Aphanosauria

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Aphanosauria
Temporal range: Middle Triassic, Anisian–Ladinian
Teleocrater v1.png
Illustration of Teleocrater rhadinus
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Archosauria
Clade: Avemetatarsalia
Clade: Aphanosauria
Nesbitt et al., 2017
Genera

Aphanosauria ("hidden lizards") is an extinct group of reptiles distantly related to dinosaurs (including birds). They are at the base of a group known as Avemetatarsalia, one of two main branches of archosaurs. The other main branch, Pseudosuchia, includes modern crocodilians. Aphanosaurs possessed features from both groups, indicating that they are the oldest and most primitive known clade of avemetatarsalians, at least in terms of their position on the archosaur family tree. Other avemetatarsalians include the flying pterosaurs, small bipedal lagerpetids, herbivorous silesaurids, and the incredibly diverse dinosaurs, which survive to the present day in the form of birds. Aphanosauria is formally defined as the most inclusive clade containing Teleocrater rhadinus and Yarasuchus deccanensis but not Passer domesticus (house sparrow) or Crocodylus niloticus (Nile crocodile). This group was first recognized during the description of Teleocrater. [1] Although only known by a few genera, Aphanosaurs had a widespread distribution across Pangaea in the Middle Triassic. [2] They were fairly slow quadrupedal long-necked carnivores, a biology more similar to basal archosaurs than to advanced avemetatarsalians such as pterosaurs, lagerpetids, and early dinosaurs. In addition, they seemingly possess 'crocodile-normal' ankles (with a crurotarsal joint), showing that 'advanced mesotarsal' ankles (the form acquired by many dinosaurs, pterosaurs, lagerpetids, and advanced silesaurids) were not basal to the whole clade of Avemetatarsalia. Nevertheless, they possessed elevated growth rates compared to their contemporaries, indicating that they grew quickly, more like birds than other modern reptiles. Despite superficially resembling lizards, the closest modern relatives of aphanosaurs are birds. [1]

Contents

Description

Members of this group were lightly-built and moderately-sized reptiles. They do not show any adaptations for bipedalism, which became much more common in other avemetatarsalians. In addition, their leg proportions indicate that they were not capable of sustained running, meaning that they were also slow by avemetatarsalian standards. [1]

Skull

Very little skull material is known for the group as a whole. The only skull bones which can be confidently referred to this group consist of a few pterygoid and postorbital fragments belonging to Yarasuchus as well as some fragmentary material considered to belong to Teleocrater. These bones include a maxilla (tooth-bearing bone of the middle of the snout), frontal (part of the skull roof above the eyes), and a quadrate (part of the cranium's jaw joint). Although these fragments make it difficult to reconstruct the skull of aphanosaurs, they do show several notable features. For example, the shape of the maxilla shows that aphanosaurs had an antorbital fenestra, a large hole on the snout just in front of the eyes. Coupled with an antorbital depression (a collapsed area of bone which surrounded the fenestra), these indicate that aphanosaurs belonged to the group Archosauria. A partially-erupted tooth was also preserved on the lower edge of the maxilla. This tooth was flattened from the sides, slightly curved backwards, and serrated along its front edge. These tooth features indicate that aphanosaurs were carnivorous, as many meat-eating reptiles (including theropod dinosaurs such as Velociraptor and Deinonychus ) had the same features. The front edge of the maxilla also has a small pit, similar to some silesaurids. The rear part of the frontal possessed a round, shallow pit known as a supratemporal fossa. In the past it was believed that only dinosaurs possessed supratemporal fossae, but its presence in aphanosaurs (and Asilisaurus , a silesaurid) shows that it was variable among many avemetatarsalians. As a whole, known aphanosaurian skull material possessed no unique features, meaning that the rest of the skeleton would have to be used to characterize the group. [2]

