 by the discovery of melanosomes, black melanin-bearing structures, in the skin of ichthyosaur specimen YORYM 1993.338 by Johan Lindgren of Lund University. It was concluded that ichthyosaurs were likely uniformly dark coloured for thermoregulation and to camouflage them in deep water while hunting. This is in contrast to mosasaurids and prehistoric leatherback turtles, which were found to be countershaded. However, a 2015 study doubted Lindgren's interpretation. This study noted that a basal layer of melanosomes in the skin is ubiquitous in reptile coloration, but does not necessarily correspond to a dark appearance. Other chromatophore structures (such as iridiophores, xanthophores, and erythrophores) affect coloration in extant reptiles but are rarely preserved or identified in fossils. Thus, due to the unknown presence of these chromatophores, YORYM 1993.338, could have been countershaded, green, or various other colors or patterns.


title: Ichthyosaur

text: Gastroliths, stomach stones that might have assisted digestion or regulated buoyancy, have only on a few occasions been found associated with ichthyosaur skeletons, once with a specimen of "Nannopterygius" and a second time in a "Panjiangsaurus" fossil. Ichthyosaur coproliths, petrified faeces, are very common, though, already being sold by Mary Anning.


title: Ichthyosaur

text: Apart from the obvious similarities to fish, ichthyosaurs also shared parallel developmental features with dolphins, lamnid sharks, and tuna. This gave them a broadly similar appearance, possibly implied similar activity levels (including thermoregulation), and presumably placed them broadly in a similar ecological niche. Ichthyosaurs were not primarily coastal animals; they also inhabited the open ocean. They have been found in all Mesozoic oceans. This is even true of the earliest Ichthyopterygia, making identification of a certain area as their place of origin impossible.


title: Ichthyosaur

text: Ichthyosaurs were carnivorous; they ranged so widely in size, and survived for so long, that they are likely to have had a wide range of prey. Species with pointed snouts were adapted to grab smaller animals. McGowan speculated that forms with protruding upper jaws, in the Eurhinosauria, would have used their pointy snouts to slash prey, as has been assumed for swordfish. The most commonly preserved gut contents in ichthyosaurs are the remains of cephalopods. Less commonly, they fed on fish and other vertebrates, including smaller ichthyosaurs. The large Triassic form "Thalattoarchon" had large, bladed teeth and was probably a macropredator, capable of killing prey its own size, and "Himalayasaurus" and several species of "Temnodontosaurus" also shared adaptations for killing very large prey. These food preferences have been confirmed by coproliths which indeed contain the remains of fishes and cephalopods. Another confirmation is provided by fossilised stomach contents. Buckland in 1835 described the presence in a specimen of a large mass of partly digested fishes, recognisable by their scales. Subsequent research in 1968 determined that these belonged to the fish genus "Pholidophorus", but also that cephalopod beaks and sucker hooks were present. Such hard food particles apparently were retained by the stomach and regularly regurgitated. Carcasses of drowned animals were eaten as well: in 2003 a specimen of "Platypterygius longmani" was reported having besides fishes and a turtle the bones of a land bird in its stomach.


title: Ichthyosaur

text: Some early ichthyosaurs were durophagous and had flat convex teeth adapted for crushing shellfish. They thus ate benthos from the floor of shallow seas. Other species were perhaps suction feeders, sucking animals into their mouths by quickly opening their relatively short jaws. This was first assumed for "Shonisaurus", which giant by this means might have secured a constant food supply for its huge body, and in 2011 for the short-snouted "Guanlingsaurus liangae". However, in 2013 a study concluded that the hyoid bone of ichthyosaurs, at the tongue base, was insufficiently ossified to support a suction feeding movement and suggested the alternative that such species were ram feeders, gathering food by constantly swimming forwards with a wide-open mouth.


title: Ichthyosaur

text: Typical ichthyosaurs had very large eyes, protected within a bony ring, suggesting that they may have hunted at night or at great depths; the only extant animals with similarly large eyes are the giant and colossal squids. Sight thus seems to have been one of the main senses employed while hunting. Hearing might have been poor, given the very robust form of the stapes. Grooves in the palate however, suggest that smell might have been acute or even that electro-sensory organs might have been present.


title: Ichthyosaur

text: Ichthyosaurs themselves served as food for other animals. During the Triassic their natural predators mainly consisted of sharks and other ichthyosaurs; in the Jurassic these were joined by large Plesiosauria and marine Crocodylomorpha. This is again confirmed by stomach contents: in 2009 e.g., a plesiosaur specimen was reported with an ichthyosaur embryo in its gut.


title: Ichthyosaur

text: In ichthyosaurs, the main propulsion was provided by a lateral movement of the body. Early forms employed an anguilliform or eel-like movement, with undulations of the entire trunk and tail. This is usually considered rather inefficient. Later forms, like the Parvipelvia, has a shorter trunk and tail and probably used a more efficient carangiform or even thunniform movement, in which the last third of the body, respectively, the tail end, is flexed only. The trunk in such species is rather stiff.


title: Ichthyosaur

text: The tail was bi-lobed, with the lower lobe being supported by the caudal vertebral column, which was "kinked" ventrally to follow the contours of the ventral lobe. Basal species had a rather asymmetric or "heterocercal" tail fin. The asymmetry differed from that of sharks in that the lower lobe was largest, instead of the upper lobe. More derived forms had a nearly vertical symmetric tail fin. Sharks use their asymmetric tail fin to compensate for the fact that they are negatively buoyant, heavier than water, by making the downward pressure exerted by the tail force the body as a whole in an ascending angle. This way, swimming forwards will generate enough lift to equal the sinking force caused by their weight. In 1973, McGowan concluded that, because ichthyosaurs have a reversed tail fin asymmetry compared to sharks, they were apparently positively buoyant, lighter than water, which would be confirmed by their lack of gastroliths and of pachyostosis or dense bone. The tail would have served to keep the body in a descending angle. The front flippers would be used to push the front of the body further downwards and control pitch. In 1987 however, Michael A. Taylor suggested an alternative hypothesis: as ichthyosaurs could vary their lung content, contrary to sharks (which lack a swimming bladder), they could also regulate their buoyancy. The tail thus mainly served for a neutral propulsion, while small variations in buoyancy were stabilised by slight changes in the flipper angles. In 1992, McGowan accepted this view, pointing out that shark tails are not a good analogy of derived ichthyosaur tails that have more narrow lobes, and are more vertical and symmetric. Derived ichthyosaur tail fins are more like those of tuna fish and indicate a comparable capacity to sustain a high cruising speed. A comparative study by Motani in 2002 concluded that, in extant animals, small tail fin lobes positively correlate with a high beat frequency. Modern researchers generally concur that ichthyosaurs were negatively buoyant.


