ArticleLiterature Review

Comparative anatomy of the extraocular muscles in four Myliobatoidei rays (Batoidea, Myliobatiformes)

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Abstract

Extraocular muscles are classically grouped as four rectus and two oblique muscles. However, their description and potential associations with species behavior are limited. The objective was to characterize extraocular muscles in four Myliobatoidei rays from diverse habitats with divergent behaviors. Heads (10 per species) of Dasyatis hypostigma, Gymnura altavela, Mobula thurstoni and Pteroplatytrygon violacea were decalcified and dissected to characterize and describe extraocular muscles. Principal component analysis (PCA) was used to evaluate relationships between muscle length and species; for P. violacea, D. hypostigma and G. altavela, these were qualitatively and quantitatively consistent with the general pattern of extraocular muscles in vertebrates. In contrast, for M. thurstoni, the two oblique muscles were completely fused and there was a seventh extraocular muscle, named m. lateral rectus b (both were apparently novel findings in this species). There were also significant differences in eye disposition in the chondrocranium. The PCA axis 1 (rectus muscles) and PCA axis 2 (oblique muscles) accounted for 98.47% of data variability. Extraocular muscles had significant differences in length and important anatomical differences among sampled species that facilitated grouping species according to their life history. In conclusion, extraocular muscles are not uniform in all vertebrate species, thereby providing another basis for comparative studies.

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... Similar to mammals, in fish, the eye muscles are responsible for the movement of the eye. The eye muscles are not uniform among species and vary in number and shape according to differences in fish species' habitats; however, information on the eye muscles in different fish species is scarce (Cunha, Oliveira, & Kfoury Jr, 2016). Generally, there are four rectus and two oblique muscles; during eyeball movement, the inferior and superior rectus muscles are responsible for movement around the horizontal axis of the eyeball, while the medial and lateral rectus muscles are responsible for movement around the vertical axis. ...
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The strict consensus reveals the following hierarchical structure: Hexatrygon + (†Asterotrygon, n.gen., Plesiobatis, Urolophidae + (Urotrygonidae + (†Heliobatis + (Potamotrygonidae + (amphi-American Himantura, Pteroplatytrygon, Himantura, Taeniura, Dasyatis + (Gymnuridae + Myliobatidae)))))). Our resulting tree has nodes in common with previous phylogenetic analyses of stingrays (e.g., Hexatrygon is the most basal stingray genus; gymnurids and myliobatids [pelagic stingrays] are well-supported sister-groups), but includes novel components, such as a clade that includes all dasyatid genera (as a polytomy) and the component Gymnuridae + Myliobatidae. “Dasyatidae” is not monophyletic in any of the minimum-length trees obtained; Urolophidae (Urolophus and Trygonoptera) and Urotrygonidae (Urobatis and Urotrygon) are both monophyletic, but are not sister-groups. †Asterotrygon, n.gen. forms a clade with urolophids in 21 of the 35 equally most parsimonious trees. Successive approximations weighting adds only one additional node in relation to the strict consensus, which unites Pteroplatytrygon, Dasyatis, and Himantura sensu stricto (in a polytomy) with Gymnuridae + Myliobatidae. The resulting stingray phylogeny is at odds with previous phylogenies mostly regarding the affinities of amphi-American Himantura and Taeniura, which do not form a monophyletic group with the South American freshwater stingrays (Potamotrygonidae) in any of the minimum-length trees obtained. Similar to most elasmobranch groups, stingrays display much character conflict, and cladogram topologies are very sensitive to changes in character coding. Due to a high degree of character variation present in certain generic-level terminal taxa, a more fully representative species-level phylogeny is necessary to clarify the systematic importance of tail-fold configuration, ceratobranchial fusion patterns, and other characters discussed in our study. Three additional synapomorphies of stingrays were uncovered by our study, pertaining to the configuration of the basihyal, first pair of hypobranchial cartilages, and to the forward extension of the basibranchial copula. Our phylogenetic results imply the following biogeographic patterns: the relationships of †Asterotrygon, n.gen. demonstrate a strong Indo-west Pacific historical correlation, while †Heliobatis displays an affinity with the Americas; the node containing the greatest diversity of modern stingrays (“Dasyatidae” + (Gymnuridae + Myliobatidae)) evolved only after an American stingray lineage was established sometime earlier than the early Eocene; and potamotrygonids date at least from the late early Eocene, and not the Miocene, as previous studies have implied. The mechanism responsible for the invasion of the potamotrygonid ancestor into South America could indeed have been a marine transgression as advocated by other authors, albeit a much earlier (pre-Miocene) one, during either the Late Cretaceous or the late Paleocene to early Eocene.
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