Vertebrae

Aphanosaurs have many distinguishing features of their cervicals (neck vertebrae). The cervicals are very long compared to those of other early avemetatarsalians. As with most other reptiles, the vertebrae are composed of a roughly cylindrical main body (centrum) and a plate-like neural spine jutting out of the top. In the anterior cervicals (vertebrae at the front of the neck), a pair of low ridges run down the underside of the centrum. These ridges are separated by a wide area with other shallower ridges, making the centrum roughly rectangular in cross-section. The neural spines of the cervicals are also unique in aphanosaurs. They are hatchet shaped, with front edges that taper to a point and drastically overhang the centrum, at least in the front and middle parts of the neck. The upper edge of the neural spine is thin and blade-like, but the area immediately below the edge acquires a rough texture and forms a low, rounded ridge. These features are all unique to aphanosaurs. [2]

Cervical vertebrae from Teleocrater, showing features characteristic of aphanosaurs. Teleocrater vertebrae.png
Cervical vertebrae from Teleocrater, showing features characteristic of aphanosaurs.

As in other reptiles, aphanosaurian vertebrae also have small structures which articulate with either other vertebrae or the ribs which connect to each vertebra. The structures which connect to vertebrae in front of them are called prezygapophyses, while those that connect to vertebrae behind them are called postzygapophyses. The structures which connect to the ribs also have different names. In most archosaurs, the heads of the ribs are two-pronged. As a result, there are two areas on the side of each vertebra for connecting to a rib: the diapophysis in the upper part of the centrum and the parapophysis in a lower position. However, some cervical ribs are very unusual in aphanosaurs due to possessing a three-pronged head, although this feature only occurs in ribs at the base of the neck. In conjunction with this feature, the vertebrae in that area have a facet for the third prong just above the parapophysis, which has sometimes been classified as a 'divided parapophysis'. [2] The only other archosaurs with this feature were the poposauroids, which explains how Yarasuchus had been mistaken for a poposauroid in the past. [3]

In addition to these features which are unique among avemetatarsalians, aphanosaurs also have a few more traits present in other groups. In vertebrae at the front and middle of the neck, the postzygapophyses have additional small prongs just above the articulating plates. These additional prongs are termed epipophyses, and are common in dinosaurs but likely independently evolved due to being absent in other groups of avemetatarsalians. The body vertebrae have a different type of secondary structure. A small structure (hyposphene) below the postzygapophyses fits into a lip (hypantrum) between the prezygapophyses of the following vertebra, forming additional articulations to assist the zygapophyses. These hyposphene-hypantrum articulations are present in saurischian dinosaurs as well as raisuchids, and are often considered to help make the spine more rigid. [2]

Forelimbs

Aphanosaurs have several characteristic features of the humerus (upper arm bone). This bone was robust, thin when seen from the side but wide when seen from the front. In anterior (front) view, its midshaft was pinched while the proximal (near) and distal (far) ends were wide, making the bone hourglass-shaped. The edge of the upper part of the humerus which faces away from the body has a rounded crest, known as a deltopectoral crest. This crest points forward and is fairly elongated, extending down about a third the length of the bone. Overall, the humerus of aphanosaurs closely resemble that of sauropod dinosaurs and Nyasasaurus, an indeterminate early dinosaur or dinosaur relative. The arm as a whole was robustly-built and somewhat shorter than the leg, but only the humerus possessed unique features. [2] The hand is mostly unknown in members of this group, but it was presumably small and five-fingered as in most archosaurs (apart from specialized forms like pterosaurs or theropod dinosaurs). [4]

Hip and hindlimbs

Pelvic girdle

The pelvis (hip) of aphanosaurs shares many similarities with those of early dinosaurs and silesaurids as well as the unrelated poposauroids. Most of these traits can be found in the ischium, a plank-shaped bone which makes up the lower rear branch of the hip. For example, each ischium (on either side of the hip) contacts each other at the hip's midline. This contact is very extensive, although they are not completely fused due to the contact not extending to the upper edge of each bone. In contrast, pterosaurs, lagerpetids, and Marasuchus (other avemetatarsalians) have their ischia only slightly contact at the middle portion of each bone. The tip of the ischium is also rounded and semi-triangular in cross-section, with the lateral (outer) face of each ischium thinning towards the lower edge of the bone while the medial (inner) face is flat and contacts the other ischium. Poposauroids and dinosaurs also have rounded ischia, but lack the semi-triangular shape, which is also known in Asilisaurus. The ischium also has a groove on the upper part of the shaft. Unlike dinosaurs, aphanosaurs have an acetabulum (hip socket) which is closed up by bone, although perhaps a small portion was open according to a notch near where the ischium contacts the ilium (upper blade of the hip). [2] [1]