title: Ichthyosaur

text: In 1994, Judy Massare concluded that ichthyosaurs had been the fastest marine reptiles. Their length/depth ratio was between three and five, the optimal number to minimise water resistance or drag. Their smooth skin and streamlined bodies prevented excessive turbulence. Their hydrodynamic efficiency, the degree to which energy is converted into a forward movement, would approach that of dolphins and measure about 0.8. Ichthyosaurs would be a fifth faster than plesiosaurs, though half of the difference was explained by assuming a 30% higher metabolism for ichthyosaurs. Together, within Massare's model these effects resulted in a cruising speed of slightly less than five kilometres per hour. However, in 2002, Motani corrected certain mistakes in Massare's formulae and revised the estimated cruising speed to less than two kilometres per hour, somewhat below that of modern Cetacea. However, as the speeds estimated for plesiosaurs and mosasaurids were also revised downwards, ichthyosaurs maintained their relative position.


title: Ichthyosaur

text: Ichthyosaurs had fin-like limbs of varying relative length. The standard interpretation is that these, together with the dorsal fin and tail fin, were used as control surfaces for directional stability, controlling yaw, and for stabilising pitch and roll, rather than propulsion. However, during the 1980s the German paleontologist Jürgen Riess proposed an alternative model. After having studied the flying movement made by the forelimbs of plesiosaurs, he suggested that at least those ichthyosaurs that had long flippers used them for a powerful propulsive stroke, moving them up and down. This would explain the non-degenerated shoulder girdle and the evolution of the hand bones, whose perfect hydrofoil profile would have been useless if it was not functionally employed. He thought to have discovered modern analogues in the Queensland lungfish and the Amazon river dolphin, which he presumed also used their long fins for propulsion. Riess expounded upon this hypothesis in a series of articles. This alternative interpretation was generally not adopted by other workers. In 1998, Darren Naish pointed out that the lungfish and the river dolphin actually do not use their fins in this way and that e.g. the modern humpback whale has very long front flippers, supported by a mosaic of bones, but that these nevertheless mainly serve as rudders. In 2013, a study concluded that broad ichthyosaur flippers, like those of "Platyptergygius", were not used for propulsion but as a control surface.


title: Ichthyosaur

text: Many extant lung-breathing marine vertebrates are capable of deep diving. There are some indications about the diving capacity of ichthyosaurs. Quickly ascending from a greater depth can cause decompression sickness. The resulting bone necrosis has been well documented with Jurassic and Cretaceous ichthyosaurs, where it is present in 15% and 18% of specimens, respectively, but is rare in Triassic species. This could be a sign that basal forms did not dive as deeply, but might also be explained by a greater predation pressure during the later epochs, more often necessitating a fast flight to the surface. However, this last possibility is contradicted by the fact that, with modern animals, damage is not caused by a limited number of rapid ascension incidents, but by a gradual accumulation of non-invalidating degeneration during normal diving behaviour.


title: Ichthyosaur

text: Additional evidence is provided by the eyes of ichthyosaurs that among vertebrates are both relatively and absolutely the largest known. Modern leopard seals can dive to up to hunting on sight. Motani suggested that ichthyosaurs, with their relatively much larger eye sockets, should have been able to reach even greater depths. "Temnodontosaurus", with eyes that had a diameter of twenty-five centimetres, could probably still see at a depth of 1,600 metres. At these depths, such eyes would have been especially useful to see large objects. Later species, such as "Ophthalmosaurus", had relatively larger eyes, again an indication that diving capacity was better in late Jurassic and Cretaceous forms.


title: Ichthyosaur

text: Similar to modern cetaceans, such as whales and dolphins, ichthyosaurs were air-breathing. Whales and dolphins are mammals and warm-blooded. Of ichthyosaurs it was traditionally assumed that they were cold-blooded, being reptiles. However, since the 1970s many dominant reptile groups of the Mesozoic, such as theropod dinosaurs, pterosaurs and plesiosaurs, have been considered warm-blooded, as this offers an elegant explanation of their dominance. Some direct evidence is available that ichthyosaurs too might have been endothermic. In 1990, Vivian de Buffrénil published a histological study, indicating that ichthyosaurs possessed a fibrolamellar bone structure, as with warm-blooded animals in general, typified by fast growth and a strong vascularisation. Early Triassic species already show these traits. In 2012, it was reported that even the very basal form "Utatsusaurus" had this bone type, indicating that the ancestors of ichthyosaurs were already warm-blooded. Additional direct proof for a high metabolism is the isotopes of oxygen ratio in the teeth, which indicates a body temperature of between 35 and 39 °C, about 20° higher than the surrounding seawater. Blubber is consistent with warm-bloodedness as the insulating qualities require the animal to generate its own heat.


title: Ichthyosaur

text: Indirect evidence for endothermy is provided by the body shape of derived ichthyosaurs, which with its short tail and vertical tail fin seems optimised for a high cruising speed that can only be sustained by a high metabolism: all extant animals swimming this way are either fully warm-blooded or, like sharks and tuna, maintain a high temperature in their body core. This argument does not cover basal forms with a more eel-like body and undulating swimming movement. In 1996, Richard Cowen, while accepting endothermy for the group, presumed that ichthyosaurs would have been subject to "Carrier's constraint", a limitation to reptilian respiration pointed out in 1987 by David Carrier: their undulated locomotion forces the air out of the lungs and thus prevents them from taking breath while moving. Cowen hypothesised that ichthyosaurs would have overcome this problem by porpoising: constantly jumping out of the water would have allowed them to take a gulp of fresh air during each jump. Other researchers have tended to assume that for at least derived ichthyosaurs Carrier's constraint did not apply, because of their stiff bodies, which seems to be confirmed by their good diving capacity, implying an effective respiration and oxygen storage system. For these species porpoising was not a necessity. Nevertheless, ichthyosaurs would have often surfaced to breathe, probably tilting their heads slightly to take in air, because of the lower position of the nostrils compared to that of dolphins.


title: Ichthyosaur

text: Ichthyosaurs were viviparous, i.e. bore live young instead of laying eggs. Although they were reptiles and descended from egg-laying, oviparous, ancestors, viviparity is not as unexpected as it first appears. Air-breathing marine creatures must either come ashore to lay eggs, like turtles and some sea snakes, or else give birth to live young in surface waters, like whales and dolphins. Given their streamlined and transversely flattened bodies, heavily adapted for fast swimming, it would have been difficult, if not impossible, for ichthyosaurs to move far enough on land to lay eggs. This was confirmed as early as 9 December 1845 when naturalist Joseph Chaning Pearce reported a small embryo in a fossil of "Ichthyosaurus communis". The embryo, with a length of eleven centimetres, was positioned in the birth canal of its two-and-a-half metre long mother, with its head pointed to the rear. Pearce concluded from the fossil that ichthyosaurs had to have been viviparous.


