Scleral ring
{{Short description|Ring of bone supporting the eye}}
File:Coloradisaurus skull reconstruction.png showing its sclerotic ring]]
The scleral ring or sclerotic ring is a hardened ring of plates, often derived from bone, that is found in the eyes of many animals in several groups of vertebrates. Mammals, amphibians, and crocodilians lack scleral rings.{{cite web |last=Motani |first=Ryosuke |title=Eyes of Ichthyosaurs|date=15 November 2001|url=http://www.ucmp.berkeley.edu/people/motani/ichthyo/eyes.html |archive-url=https://web.archive.org/web/20011217152147/http://www.ucmp.berkeley.edu/people/motani/ichthyo/eyes.html |url-status=dead |archive-date=17 December 2001 |publisher=UC Museum of Paleontology|accessdate=15 October 2013}} The ring is in the fibrous outer layer of the eye, called the sclera. 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 material (scleral cartilage) or bony material (scleral ossicles), or often a combination of both, that comes together to form a ring.{{Cite journal |last1=Atkins |first1=Jade B. |last2=Franz-Odendaal |first2=Tamara A. |date=October 2016 |title=The sclerotic ring of squamates: an evo-devo-eco perspective |journal=Journal of Anatomy |language=en |volume=229 |issue=4 |pages=503–513 |doi=10.1111/joa.12498|doi-access=free |pmid=27240556 |pmc=5013065 }} The arrangement, size, shape, and number of ossicles vary by group.{{Cite journal |last=Franz-Odendaal |first=Tamara Anne |date=December 2018 |title=Skeletons of the Eye: An Evolutionary and Developmental Perspective |journal=The Anatomical Record |language=en |volume=303 |issue=1 |pages=100–109 |doi=10.1002/ar.24043 |issn=1932-8486|doi-access=free |pmid=30548203 }} They are believed to have a role in supporting the eye, especially in animals whose eyes are not spherical, or which live underwater. Fossil scleral rings are known for a variety of extinct animals, including ichthyosaurs, pterosaurs, and non-avian dinosaurs,{{cite web |last=Pigdon |first=Dann |url=http://dml.cmnh.org/1997Oct/msg00393.html |title=Re: Sclerotic ring in eyes |accessdate=2007-07-06}}{{cite web|last=Milner|first=Angela|title=Ophthalmosaurus icenicus: Why did it have such large eyes?|url=http://www.nhm.ac.uk/nature-online/species-of-the-day/evolution/ophthalmosaurus%20icenicus/eyes/index.html|publisher=Natural History Museum, London|accessdate=15 October 2013}} 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, lizards, tuatara, and relatives), scleral rings have been found in all major lineages except Serpentes, or snakes, and two families within Anguimorpha: Dibamidae and Rhineuridae, which are both legless lizard families. All of these clades 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 (turtles, birds, crocodilians, and relatives), only birds and turtles retain the scleral rings. Fossil evidence shows that extinct marine crocodiles living in the Mesozoic had scleral rings, so the trait was lost over time.{{Cite book |last=Walls |first=Gordon L. |url=http://dx.doi.org/10.5962/bhl.title.7369 |title=The vertebrate eye and its adaptive radiation [by] Gordon Lynn Walls. |date=1942 |publisher=Cranbrook Institute of Science |location=Bloomfield Hills, Mich.|doi=10.5962/bhl.title.7369 }} Scleral rings of varying lengths, curvatures, numbers of ossicles, and thickness are found in all birds.{{Cite journal |last=Hall |first=Margaret I. |date=June 2008 |title=The anatomical relationships between the avian eye, orbit and sclerotic ring: implications for inferring activity patterns in extinct birds |journal=Journal of Anatomy |language=en |volume=212 |issue=6 |pages=781–794 |doi=10.1111/j.1469-7580.2008.00897.x |pmid=18510506 |issn=0021-8782|pmc=2423400 }} Birds typically have 12-18 ossicles, with 14 being the most common number.
= Fish =
While all fish have scleral cartilage, teleost fish are the only family to retain scleral rings, with the rings being absent in the more basal clades Cladistia, Chondrostei, Lepisosteiformes, and Amiiformes.
Teleost fish typically have only one or two ossicles per ring, and fish with no ossicles still retain cartilage.{{Cite journal |last=Franz-Odendaal |first=Tamara A. |date=22 January 2008 |title=Scleral Ossicles of Teleostei: Evolutionary and Developmental Trends |url=https://onlinelibrary.wiley.com/doi/10.1002/ar.20639 |journal=The Anatomical Record: Advances in Integrative Anatomy and Evolutionary Biology |language=en |volume=291 |issue=2 |pages=161–168 |doi=10.1002/ar.20639|pmid=18213703 |url-access=subscription }} 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
File:Mosasaurus_sp_sclerotic_ring.JPG|Partial scleral ring of Mosasaurus.
File:Viatkogorgon skull.png|Skull and scleral ring of the gorgonopsian therapsid Viatkogorgon.
File:Prosaurolophus maximus3.JPG|Scleral ring of the hadrosaur Prosaurolophus.
References
{{Portal|Paleontology|Anatomy}}
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External links
{{Commonscat|Sclerotic rings}}
{{Tetrapod osteology|S.}}