Leg

The gracile femur (thigh bone) of aphanosaurs possesses a characteristic set of features which can be used to diagnose the group. The proximal (near) surface of the bone, which connects to the hip socket, has a deep groove on it, rather than simply being a flat articulation surface. In addition, the bone's distal (far) articulation, which connects to the lower leg bones, is concave. The proximal part of the femur also has several bumps (tubers) on either the outer or inner edge of the bone. Many avemetatarsalians have two of these tubers on the inner edge, a small anteromedial tuber in front and a larger posteromedial tuber further back. However, aphanosaurians seem to have completely lost (or never even possessed) the anteromedial tuber. This is nearly unprecedented among archosaurs, but similar to the case in archosaur relatives such as Euparkeria . [2]

A small ridge is present on the inner part of the bone, about a quarter the way down the shaft. This ridge, called a fourth trochanter, is an attachment point for the M. caudofemoralis , a tail muscle which helps to retract the hindlimbs. A scar on the anterolateral (front and outer) edge of the femur may have attached to the M. iliotrochantericus caudalis, a muscle which connects to the hip and helps to stabilize the thigh. This particular scar may be the same thing as the anterior (or lesser) trochanter, a specific structure present in dinosaurs and their close relative. A different scar is located somewhat further back on the bone and lower on the shaft. This scar may have attached to the M. iliofemoralis externus, a muscle which has a similar role to the M. iliotrochantericus caudalis. Likewise, its supposed equivalent in dinosaurs is a structure known as the trochanteric shelf. Aphanosaurs are unique among other avemetatarsalians in the fact that these two scars are separate from each other. In more advanced avemetatarsalians such as dinosaurs, the two structures and their corresponding muscles merge, a condition which is retained in modern birds. [2]

The thin tibia and fibula (lower leg bones) of aphanosaurs do not possess unique traits to the same extent as the femur. However, they are also shorter than the femur. These proportions are rare among early avemetatarsalians, but more common among pseudosuchians and non-archosaur archosauriformes. A short lower leg is inversely correlated with running abilities, indicating that aphanosaurs were not as fast or agile as more advanced members of Avemetatarsalia. [1]

Ankle

The calcanea of Yarasuchus (A) and Teleocrater (B) seen from above, showing 'crocodile-normal' features Aphanosaur calcanea.png
The calcanea of Yarasuchus (A) and Teleocrater (B) seen from above, showing 'crocodile-normal' features

Two different aphanosaurs (Yarasuchus and Teleocrater) each preserve a calcaneum, also known as a heel bone. Most avemetatarsalians have simple calcaneums which are firmly connected to a large bone known as an astragalus next to them. This type of heel, known as the 'advanced mesotarsal' condition, allows for more stability but less flexibility in the foot as it means the different bones of the ankle cannot flex against each other. Pseudosuchians (including modern crocodiles), as well as the crocodile-like phytosaurs have a different configuration, where the calcaneum is much larger and more complex, connecting to the astragalum with a joint that allows for movement between the two. This configuration is called a 'crocodile-normal' ankle, and reptiles which possess it are called crurotarsans. Some recent studies have suggested that phytosaurs are not actually archosaurs, but instead close relatives of the group. [5] This indicates that 'crocodile-normal' ankles were the plesiomorphic (default) state in the first archosaurs, with 'advanced mesotarsal' ankles only later evolving within Avemetatarsalia, rather than at the base of the group. [2]

The calcaneum of aphanosaurs supports this idea, as it more closely resembles that of 'crocodile-normal' ankles than 'advanced mesotarsal' ankles. The calcaneum lies on the outer side of the ankle, with its front or inner edge connecting to the astragalus, the upper surface connecting to the fibula, and the underside connecting to the fourth tarsal (a minor foot bone). In aphanosaurs, the socket for the astragalus is concave while the connection to the fibula manifests as a rounded dome. These are both characteristics of a 'crocodile-normal' ankle. In addition, the rear part of the calcaneum has a cylindrical structure known as a calcaneal tuber. Although this structure is smaller in aphanosaurs than in pseudosuchians, it is still much larger than in other avemetatarsalians, most of which don't even possess the structure. A few dinosauriformes also have small calcaneal tubers, although aphanosaurs have larger and rounder tubers than these taxa ( Marasuchus and a few basal silesaurids). In cross-section, the calcaneal tubers of aphanosaurs are oval-shaped, taller than wide. Most foot material is fragmentary in this group, with only a few phalanges (toe bones) and metatarsals (primary elongated foot bones) known. Based on the length of the preserved metatarsals, the foot was likely rather elongated. [2]