title: Ichthyosaur

text: Later, from the Holzmaden deposits numerous adult fossils were found containing fetuses. In 1880, Harry Govier Seeley, heading a special British paleontological committee studying the problem of ichthyosaur reproduction, concluded that birth was given in the water and that fossils containing fetuses in the birth canal probably represented cases of premature death of the juvenile, causing the demise of the mother animal as well. A comparison has been made with dolphins and whales, whose young need to be born tail-first to prevent drowning; if the juvenile is born head-first, it dies and the mother with it if the corpse gets stuck in the birth canal. However, an alternative explanation is that such fossils actually represent females that had died for other reasons while pregnant, after which the decomposition gasses drove out the fetuses head-first. In 2014, a study reported the find of a fossilized "Chaohusaurus" female that had died while giving birth to three neonates. Two had already been expelled while a third was present in the birth canal. The fossil also documented that early ichthyosaurs were also born head first, perhaps opposed to later genera. As "Chaohusaurus" is a very basal ichthypterygian—previously, the most basal genus of which fetuses were known, had been "Mixosaurus"—this discovery suggests that the earliest land-dwelling ancestors of ichthyosaurs had already been viviparous.


title: Ichthyosaur

text: Compared with placental mammals or plesiosaurs, ichthyosaur fetuses tend to be very small and their number per litter is often high. In one female of "Stenopterygius" seven have been identified, in another eleven. The fetuses have at most a quarter of the length of the mother animal. The juveniles have about the same body proportions as adult individuals. The main ontogenetical changes during growth consist in the fusion and greater robustness of the skeletal elements.


title: Ichthyosaur

text: Crocodiles, most sea turtles and some lizards control offspring gender by manipulating the temperature of the developing eggs' environment; i.e. they do not have distinct sex chromosomes. Live-bearing reptiles do not regulate sex through incubation temperature. A study in 2009, which examined 94 living species of reptiles, birds and mammals, found that the genetic control of sex appears to be crucial to live birth. It was concluded that with marine reptiles such control predated viviparity and was an adaptation to the stable sea-climate in coastal regions. Genetics likely controlled sex in ichthyosaurs, mosasaurs and plesiosaurs.


title: Ichthyosaur

text: Ichthyosaurs are often assumed to have lived in herds or hunting groups. Little evidence is available about the nature of ichthyosaur social behaviour. Some indications exist that a level of sexual dimorphism was present. Skeletons of "Eurhinosaurus" and "Shastasaurus" show two morphotypes. Individuals with a longer snout, larger eyes, a longer trunk, a shorter tail, and longer flippers with additional phalanges, could have represented the females; the longer trunk may have provided room for the embryos.


title: Ichthyosaur

text: Generally, the brain shows the limited size and elongated shape of that of modern cold-blooded reptiles. However, in 1973, McGowan, while studying the natural endocast of a well-preserved specimen, pointed out that the telencephalon was not very small. The visual lobes were large, as could be expected from the eye size. The olfactory lobes were, though not especially large, well-differentiated; the same was true of the cerebellum.


title: Ichthyosaur

text: Though fossils revealing ichthyosaur behavior remain rare, one ichthyosaur fossil is known to have sustained bites to the snout region. Discovered in Australia, and analyzed by Benjamin Kear "et alii" in 2011, measurements of the wounds reveal that the bite marks were inflicted by another ichthyosaur, likely of the same species, a probable case of face biting during a conflict. The wounds show signs of healing in the form of bone growth, meaning that the victim survived the attack. Another, very large ichthyosaur close to nine metres in length was found in Svalbard; it was nearly complete save for its tail. Scrutiny of the find revealed that while hunting ammonites (as evidenced by an ammonite shell in the throat region), the ichthyosaur was ambushed and attacked, likely by a pliosaurid (known from the same habitat), which severed its tail. The ichthyosaur then sank to the depths, drowning and eventually becoming fossilized in the deep water. The find was revealed to the public in the National Geographic special "Death of a Sea Monster".


title: Ian McElhinney

text: Ian McElhinney (born 19 August 1948) is a Northern Irish actor and director. He has appeared in many television series in a career spanning more than forty years; notable appearances include "Taggart", "Hornblower", "Cold Feet", and "The Tudors". In recent times his best known roles are as Barristan Selmy in "Game of Thrones", Morgan Monroe in "The Fall", and Granda Joe in "Derry Girls".


title: Ian McElhinney

text: McElhinney was born in Belfast, the son of a Church of Ireland (Anglican) clergyman and teacher. He studied international affairs at Brandeis University in Waltham, Massachusetts. He worked as a teacher at Goole Grammar School, now known as Goole Academy, in Yorkshire, England, for several years before becoming an actor. He started acting professionally at the age of 30, playing Bill Sykes in a theatre production of "Oliver!".


title: Ian McElhinney

text: He is married to playwright and actress Marie Jones; in 2009 the couple started their own company, Rathmore Productions Ltd. They live in Belfast and have three sons.


title: Ian McElhinney

text: At the end of 2008, McElhinney formed another company, Stones Film Productions Ltd, with Mary Crossan Magowan, Chris Parr, and Anthony E Rowe. In 2013, Rowe departed the company. As of November 2022, the Internet Movie Database lists nothing made by Stones Film.


title: Paul F. Tompkins

text: Paul Francis Tompkins (born September 12, 1968) is an American comedian, actor, and writer. He is known for his work in television on such programs as "Mr. Show with Bob and David", "Real Time with Bill Maher", and "Best Week Ever", later renamed "Best Week Ever with Paul F. Tompkins".


title: Paul F. Tompkins

text: He is known for his numerous appearances on podcasts, including his 200-plus appearances on "Comedy Bang! Bang!" He has also been the host of the Fusion Channel talk show "No, You Shut Up!", "The Dead Authors Podcast", the online Made Man interview series "Speakeasy with Paul F. Tompkins", the Earwolf podcast "Spontaneanation with Paul F. Tompkins", and "The Pod F. Tompkast", which was ranked #1 by "Rolling Stone" on their list of "The 10 Best Comedy Podcasts of the Moment" in 2011.


title: Paul F. Tompkins

text: He is also a main cast member of the "Superego" podcast and was a regular player on "Thrilling Adventure Hour" podcast, which ended in 2015. Tompkins was the voice of Mr. Peanutbutter, an anthropomorphic yellow , on the Netflix animated series "BoJack Horseman" from 2014 to 2020. He has also appeared in drama films like "There Will Be Blood" and "The Informant!". In 2021, he had a recurring role on the sitcom "Rutherford Falls".


