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. 2015 Feb 18;10(2):e0117079.
doi: 10.1371/journal.pone.0117079. eCollection 2015.

Sclerotic rings in mosasaurs (Squamata: Mosasauridae): structures and taxonomic diversity

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Sclerotic rings in mosasaurs (Squamata: Mosasauridae): structures and taxonomic diversity

Momo Yamashita et al. PLoS One. .

Abstract

Mosasaurs (Squamata: Mosasauridae) were a highly diverse, globally distributed group of aquatic lizards in the Late Cretaceous (98-66 million years ago) that exhibited a high degree of adaptation to life in water. To date, despite their rich fossil record, the anatomy of complete mosasaur sclerotic rings, embedded in the sclera of the eyeball, has not been thoroughly investigated. We here describe and compare sclerotic rings of four mosasaur genera, Tylosaurus, Platecarpus, Clidastes, and Mosasaurus, for the first time. Two specimens of Tylosaurus and Platecarpus share an exact scleral ossicle arrangement, excepting the missing portion in the specimen of Platecarpus. Furthermore, the exact arrangement and the total count of 14 ossicles per ring are shared between Tylosaurus and numerous living terrestrial lizard taxa, pertaining to both Iguania and Scleroglossa. In contrast, two species of Mosasaurus share the identical count of 12 ossicles and the arrangement with each other, while no living lizard taxa share exactly the same arrangement. Such a mosaic distribution of these traits both among squamates globally and among obligatorily aquatic mosasaurs specifically suggests that neither the ossicle count nor their arrangement played major roles in the aquatic adaptation in mosasaur eyes. All the mosasaur sclerotic rings examined consistently exhibit aperture eccentricity and the scleral ossicles with gently convex outer side. Hitherto unknown to any squamate taxa, one specimen of Platecarpus unexpectedly shows a raised, concentric band of roughened surface on the inner surface of the sclerotic ring. It is possible that one or both of these latter features may have related to adaptation towards aquatic vision in mosasaurs, but further quantitative study of extant reptilian clades containing both terrestrial and aquatic taxa is critical and necessary in order to understand possible adaptive significances of such osteological features.

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Conflict of interest statement

Competing Interests: The authors have declared that no competing interests exist.

Figures

Figure 1
Figure 1. Sclerotic rings of five specimens of mosasaurs representing four genera.
All showing left ring in outer view, where vertical arrows point dorsally and horizontal ones anteriorly. A, Tylosaurus proriger (FFHM 1997–10); B, Platecarpus tympaniticus (LACM 128319); C, P. tympaniticus (KU 1001); D, Clidastes propython (CFDC M74.10.06); E. Mosasaurus sp., cf. M. missouriensis (TMP 2012.010.0001). Abbreviations: f, frontal; j, jugal; os, orbitosphenoid; pra, prearticular; pof, postorbital frontal; prf, prefrontal. Scale bars equal 1 cm.
Figure 2
Figure 2. Structure of a generalized reptilian eye.
A, Cross section of a lizard eye; B, Corneal section of an eye. Abbreviations: Bm, Brücke’s muscle; c, cornea; Cm, Crampton’s muscle; cp, corneal process; i, iris; IED, distance between internal and external borders of sclerotic ring; l, lens; s, sclera; sr, sclerotic ring; sm, sphincter muscle; ss, sulcus.
Figure 3
Figure 3. Observed arrangement of scleral ossicles in mosasaurs and a modern monitor lizard.
A, Tylosaurus proriger (FFHM 1997–10, outer view of left ring); B, Platecarpus tympaniticus (LACM 128319, ditto); C, Clidastes propython (CFDC M74.10.06, ditto); D, Mosasaurus sp., cf. M. missouriensis (TMP 2012.010.0001, ditto); E, Mosasaurus sp., cf. M. hoffmannii (NHMM 2013001, ditto); F, Varanus dumerilii (NSM PO 391, ditto). Dark gray, light gray, and white indicate the ossicles of the minus-, the plus- and the imbricating-types, respectively. Except for final ossicle, only the plus- and the minus-type are numbered. Cross-hatching indicates broken parts. See main text for further explanation. Figures are not to scale. Abbreviations: d, dorsal; n, nasal; t, temporal; v, ventral.
Figure 4
Figure 4. Difference of surface conditions between inner and outer side of the sclerotic ring in Platecarpus tympaniticus.
A, interpretative drawing of inner side of the sclerotic ring based on KU 1001; B, cross sectional view of a single scleral ossicle along the line I-I’ in A; C, cross sectional view of a boundary of adjacent scleral ossicles along the line II-II’ in A. On the inner side, the raised middle area shaded in blue is roughened/asperate, resulting in obliterating ossicle boundaries.
Figure 5
Figure 5. Sclerotic ring of a modern monitor lizard.
A, outer surface; B, inner surface; C, lateral view. White arrow indicates the sigmoid flexure that supports a circular depression/sulcus which surrounds the cornea of the lizard eyeball.

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