Classification

Aphanosauria is a recently named group, so it has a fairly short taxonomic history. Before it was named, its constituent genera were shuffled around Archosauria and its somewhat larger parent group, Archosauriformes. For example, Yarasuchus was first considered a prestosuchid [6] and later a poposauroid by different analyses, [3] with Martin Ezcurra (2016) placing both it and Dongusuchus as Euparkeria-grade archosaur relatives in his analysis. [7] At the time of these analyses, Teleocrater (the most completely known aphanosaur) was not yet described.

In 2017, Aphanosauria was named and defined by Nesbitt et al. during the formal description of Teleocrater. The description was accompanied by two separate phylogenetic analyses, one derived from Nesbitt (2011)'s [5] broad study on archosaurs and the other from Ezcurra (2016). [7] Both analyses, reapplied with new information, gave a similar result for the position of aphanosaurs. They each placed the group at the base of Avemetatarsalia, outside of Ornithodira (the group containing pterosaurs, dinosaurs, and most other avemetatarsalians). A simplified strict consensus tree (a family tree with the fewest steps in evolution) using the Nesbitt (2011) analysis is given below: [1]

Archosauriformes

Related Research Articles

<span class="mw-page-title-main">Archosaur</span> Group of diapsids broadly classified as reptiles

Archosauria is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term, which includes birds. Extinct archosaurs include non-avian dinosaurs, pterosaurs and extinct relatives of crocodilians. Modern paleontologists define Archosauria as a crown group that includes the most recent common ancestor of living birds and crocodilians, and all of its descendants. The base of Archosauria splits into two clades: Pseudosuchia, which includes crocodilians and their extinct relatives; and Avemetatarsalia, which includes birds and their extinct relatives.

<i>Lagosuchus</i> Genus of fossil bipedal reptile closely related to dinosaurs

Lagosuchus is an extinct genus of avemetatarsalian archosaur from the Late Triassic of Argentina. The type species of Lagosuchus, Lagosuchus talampayensis, is based on a small partial skeleton recovered from the early Carnian-age Chañares Formation. The holotype skeleton of L. talampayensis is fairly fragmentary, but it does possess some traits suggesting that Lagosuchus was a probable dinosauriform, closely related to dinosaurs.

<i>Marasuchus</i> Extinct genus of reptiles

Marasuchus is a genus of basal dinosauriform archosaur which is possibly synonymous with Lagosuchus. Both genera lived during the Late Triassic in what is now La Rioja Province, Argentina. Marasuchus contains a single species, Marasuchus lilloensis.

<i>Nyasasaurus</i> Extinct genus of reptiles

Nyasasaurus is an extinct genus of avemetatarsalian archosaur from the putatively Middle Triassic Manda Formation of Tanzania that may be the earliest known dinosaur. The type species Nyasasaurus parringtoni was first described in 1956 in the doctoral thesis of English paleontologist Alan J. Charig, but it was not formally described until 2013.

<i>Teleocrater</i> Extinct genus of reptiles

Teleocrater is a genus of avemetatarsalian archosaur from the Middle Triassic Manda Formation of Tanzania. The name was coined by English paleontologist Alan Charig in his 1956 doctoral dissertation, but was only formally published in 2017 by Sterling Nesbitt and colleagues. The genus contains the type and only species T. rhadinus. Uncertainty over the affinities of Teleocrater have persisted since Charig's initial publication; they were not resolved until Nesbitt et al. performed a phylogenetic analysis. They found that Teleocrater is most closely related to the similarly enigmatic Yarasuchus, Dongusuchus, and Spondylosoma in a group that was named the Aphanosauria. Aphanosauria was found to be the sister group of the Ornithodira, the group containing dinosaurs and pterosaurs.