title: Paul F. Tompkins

text: In December 2014, "Paste" named his Twitter one of "The 75 Best Twitter Accounts of 2014", ranking it at #70.


title: Paul F. Tompkins

text: Paul Francis Tompkins was born September 12, 1968, in Mount Airy, Philadelphia, Pennsylvania. He has two brothers (one older, one younger) and three sisters (all older).


title: Paul F. Tompkins

text: In 1986, Tompkins first performed comedy at 17 years of age at The Comedy Works in Philadelphia (a club now located in Bristol, Pennsylvania), where he performed as half of a sketch comedy duo with Rick Roman. Tompkins attended Temple University; however, he dropped out and left for Los Angeles, California, in 1994.


title: Paul F. Tompkins

text: Tompkins met actor Jay Johnston in L.A. through their mutual friend, actor and director Adam McKay. McKay and Tompkins had become friends in Philadelphia, where they had both started to perform stand-up at around the same time. McKay later moved to Chicago and met Johnston; Johnston moved to L.A. at around the same time as Tompkins and McKay introduced the two. Tompkins and Johnston went on to create a live sketch comedy show called "The Skates" that was seen by Bob Odenkirk and David Cross and helped get them hired to work on "Mr. Show with Bob and David" in 1996.


title: Paul F. Tompkins

text: Tompkins' comedy career has included stand-up, sketch comedy and a variety of other live performances.


title: Paul F. Tompkins

text: Tompkins' stand-up comedy performances are of a storytelling and observationalist style. His shows often consist of extended riffs and long anecdotes. Tompkins deals with topics of the bizarre and the absurd—such as a rant about peanut brittle, a discussion about cake versus pie, and smashed coins—in addition to recounting stories about his own life experiences and family. His comedic style has been described as alternative comedy; Tompkins has stated that he is not bothered by the label and that he likes the term.


title: Paul F. Tompkins

text: Tompkins is known for his style of dress during his live comedic performances, always performing in suit and tie, sometimes in pinstripes and with a bowtie; his look has been described by some in the press as "dapper". Tompkins has described his look as "foppish" and "just this side of Cedric the Entertainer."


title: Paul F. Tompkins

text: Tompkins is based in Los Angeles and performs regularly in the city. Since 2002 he has performed a monthly show called "The Paul F. Tompkins Show" at Largo, an L.A. nightclub and cabaret. His show has featured such guests as Fiona Apple, Jack Black, Dave Foley, Zach Galifianakis, Ed Helms, Aimee Mann, and Weird Al Yankovic. Since its inception in 2005, Tompkins has taken part in the "Thrilling Adventure Hour", a staged production in the style of old-time radio that is also held monthly at Largo. The show began podcasting in January 2011; in October of that same year the show's podcasts moved to the Nerdist Industries podcast network created by Chris Hardwick. Tompkins is a member of the Upright Citizens Brigade Theatre (UCB) Los Angeles. His comedy album "Impersonal" was recorded live at the UCB Theatre. He also performs monthly at the "Dead Authors" show at UCB Theatre in support of the nonprofit organization 826LA; Tompkins plays the role of H.G. Wells who serves as the host of the show.


title: Paul F. Tompkins

text: Tompkins has toured in the US and Canada and prefers to perform in independent venues, rather than conventional comedy clubs. Starting in 2009 he embarked on his "Tompkins 300" tour. Tompkins had been preparing for his one-hour Comedy Central special "You Should Have Told Me" at the Laughing Skull Lounge theatre in Atlanta, Georgia—a small theatre that seats about 74 people. In order to fill the seats for the recording of his special, Tompkins required about 280 people in the audience over the course of four nights for the recording of his show. Tompkins decided to announce on Twitter that he needed 300 people to fill the seats each night.


title: Paul F. Tompkins

text: Bob Kerr, a Canadian comedian, saw the Twitter post and asked if Tompkins would like to perform in Toronto. Tompkins advised Kerr that if he was able to get 300 people to state that they would definitely see his show he would come to Toronto. Kerr then started a Facebook group called "I Wanna See Paul F. Tompkins in Toronto" and managed to get 300 people to join. In October of that same year Tompkins performed at The Rivoli night club in Toronto, the same club in which the sketch comedy troupe The Kids in the Hall got their start. Facebook groups were subsequently started in other North American cities and in 2010 he stated that he had stopped promoting his shows on the radio. In 2011 he said that the Facebook 300 groups had become his main method of booking comedy shows.


title: Paul F. Tompkins

text: Tompkins wrote and performed in his one-man show, "Driven to Drink", which aired on HBO in 1998. He appeared on 6 episodes of "Late Night with Conan O'Brien" between 1998 and 2008 as well as two episodes of "Conan" in 2011 and 2012. He has recorded three comedy albums: "Impersonal" in 2007, "Freak Wharf" in 2009, and "Laboring Under Delusions" in 2012. His stand-up appearances on the Comedy Central network include being featured in episodes of "Comedy Central Presents" in 2003 and 2007, hosting an episode of "Live at Gotham" in 2009, performing on "John Oliver's New York Stand Up Show" in 2010, and recording two original one-hour comedy specials—"You Should Have Told Me" which aired in 2010 and "Paul F. Tompkins: Laboring Under Delusions" in 2012. He also appeared in the RiffTrax live broadcast of "House on Haunted Hill".


title: Paul F. Tompkins

text: Tompkins wrote for and performed on "Mr. Show with Bob and David" from 1995 to 1998; the show's writers, including Tompkins, were nominated for an Emmy Award in 1998 for "Outstanding Writing for a Variety or Music Program".


title: Paul F. Tompkins

text: Tompkins's work with Mr. Show's creators Bob Odenkirk and David Cross also led to his recurring role on the "Tenacious D" TV series. Tompkins played the character of a nightclub manager who is duped into reading Tenacious D's ridiculous introductions during their open mic performances. He revived the role in the comedic band's film "Tenacious D in The Pick of Destiny" released in 2006.


title: Paul F. Tompkins

text: Longtime friend Adam McKay consulted Tompkins regarding the screenplay for "Talladega Nights". Tompkins also played the MC of a cat show in McKay's "".


title: Paul F. Tompkins

text: Tompkins has appeared on television programs including "NewsRadio", "Frasier", "Weeds", "The Sarah Silverman Program", "Pushing Daisies", "Community", and "Curb Your Enthusiasm". Tompkins played the role of Prescott in Paul Thomas Anderson's film "There Will Be Blood" (2007); Anderson had previously cast Tompkins in a small role in the 1999 film "Magnolia" after watching Tompkins perform at Largo. Tompkins also played FBI Agent Anthony D’Angelo in Steven Soderbergh's "The Informant!" (2009). He has a recurring role in the Canadian TV series "The L.A. Complex" as a fictionalized version of himself. He also appeared in the music video for Nick Lowe's song "Stoplight Roses" and in the Ted Leo and the Pharmacists song "Bottled In Cork". Tompkins wrote for "Real Time with Bill Maher" in 2003 and 2009, in addition to being a show correspondent in the show's first season. In 2011 Tompkins was asked to write humorous recaps of "American Idol" episodes for "New York" magazine's online blog "Vulture".


