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The scleral ring or sclerotic ring is a hardened ring of plates, often derived from
bone A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, ...
, that is found in the eyes of many
animal Animals are multicellular, eukaryotic organisms in the Biology, biological Kingdom (biology), kingdom Animalia (). With few exceptions, animals heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, ...
s in several groups of
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
s. Some species of
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s,
amphibian Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s, and
crocodilia Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
ns lack scleral rings. The ring is in the fibrous outer layer of the eye, called the
sclera The sclera, also known as the white of the eye or, in older literature, as the tunica albuginea oculi, is the opaque, fibrous, protective outer layer of the eye containing mainly collagen and some crucial elastic fiber. In the development of t ...
. The structure is commonly referred to as the sclerotic ring; but, because the word ''sclerotic'' often implies pathology of the sclera (''see'' " sclerosis", an unrelated medical condition), recent authors have urged avoiding the use of this term, to avoid confusion and to increase the utility of character comparisons. Scleral rings can be made of
cartilaginous Cartilage is a resilient and smooth type of connective tissue. Semi-transparent and non-porous, it is usually covered by a tough and fibrous membrane called perichondrium. In tetrapods, it covers and protects the ends of long bones at the joints ...
material (scleral cartilage) or bony material (scleral ossicles), or often a combination of both, that comes together to form a ring. The arrangement, size, shape, and number of ossicles vary by group. They are believed to have a role in supporting the eye, especially in animals whose eyes are not spherical, or which live underwater.
Fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
scleral rings are known for a variety of
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
animals, including ichthyosaurs,
pterosaur Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosaurs are the earli ...
s, and non-avian
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s, but are often not preserved. Scleral rings may help support inner structures of the eye, especially in animals that do not have round eyes. Animals that move rapidly, including both fast flying birds and fast swimming fish have the most robust scleral rings, indicating that these thick rings are used to protect the eye during intense changes in pressure in the air and in the water. Additionally, scleral rings may help the eye adjust to different viewing distances, also known as visual accommodation. Muscles are used to adjust the shape of the eye for accommodation, and the rings provide attachment sites for these muscles. In aquatic animals, the lens is squeezed in a different way to compensate for differences in light refraction underwater, and so the shape of the ring can be different than those in terrestrial animals.


Extant Animals


Reptiles

A combination of scleral cartilage and ossicles are present, in which the cartilage acts as a cup around the posterior (rear) position of the eye and ossicles at the anterior (front) position of the eye form the ring. Within Lepidosaurs (
snakes Snakes are elongated Limbless vertebrate, limbless reptiles of the suborder Serpentes (). Cladistically Squamata, squamates, snakes are ectothermic, amniote vertebrates covered in overlapping Scale (zoology), scales much like other members of ...
,
lizards Lizard is the common name used for all squamate reptiles other than snakes (and to a lesser extent amphisbaenians), encompassing over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The ...
,
tuatara The tuatara (''Sphenodon punctatus'') is a species of reptile endemic to New Zealand. Despite its close resemblance to lizards, it is actually the only extant member of a distinct lineage, the previously highly diverse order Rhynchocephal ...
, and relatives), scleral rings have been found in all major lineages except Serpentes, or snakes, and two families within Anguimorpha:
Dibamidae Dibamidae or blind skinks is a family of lizards characterized by their elongated cylindrical body and an apparent lack of limbs. Female dibamids are entirely limbless and the males retain small flap-like hind limbs, which they use to grip their ...
and Rhineuridae, which are both legless lizard families. All of these
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
s that lack a scleral ring share either a burrowing lifestyle or lack of limbs, indicating a possible correlation among these traits and loss of the scleral ring. Lizards typically have 14 ossicles in the ring, though this can vary. Within Archelosauria (
turtle Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
s,
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s, crocodilians, and relatives), only birds and turtles retain the scleral rings. Fossil evidence shows that extinct marine crocodiles living in the
Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...
had scleral rings, so the trait was lost over time. Scleral rings of varying lengths, curvatures, numbers of ossicles, and thickness are found in all birds. Birds typically have 12-18 ossicles, with 14 being the most common number.


Fish

While all fish have scleral cartilage,
Teleost Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all neontology, extant species of f ...
fish are the only family to retain scleral rings, with the rings being absent in the more basal clades Cladistia, Chondrostei, Lepisosteiformes, and
Amiiformes The Amiiformes order (biology), order of fish has only two extant taxa, extant species, the bowfins: ''Amia calva'' and ''Amia ocellicauda'', the latter recognized as a separate species in 2022. These Amiiformes are found in the freshwater syste ...
. Teleost fish typically have only one or two ossicles per ring, and fish with no ossicles still retain cartilage. Most teleosts do not have ossicles, but this can vary even within groups. As a general trend, more basal groups (such as Elopomorpha and Osteoglossomorpha) tend to have no ossicles, while more derived groups (such as Percomorpha) are likely to have a variable number of ossicles (zero to two). More active fish are more likely to have scleral rings, indicating that the rings help keep the eye stable during rapid swimming.


Gallery

File:Tawny Frogmouth Skull.jpg, A skull of an extant tawny frogmouth, showing large scleral rings. File:Uroplatus_phantasticus_skull1.jpg, A skull of an extant satanic leaf-tailed gecko, showing large scleral rings. File:Ophthalmosaurus icenicus sclerotic ring.jpg, Virtually complete scleral ring of the ichthyosaur ''
Ophthalmosaurus ''Ophthalmosaurus'' (Greek ὀφθάλμος ''ophthalmos'' 'eye' and σαῦρος ''sauros'' 'lizard') is a genus of ichthyosaur known from the Middle-Late Jurassic. Possible remains from the earliest Cretaceous, around 145 million years ago, a ...
'' File:Mosasaurus_sp_sclerotic_ring.JPG, Partial scleral ring of ''
Mosasaurus ''Mosasaurus'' (; "lizard of the Meuse (river), Meuse River") is the type genus (defining example) of the mosasaurs, an extinct group of aquatic Squamata, squamate reptiles. It lived from about 82 to 66 million years ago during the Campanian an ...
''. File:Viatkogorgon skull.png, Skull and scleral ring of the gorgonopsian therapsid '' Viatkogorgon''. File:Prosaurolophus maximus3.JPG, Scleral ring of the hadrosaur '' Prosaurolophus''.


References

{{Tetrapod osteology, S. Dinosaur anatomy Eye Skull bones