<span class="mw-page-title-main">Dinosauromorpha</span> Clade of reptiles

Dinosauromorpha is a clade of avemetatarsalians that includes the Dinosauria (dinosaurs) and some of their close relatives. It was originally defined to include dinosauriforms and lagerpetids, with later formulations specifically excluding pterosaurs from the group. Birds are the only dinosauromorphs which survive to the present day.

<span class="mw-page-title-main">Avemetatarsalia</span> Clade of archosaur reptiles

Avemetatarsalia is a clade of diapsid reptiles containing all archosaurs more closely related to birds than to crocodilians. The two most successful groups of avemetatarsalians were the dinosaurs and pterosaurs. Dinosaurs were the largest terrestrial animals for much of the Mesozoic Era, and one group of small feathered dinosaurs has survived up to the present day. Pterosaurs were the first flying vertebrates and persisted through the Mesozoic before dying out at the Cretaceous-Paleogene (K-Pg) extinction event. Both dinosaurs and pterosaurs appeared in the Triassic Period, shortly after avemetatarsalians as a whole. The name Avemetatarsalia was first established by British palaeontologist Michael Benton in 1999. An alternate name is Pan-Aves, or "all birds", in reference to its definition containing all animals, living or extinct, which are more closely related to birds than to crocodilians.

<i>Turfanosuchus</i> Extinct genus of reptiles

Turfanosuchus is a genus of archosauriform reptile, likely a gracilisuchid archosaur, which lived during the Middle Triassic (Anisian) of northwestern China. The type species, T. dabanensis, was described by C.C. Young in 1973, based on a partially complete but disarticulated fossil skeleton found in the Kelamayi Formation of the Turfan Basin.

Yarasuchus is an extinct genus of avemetatarsalian archosaur that lived during the Anisian stage of the Middle Triassic of India. The genus was named and described in 2005 from a collection of disarticulated but fairly complete fossil material found from the Middle Triassic Yerrapalli Formation. The material is thought to be from two individuals, possibly three, with one being much more complete and articulated than the other. The type and only species is Y. deccanensis. Yarasuchus was a quadruped roughly 2–2.5 metres (6.6–8.2 ft) long, with an elongated neck and tall spines on its vertebrae. Unlike other quadrupedal Triassic reptiles, the limbs and shoulders of Yarasuchus were slender, and more like those of ornithodirans.

<i>Mandasuchus</i> Extinct genus of reptiles

Mandasuchus is an extinct genus of loricatan pseudosuchian from the Manda Formation of Tanzania, which dates back to the Anisian stage of the Middle Triassic. Although this genus was first mentioned by Alan Charig in 1956, a formal description was not published until 2018.

<i>Asilisaurus</i> Genus of reptiles (fossil)

Asilisaurus ; from Swahili, asili, and Greek, σαυρος is an extinct genus of silesaurid archosaur. The type species is Asilisaurus kongwe.Asilisaurus fossils were uncovered in the Manda Beds of Tanzania and date back to the early Carnian, making it one of the oldest known members of the Avemetatarsalia. It was the first non-dinosaurian dinosauriform recovered from Africa. The discovery of Asilisaurus has provided evidence for a rapid diversification of avemetatarsalians during the Middle Triassic, with the diversification of archosaurs during this time previously only documented in pseudosuchians.

<span class="mw-page-title-main">Lagerpetidae</span> Extinct family of reptiles

Lagerpetidae is a family of basal avemetatarsalians. Though traditionally considered the earliest-diverging dinosauromorphs, fossils described in 2020 suggest that lagerpetids may instead be pterosauromorphs. Lagerpetid fossils are known from theTriassic of Argentina, Arizona, Brazil, Madagascar, New Mexico, and Texas. They were typically small, although some lagerpetids, like Dromomeron gigas and a specimen from the Santa Rosa Formation attributed to Dromomeron sp., were able to get quite large. Lagerpetid fossils are rare; the most common finds are bones of the hindlimbs, which possessed a number of unique features.