title: Paul F. Tompkins

text: Tompkins has expressed in interviews that he dislikes writing (particularly writing for others), preferring instead to perform in front of a camera.


title: Paul F. Tompkins

text: Tompkins has done voice work for many animated television series including "Dr. Katz, Professional Therapist", "King of the Hill", and "Bob's Burgers", in which he voices the recurring character Randy. He lent his voice to a character in an unaired 2007 episode of "Aqua Teen Hunger Force" titled "Boston" that was supposed to be the premiere episode of the show's fifth season, but it was pulled by Turner Broadcasting System to avoid further controversy surrounding the 2007 Boston bomb scare.


title: Paul F. Tompkins

text: Tompkins later appeared in an episode during the show's 7th season. He was also the voice of one of the thugs in Walt Disney Animation Studios' 2010 computer animated film "Tangled". Tompkins was the voice of Benton Criswell, a character in MTV series "Super Adventure Team" which featured marionettes in the style of the 1960s British series "Thunderbirds"; the role was credited under the stage name Francis Mt. Pleasant. He was the voice of a puppet in ads for the Ford Focus.


title: Paul F. Tompkins

text: Tompkins played Mr. Peanutbutter in the 2014 Netflix original animated series "BoJack Horseman". Tompkins voices the recurring character, Gladstone Gander, in the reboot series of "DuckTales". In 2020, he voiced Dr. Migleemo, a therapist aboard the USS Cerritos in the CBS All Access animated series "".


title: Paul F. Tompkins

text: Tompkins has appeared on several television programs devoted to discussing politics, popular culture and current events; however, he says he does not consider himself to be a political comic.


title: Paul F. Tompkins

text: Tompkins was a contributor to the "Us People's Weekly Entertainment" segment of "The Daily Show" in 1998. In 2003 he was a writer and correspondent for "Real Time with Bill Maher" in the show's first season and wrote again for the show in 2009. He appeared on "Tough Crowd with Colin Quinn" in 2004. In 2004 he also became a pop culture analyst on VH1's "Best Week Ever"; in 2008 the show was retooled and relaunched as "Best Week Ever with Paul F. Tompkins" with Tompkins as host. From 2006 to 2008 he was a regular guest on "Countdown with Keith Olbermann". In 2008 he appeared on "Lewis Black's Root of All Evil" and took part in a panel on "Larry King Live" in an episode titled "Politics & Humor".


title: Paul F. Tompkins

text: Tompkins has appeared in documentaries such as Jamie Kennedy's "Heckler" (2007) and Doug Benson's "Super High Me" (2007). He also appeared in "The Bitter Buddha" (2013), a documentary about the career of actor and comedian Eddie Pepitone.


title: Paul F. Tompkins

text: Tompkins later became the host of a discussion show called "No, You Shut Up!" by The Jim Henson Company under its Henson Alternative banner.


title: Paul F. Tompkins

text: In 2010, Tompkins launched his podcast called "The Pod F. Tompkast". The podcast was a mixture of Tompkins discussing various topics, clips from his live show at Largo, and segments where Tompkins voices a variety of celebrities speaking with one another. Comedian Jen Kirkman was a regular contributor on the show. The podcast ended in 2012.


title: Paul F. Tompkins

text: The "Thrilling Adventure Hour" comedy show at Largo began podcasting in January 2011; in October of that same year the show's podcasts moved to the Nerdist Industries podcast network created by Chris Hardwick. The show ended in 2015.


title: Paul F. Tompkins

text: "Dead Authors", a live show that Tompkins hosts at the UCB Theatre in Los Angeles, also began podcasting in September 2011.


title: Paul F. Tompkins

text: In May 2012 Tompkins started a weekly web series called "Speakeasy". Hosted by the Break Media site MadeMan.com, the series features Tompkins interviewing various guests in the entertainment industry, such as Ty Burrell, Nathan Fillion, Zach Galifianakis, Chris Hardwick, Oscar Nunez, Weird Al Yankovic, and Alison Brie. The interviews are conducted as casual conversations between Tompkins and his guests over cocktails at various bars in the L.A. area.


title: Paul F. Tompkins

text: Tompkins has appeared well over 200 times as a guest, and occasionally as a guest host, on "Comedy Bang! Bang!" (formerly "Comedy Death-Ray Radio"), a weekly audio podcast hosted by Scott Aukerman, a comedian who also wrote for "Mr. Show with Bob and David". The show's format mixes conversation between the host and guests, and usually includes improv games. Some guests play characters or impersonate certain celebrities, usually for the entirety of the episode; Paul F. Tompkins has impersonated celebrities such as rapper Ice-T, composer Andrew Lloyd Webber, and Buddy Valastro from the reality television series "Cake Boss (cakeboss.)".


title: Paul F. Tompkins

text: In addition to Aukerman's "Comedy Bang! Bang!", Tompkins has appeared on the podcasts of other fellow comedians such as "WTF with Marc Maron", Jimmy Pardo's "Never Not Funny", Jessica Chaffin and Jamie Denbo's "Ronna and Beverly" podcast, and the "Superego" podcast with Jeremy Carter, Matt Gourley, and Mark McConville. Tompkins has also been a regular guest on the radio show and podcast "The Best Show on WFMU with Tom Scharpling".


title: Paul F. Tompkins

text: In 2015, Tompkins created his own podcast on the Earwolf podcast network called "Spontaneanation with Paul F. Tompkins". This podcast is similar to the "Pod F. Tompkast"; however, "Spontaneanation" is fully improvised and in-the-moment, as opposed to the "Tompkast", which was highly produced. "Spontaneanation" begins with an improvised monologue, accompanied on piano by Eban Schletter, much like the "Pod F. Tompkast". The next segment is an interview with one of Tompkins' famous friends. The final segment is one long improvised story performed by Paul and guest improvisers, based on ideas discussed in the interview segment. The podcast ended in 2019.


title: Paul F. Tompkins

text: Since 2019, Tompkins has cohosted, with Nicole Parker, "The Neighborhood Listen" on Stitcher Premium. Along with Tawny Newsome, he also hosts "The Pod Directive", the official Star Trek podcast, which launched in September 2020.


title: Paul F. Tompkins

text: In March 2020, during the COVID-19 pandemic, Paul started a weekly podcast with his wife, Janie Haddad Tompkins, called "Stay F. Homekins". The first season ran for 41 episodes throughout the rest of 2020, and season two started in January 2021.


title: Paul F. Tompkins

text: In 2017, Tompkins stated on Twitter that he was once a Catholic, but later went on to become an atheist.


title: Paul F. Tompkins

text: He is a member of the Democratic Socialists of America and helped canvass for Eunisses Hernandez and Hugo Soto-Martinez during the 2022 Los Angeles elections.


title: Paul F. Tompkins

text: Tompkins is known for his many podcast appearances, as well as hosting a few of his own. He is often referred to as the mayor of podcasts.


title: Type 212 submarine

text: The German Type 212 class (German: U-Boot-Klasse 212 A), also Italian Todaro class, is a diesel-electric submarine developed by Howaldtswerke-Deutsche Werft AG (HDW) for the German and Italian navies. It features diesel propulsion and an additional air-independent propulsion (AIP) system using Siemens proton exchange membrane (PEM) compressed hydrogen fuel cells. The submarines can operate at high speed on diesel power or switch to the AIP system for silent slow cruising, staying submerged for up to three weeks with little exhaust heat. The system is also said to be vibration-free, extremely quiet and virtually undetectable.


title: Type 212 submarine

text: At the beginning of the 1990s the German Navy was seeking a replacement for the Type 206 submarines. Initial study started on a Type 209 improved design, with AIP capability, called Type 212.


title: Type 212 submarine

text: The final programme started in 1994 as the two navies of Germany and Italy began working together to design a new conventional submarine, respectively to operate in the shallow and confined waters of the Baltic Sea and in the deeper waters of the Mediterranean Sea. The two different requirements were mixed into a common one and, because of significant updates to the design, the designation has been changed to Type 212A since then.


title: Type 212 submarine

text: On 22 April 1996 a Memorandum of Understanding (MOU) gave the start to the cooperation for building four vessels for German Navy and four vessels for Italian Navy. Its main aim was the construction of identical boats and the start of a collaboration in logistic and life-cycle support for the two navies.


title: Type 212 submarine

text: The German government placed an initial order of four "Type 212A" submarines in 1998. The German Submarine Consortium built them at the shipyards of HDW and Thyssen Nordseewerke GmbH (TNSW) of Emden. Different sections of the submarines were constructed at both sites at the same time and then half of them were shipped to the respective other yard so that both HDW and Thyssen Nordseewerke assembled two complete submarines each.


title: Type 212 submarine

text: In the same year the Italian government placed an order of two "U212A" submarines built by Fincantieri for the Italian Navy ("Marina Militare") at Muggiano shipyard, designated as the Todaro class.


title: Type 212 submarine

text: The German Navy ordered two additional, improved submarines in 2006, to be delivered from 2012 on. They will be 1.2 meters longer to give additional space for a new reconnaissance mast.


title: Type 212 submarine

text: On 21 April 2008 the Italian Navy ordered the optional second batch of submarine, in the same configuration of the original ones. Some upgrading should involve materials and components of commercial derivation, as well as the software package of the CMS. The intention is to keep the same configuration of the first series and reduce maintenance costs.


title: Type 212 submarine

text: The export-oriented Type 214 submarine succeeds the Type 209 submarine and shares certain features with the Type 212A, such as the AIP fuel cell propulsion.


title: Type 212 submarine

text: Poland announced in December 2013 they will not buy, but only lease, two U212-A's, on account of not meeting "requirements of tactical and technical equipment developed by the military, including in particular the propulsion system, missile weapons and rescue system".


title: Type 212 submarine

text: On 22 December 2015 Admiral Giuseppe De Giorgi, Commander in Chief of the Italian Navy, announced plans to build another two U212A submarines.


title: Type 212 submarine

text: In October 2016, during the celebration of the commissioning of "U36", the German Navy announced the intent to procure another batch of two U212A within the next decade.


title: Type 212 submarine

text: In February 2017, it was announced that the Royal Norwegian Navy will procure four submarines based on Type 212. Initial plans envisaged service entry between 2025 and 2028. However, the Norwegian 2020 Defence Plan later envisaged service entry "around 2030". This "CD" (Common Design) variant of the Type 212 will consist of six submarines, with the German Navy ordering two new boats alongside the four Norwegian vessels. In March 2021 it was indicated that an agreement had been reached between Norway and Germany to initiate the acquisition program, pending approval by the Bundestag. The contract for construction of the six boats was signed in July 2021 with delivery of the first boat to the Royal Norwegian Navy anticipated in 2029.


title: Type 212 submarine

text: Partly owing to the "X" arrangement of the stern planes, the "Type 212" is capable of operating in as little as 17 metres of water. This is a long-standing requirement for German submarine designs, enabling them to pass a strategic point in the Baltic Sea (the "Kadettrinne") submerged. This allows it to come much closer to shore than most contemporary submarines. This gives it an advantage in covert operations, as SCUBA-equipped commandos operating from the boat can surface close to the beach and execute their mission more quickly and with less effort.


title: Type 212 submarine

text: A notable design feature is the prismatic hull cross-section and smoothly faired transitions from the hull to the sail, improving the boat's stealth characteristics. The ship and internal fixtures are constructed of nonmagnetic materials, significantly reducing the chances of it being detected by magnetometers or setting off magnetic naval mines.


title: Type 212 submarine

text: Although hydrogen–oxygen propulsion had been considered for submarines as early as World War I, the concept was not very successful until recently due to fire and explosion concerns. In the "Type 212" this has been countered by storing the fuel and oxidizer in tanks outside the crew space, between the pressure hull and outer light hull. The gases are piped through the pressure hull to the fuel cells as needed to generate electricity, but at any given time there is only a very small amount of gas present in the crew space.


title: Type 212 submarine

text: Currently, the "Type 212A" is capable of launching the fiber optic-guided DM2A4 Seehecht ("Seahake") heavyweight torpedoes, the WASS BlackShark torpedoes and short-range missiles from its six torpedo tubes, which use a water ram expulsion system. Future capability may include tube-launched cruise missiles.


title: Type 212 submarine

text: The short-range IDAS missile (based on the IRIS-T missile), primarily intended for use against air threats as well as small or medium-sized sea- or near land targets, is currently being developed by Diehl BGT Defence to be fired from Type 212's torpedo tubes. IDAS is fiber-optic guided and has a range of approx. 20 km. Four missiles fit in one torpedo tube, stored in a magazine. First deliveries of IDAS for the German Navy were scheduled from 2014 on.


title: Type 212 submarine

text: A 30mm auto-cannon called Muräne (moray) to support diver operations or to give warning shots is being considered, too. The cannon, probably a version of the RMK30 built by Rheinmetall, will be stored in a retractable mast and can be fired without the boat emerging. The mast will also be designed to contain three Aladin UAVs for reconnaissance missions. This mast is likely to be mounted on the second batch of Type 212 submarines for the German Navy.