<span class="mw-page-title-main">Suchia</span> Clade of reptiles

Suchia is a clade of archosaurs containing the majority of pseudosuchians. It was defined as the least inclusive clade containing Aetosaurus ferratus, Rauisuchus tiradentes, Prestosuchus chiniquensis, and Crocodylus niloticus by Nesbitt (2011). Generally the only pseudosuchian group which is omitted from Suchia is the family Ornithosuchidae, although at least one analysis classifies ornithosuchids as close relatives of erpetosuchids and aetosaurs. Phytosaurs are also excluded from Suchia, although it is not certain whether they qualify as pseudosuchians in the first place.

<span class="mw-page-title-main">Poposauroidea</span> Extinct clade of reptiles

Poposauroidea is a clade of advanced pseudosuchians. It includes poposaurids, shuvosaurids, ctenosauriscids, and other unusual pseudosuchians such as Qianosuchus and Lotosaurus. It excludes most large predatory quadrupedal "rauisuchians" such as rauisuchids and "prestosuchids". Those reptiles are now allied with crocodylomorphs in a clade known as Loricata, which is the sister taxon to the poposauroids in the clade Paracrocodylomorpha. Although it was first formally defined in 2007, the name "Poposauroidea" has been used for many years. The group has been referred to as Poposauridae by some authors, although this name is often used more narrowly to refer to the family that includes Poposaurus and its close relatives.

<i>Lutungutali</i> Extinct genus of reptiles

Lutungutali is an extinct genus of silesaurid dinosauriform from the Middle Triassic of Zambia. The single type species of the genus is Lutungutali sitwensis. Lutungutali was named in 2013 and described from a fossil specimen, holotype NHCC LB32, including hip bones and tail vertebrae. The specimen was collected in 2009 from the upper Ntawere Formation, which dates to the Anisian stage of the Middle Triassic. Lutungutali is the first known silesaurid from Zambia and, along with the Tanzanian silesaurid Asilisaurus and dinosauriform Nyasasaurus, the oldest bird-line archosaur known from body fossils.

<i>Asperoris</i> Extinct genus of reptiles

Asperoris is an extinct genus of archosauriform reptile known from the Middle Triassic Manda Beds of southwestern Tanzania. It is the first archosauriform known from the Manda Beds that is not an archosaur. However, its relationships with other non-archosaurian archosauriforms are uncertain. It was first named by Sterling J. Nesbitt, Richard J. Butler and David J. Gower in 2013 and the type species is Asperoris mnyama. Asperoris means "rough face" in Latin, referring to the distinctive rough texture of its skull bones.

Nundasuchus is an extinct genus of crurotarsan, possibly a suchian archosaur related to Paracrocodylomorpha. Remains of this genus are known from the Middle Triassic Manda beds of southwestern Tanzania. It contains a single species, Nundasuchus songeaensis, known from a single partially complete skeleton, including vertebrae, limb elements, osteoderms, and skull fragments.

<i>Kongonaphon</i> Extinct genus of reptiles

Kongonaphon is an extinct genus of lagerpetid avemetatarsalians from the Middle to Late Triassic of Madagascar. It contains a single species, Kongonaphon kely, which is known from a fragmentary partial skeleton. This fossil hails from the late Ladinian or early Carnian-age "basal Isalo II beds". As the first lagerpetid found in Africa, Kongonaphon extends the range of the family significantly. It possessed a combination of features from various other lagerpetids, but developed particularly long and slender leg bones. Kongonaphon is also the first lagerpetid for which fossils of the snout and teeth are known. It was likely an insectivore based on the shape and texture of its teeth.

<i>Incertovenator</i> Extinct genus of probable archosaur

Incertovenator is an extinct genus of archosauriform reptile, likely an archosaur, of uncertain affinities. Its unstable position is a result of possessing a number features found in both the bird-line avemetatarsalian archosaurs and the crocodylian-line pseudosuchians. The type and only known species is I. longicollum, which is known from single specimen discovered in the Late Triassic Ischigualasto Formation of Argentina. Incertovenator is known almost entirely by its vertebral column. This indicates that it had a relatively long neck, leading to its uncertain classification due to the convergent evolution of elongated neck vertebrae in both avemetatarsalian and pseudosuchian archosaurs.

Mambachiton is an extinct genus of basal avemetatarsalian from the Middle/Upper Triassic Makay Formation of Madagascar. The genus contains a single species, M. fiandohana, known from a partial skeleton with articulated osteoderms.

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