title: Type 212 submarine

text: In April 2006, the German Navy's "U-32" sailed from the Baltic Sea to Rota, Spain in a journey lasting two weeks which covered without surfacing or snorkelling.


title: Type 212 submarine

text: The Italian Navy's S 526 "Todaro" was deployed, for over six months in 2008, to the United States for CONUS 2008 exercise with the United States Navy.


title: Type 212 submarine

text: The Italian Navy's S 527 "Scirè" was deployed, for over five months in 2009, to the U.S. for CONUS 2009 exercise with the United States Navy.


title: Type 212 submarine

text: The Italian Navy's S 526 "Todaro", between 1 September 2012 and 13 February 2013, for the first time was deployed to the Aden Gulf, Arabic Sea, Oman Gulf and Indian Ocean.


title: Type 212 submarine

text: In 2013, while on the way to participate in naval exercises in U.S. waters, the German Navy's "U-32" established a new record for non-nuclear submarines with 18 days in submerged transit without snorkelling.


title: Type 212 submarine

text: On 15 October 2017, the German Navy's "U-35" suffered damage to its rudder fins while conducting dives off the Norwegian coast.


title: Lola Kirke

text: Lola Clementine Kirke (born 27 September 1990) is an English-born American actress and singer-songwriter. She starred in the 2015 film "Mistress America" and the Amazon Studios television series "Mozart in the Jungle." She appeared in "Gone Girl" (2014), "Gemini" (2017), and "Lost Girls" (2020) among other films. Recent work includes HBO's "Winning Time" (2022) and upcoming Showtime limited series "Three Women".


title: Lola Kirke

text: Kirke was born in Westminster, London, and raised in New York City from the age of five. Her father, Simon Kirke, was drummer for the rock bands Bad Company and Free. Her mother is Lorraine (née Dellal) Kirke, the owner of Geminola, a vintage boutique in New York City that supplied a number of outfits for the television series "Sex and the City". ("Geminola" is an amalgam of the names of Lola and her siblings.) Kirke graduated from Saint Ann's School in Brooklyn and in 2012 from Bard College.


title: Lola Kirke

text: Her father is of English and Scottish descent (the Kirkes being a junior branch of a family of Nottinghamshire landed gentry, and descending also from the Gibson-Craig baronets) and her mother is Jewish. Kirke's maternal grandfather, Jack Dellal, was a British businessman of Iraqi-Jewish descent, and her maternal grandmother Zehava Helmer was a former Israeli stewardess; Kirke has described herself as Jewish. Kirke has two older sisters, singer Domino Kirke and actress Jemima Kirke, and an older half-brother from a previous relationship of her mother's. She is a cousin of curator Alexander Dellal, shoe designer Charlotte Olympia Dellal, and model Alice Dellal.


title: Lola Kirke

text: Kirke's breakout role came as the main character Hayley Rutledge in the Amazon series "Mozart in the Jungle". It ran for four seasons, from 2014 to 2018. In 2014, she had a supporting role in the David Fincher film "Gone Girl", before going on to star in "Mistress America" alongside Greta Gerwig. In 2017, Kirke starred opposite Zoe Kravitz in the Neon film "Gemini", a neo-noir mystery film set in Hollywood, which made its world premiere at South By Southwest on March 12, 2017. In 2018 she starred in the drama film "Untogether", playing sisters with her real-life sister Jemima Kirke. It premiered at the 2018 Tribeca Film Festival. She appears in the mystery drama "Lost Girls", which made its premiere at The Sundance Film Festival in 2020, and was later made available on Netflix. In 2021, Kirke starred in the BJ Novak anthology series "The Premise" for FX and Hulu. Kirke appears in the HBO series about the Los Angeles Lakers, "Winning Time", and will be seen in the Showtime drama series "Three Women", based on Lisa Taddeo’s "book of the same name".


title: Lola Kirke

text: Kirke is also active in music. In 2016 she released an EP. In 2018, she released three singles, including two with accompanying music videos, "Monster" and "Supposed To". Her debut album, "Heart Head West", was released on 10 August 2018 by Downtown Records.


title: Lola Kirke

text: On 12 March 2019, Kirke released a cover of Rick Danko's "Sip the Wine" on Downtown Records, produced by Matthew E. White at his Richmond, Virginia studio, Spacebomb Studios.


title: Lola Kirke

text: In April 2022, they released her sophomore LP Lady for Sale. The album was hailed as "pure delight" by Pitchfork, "immediately iconic" by AllMusic, and was said to bridge "the gap that once existed between Madonna and the Mandrell Sisters," by Uncut Magazine. "A new era of sparkle-country is upon us," declared No Depression. "Lola Kirke is here to usher it in." "Lyrically, Kirke moves into the top tier of writers," wrote The Line of Best Fit.


title: Lola Kirke

text: In August 2022, it was announced that Kirke would be supporting Swedish folk duo First Aid Kit on their UK Tour in November and December 2022.


title: Lola Kirke

text: Kirke is an activist for women's rights. In 2017 she notably wore a "Fuck Paul Ryan" pin on her gown to the Golden Globes to protest the defunding of Planned Parenthood.


title: Glory Johnson

text: Glory Bassey Johnson (born July 27, 1990) is an American basketball player for Turkish club Beşiktaş. Born in Colorado Springs, Colorado, she went to Webb School Of Knoxville and played collegiately for the University of Tennessee Lady Vols. She holds a Montenegrin passport and has represented the Montenegro national team internationally.


title: Glory Johnson

text: Johnson enrolled at the University of Tennessee-Knoxville in 2008. She recorded 36 double-doubles over the course of her college career.


title: Glory Johnson

text: Johnson completed her bachelor's degree in global studies in three years, and earned a master's degree in communications during her fourth year of basketball eligibility.


title: Glory Johnson

text: Johnson played on the team representing the US at the 2011 Summer Universiade held in Shenzhen, China. The team won all six games to earn the gold medal. Johnson scored 6.2 points per game and had nine steals, tied for second place on the team.


title: Glory Johnson

text: Johnson was selected in the first round of the 2012 WNBA Draft (4th overall) by the Tulsa Shock. In her rookie season, she was ranked fourth in steals per game (a career-high 2.1 spg).


title: Glory Johnson

text: In 2013 and 2014, her second and third seasons with the Shock, Johnson was named in the Western Conference All-Star team.


title: Glory Johnson

text: She returned one month into the 2016 WNBA season after her suspension stemming from her domestic violence incident with Brittney Griner. By this time the Tulsa Shock had relocated to Dallas and were renamed the Dallas Wings, Johnson had re-signed with the team during free agency. Johnson averaged 11.3 ppg in 18 games with 6 starts. Also during her comeback season, she recorded the league's 16th 20-point, 20-rebound performance of the season in a win against the Phoenix Mercury when she scored 23 points along with a career-high 22 rebounds.


title: Glory Johnson

text: During the 2017 season, Johnson continued to put double-doubles in rebounds and points which is what she has become statistically known for earlier on in her career. On June 18, 2017, Johnson score a season-high 27 points in an 87–83 win over the Washington Mystics. From July 31 to August 6, Johnson named the Western Conference player of the week. On August 19, 2017, Johnson recorded her 13th double-double of the season, scoring 23 points along with 13 rebounds in a 90–86 win over the Atlanta Dream. During the game, Johnson threw a punch at Dream's point guard Matee Ajavon and served a one-game suspension two days later. The Wings would finish with a 16–18 record and the number 7 seed in the league. In her first career playoff game, Johnson scored 15 points and grabbed 14 rebounds in a losing effort to the Washington Mystics in the first round elimination game.


title: Glory Johnson

text: In 2018, Johnson had a bit of a challenging season with a couple injuries sidelining her for a few games and causing her to play some games off the bench. She played 29 games with 17 starts and would average career-lows in scoring and rebounding. The Wings would still make the playoffs with a 15–19 record and the 8th seed in the league. They would once again be a first-round exit as they lost to the Phoenix Mercury.


title: Glory Johnson

text: From 2012 to 2015, Johnson played three off-seasons in Russia for two different teams; Chevakata Vologda and Nadezhda Orenburg. As of November 2016, Johnson signed with the Xinjiang Tiashan Deers of the WCBA for the 2016–17 off-season. In August 2018, Johnson signed with Hatay BB of the Turkish League for the 2018–19 off-season.


title: Glory Johnson

text: On August 14, 2014, it was announced that Johnson and fellow WNBA player Brittney Griner were engaged. On April 22, 2015, both women were arrested for physically attacking each other after police broke up a fight between the two in their Goodyear, Arizona, home. Both sustained minor injuries during the incident. Nevertheless, they stayed together and married on May 8, 2015. It was announced on June 4, 2015 that Johnson was pregnant and that she would miss the 2015 WNBA season. One day later, Griner filed for annulment which was rejected. Johnson gave birth to twin girls in October 2015, 16 weeks premature. The twins were conceived by IVF treatment using her eggs and a sperm donation. The divorce was finalized in June 2016.


title: Mid-century modern

text: Mid-century modern (MCM) is an American design movement in interior, product, graphic design, architecture, and urban development that was popular from roughly 1945 to 1969, during the United States's post–World War II period. The term was used descriptively as early as the mid-1950s and was defined as a design movement by Cara Greenberg in her 1984 book "Mid-Century Modern: Furniture of the 1950s". It is now recognized by scholars and museums worldwide as a significant design movement. The MCM design aesthetic is modern in style and construction, aligned with the Modernist movement of the period. It is typically characterized by clean, simple lines and honest use of materials, and it generally does not include decorative embellishments.


title: Mid-century modern

text: The Mid-century modern movement in the U.S. was an American reflection of the International and Bauhaus movements, including the works of Gropius, Florence Knoll, Le Corbusier and Ludwig Mies van der Rohe. Although the American component was slightly more organic in form and less formal than the International Style, it is more firmly related to it than any other. Brazilian and Scandinavian architects were very influential at this time, with a style characterized by clean simplicity and integration with nature. Like many of Wright's designs, Mid-century architecture was frequently employed in residential structures with the goal of bringing modernism into America's post-war suburbs. This style emphasized creating structures with ample windows and open floor plans, with the intention of opening up interior spaces and bringing the outdoors in. Many Mid-century houses utilized then-groundbreaking post and beam architectural design that eliminated bulky support walls in favor of walls seemingly made of glass. Function was as important as form in Mid-century designs, with an emphasis placed on targeting the needs of the average American family.


title: Mid-century modern

text: In Europe, the influence of Le Corbusier and the CIAM resulted in an architectural orthodoxy manifest across most parts of post-war Europe that was ultimately challenged by the radical agendas of the architectural wings of the avant-garde Situationist International, COBRA, as well as Archigram in London. A critical but sympathetic reappraisal of the internationalist oeuvre, inspired by Scandinavian Moderns such as Alvar Aalto, Sigurd Lewerentz and Arne Jacobsen, and the late work of Le Corbusier himself, was reinterpreted by groups such as Team X, including structuralist architects such as Aldo van Eyck, Ralph Erskine, Denys Lasdun, Jørn Utzon and the movement known in the United Kingdom as New Brutalism.


title: Mid-century modern

text: Pioneering builder and real estate developer Joseph Eichler was instrumental in bringing Mid-century modern architecture ("Eichler Homes") to subdivisions in the Los Angeles area and the San Francisco Bay region of California, and select housing developments on the east coast. George Fred Keck, his brother Willam Keck, Henry P. Glass, Mies van der Rohe, and Edward Humrich created Mid-century modern residences in the Chicago area. Mies van der Rohe's Farnsworth House is extremely difficult to heat or cool, while Keck and Keck were pioneers in the incorporation of passive solar features in their houses to compensate for their large glass windows.


title: Mid-century modern

text: The city of Palm Springs, California is noted for its many examples of Mid-century modern architecture.


title: Mid-century modern

text: Examples of 1950s Palm Springs motel architecture include Ballantines Movie Colony (1952) – one portion is the 1935 Albert Frey San Jacinto Hotel – the Coral Sands Inn (1952), and the Orbit Inn (1957). Restoration projects have been undertaken to return many of these residences and businesses to their original condition.


title: Mid-century modern

text: Scandinavian design was very influential at this time, with a style characterized by simplicity, democratic design and natural shapes. Glassware (Iittala – Finland), ceramics (Arabia – Finland), tableware (Georg Jensen – Denmark), lighting (Poul Henningsen – Denmark), and furniture (Danish modern) were some of the genres for the products created. In America, east of the Mississippi, the American-born Russel Wright, designing for Steubenville Pottery, and Hungarian-born Eva Zeisel designing for Red Wing Pottery and later Hall China created free-flowing ceramic designs that were much admired and heralded in the trend of smooth, flowing contours in dinnerware. On the West Coast of America the industrial designer and potter Edith Heath (1911–2005) founded Heath Ceramics in 1948. The company was one of the numerous California pottery manufacturers that had their heyday in post-war US, and produced Mid-Century modern ceramic dish-ware. Edith Heath's "Coupe" line remains in demand and has been in constant production since 1948, with only periodic changes to the texture and color of the glazes. The Tamac Pottery company produced a line of mid-century modern biomorphic dinnerware and housewares between 1946 to 1972.