\cite{Blevins_2012} - Rajiform locomotion: three-dimensional kinematics of the pectoral fin surface during swimming in the freshwater stingray Potamotrygon orbignyi - Rajiform locomotion in fishes is dominated by distinctive undulations of expanded pectoral fins. Unlike other fishes, which typically interact with the fluid environment via multiple fins, undulating rays modulate a single control surface, the pectoral disc, to perform pelagic locomotion, maneuvering and other behaviors.
\cite{Nakamura_2015} - Molecular mechanisms underlying the exceptional adaptations of batoid fins - With pectoral fins that surround much of the body, by fusing to the head, the skate is a cartilaginous fish that has one of the most unique appendages of all vertebrates. Here, we use an unbiased RNA screen to uncover genetic pathways underlying this morphology. Unlike tetrapods and other fishes, skates induce a second growth center in the anterior region, by the redeployment of an ancient genetic module. We find that some of the genes involved in generating the anterior–posterior fin function differently in skates than they do in limbed animals. Our data reveal the mechanisms for the unique skate fin morphology and also provide insights into the genetic origins of fin variation and morphological innovation in paired appendages.
\cite{Barry_2017} - The role of HoxA11 and HoxA13 in the evolution of novel fin morphologies in a representative batoid (Leucoraja erinacea) - Background:
Batoids exhibit unique body plans with derived fin morphologies, such as the anteriorly expanded pectoral fins that fuse to the head, or distally extended anterior pelvic fin lobes used for a modified swimming technique utilized by skates (Rajidae). The little skate (Leucoraja erinacea), exhibits both of these unique fin morphologies. These fin modifications are not present in a typical shark body plan, and little is known regarding the mechanisms underlying their development. A recent study identified a novel apical ectodermal ridge (AER) associated with the development of the anterior pectoral fin in the little skate, but the role of the posterior HoxA genes was not featured during skate fin development.
Results:
We present the first evidence for HoxA expression (HoxA11 and HoxA13) in novel AER domains associated with the development of three novel fin morphologies in a representative batoid, L. erinacea. We found HoxA13 expression associated with the recently described novel AER in the anterior pectoral fin, and HoxA11 expression in a novel AER domain in the anterior pelvic fin that we describe here. We find that both HoxA11 and HoxA13 are expressed in claspers, and while HoxA11 is expressed in pelvic fins and claspers, HoxA13 is expressed exclusively in developing claspers of males. Finally, HoxA11 expression is associated with the developing fin rays in paired fins.
Conclusion:
Overall, these results indicate that the posterior HoxA genes play an important role in the morphological evolution of paired fins in a representative batoid. These data suggest that the batoids utilize a unique Hox code, where the posterior HoxA genes exhibit distinct expression patterns that are likely associated with specification of novel fin morphologies.
\cite{Martinez_2016} - Re-evaluation of batoid pectoral morphology reveals novel patterns of diversity among major lineages. - Batoids (Chondrichthyes: Batoidea) are a diverse group of cartilaginous fishes which comprise a monophyletic sister lineage to all neoselachians or modern sharks. All species in this group possess anteroposteriorly expanded-pectoral fins, giving them a unique disc-like body form. Reliance on pectoral fins for propulsion ranges from minimal (sawfish) to almost complete dependence (skates and rays). A recent study on the diversity of planform pectoral fin shape in batoids compared overall patterns of morphological variation within the group. However, inconsistent pectoral homology prevented the study from accurately representing relationships within and among major batoid taxa. With previous work in mind, we undertook an independent investigation of pectoral form in batoids and evaluated the implications of shape diversity on locomotion and lifestyle, particularly in the skates (Rajoidei) and rays (Myliobatoidei). We used geometric morphometrics with sliding semilandmarks to analyze pectoral fin outlines and also calculate fin aspect ratios (AR), a functional trait linked to locomotion. In agreement with previous work, our results indicated that much of the evolution of batoid pectoral shape has occurred along a morphological axis that is closely related to AR. For species where kinematic data were available, both shape and AR were associated with swimming mode. This work further revealed novel patterns of shape variation among batoids, including strong bimodality of shape in rays, an intermediate location of skate species in the morphospace between benthic/demersal and pelagic rays, and approximately parallel shape trajectories in the benthic/demersal rays and skates. Finally, manipulation of landmarks verified the need for a consistent and accurate definition of homology for the outcome and efficacy of analyses of pectoral form and function in batoids.
\cite{Franklin_2014} - Pectoral fin morphology of batoid fishes (Chondrichthyes: Batoidea): explaining phylogenetic variation with geometric morphometrics. - The diverse cartilaginous fish lineage, Batoidea (rays, skates, and allies), sister taxon to sharks, comprises a huge range of morphological diversity which to date remains unquantified and unexplained in terms of evolution or locomotor style. A recent molecular phylogeny has enabled us to confidently assess broadscale aspects of morphology across Batoidea. Geometric morphometrics quantifies the major aspects of shape variation, focusing on the enlarged pectoral fins which characterize batoids, to explore relationships between ancestry, locomotion and habitat. A database of 253 specimens, encompassing 60 of the 72 batoid genera, reveals that the majority of morphological variation across Batoidea is attributable to fin aspect-ratio and the chordwise location of fin apexes. Both aspect-ratio and apex location exhibit significant phylogenetic signal. Standardized independent linear contrast analysis reveals that fin aspect-ratio can predict locomotor style. This study provides the first evidence that low aspect-ratio fins are correlated with undulatory-style locomotion in batoids, whereas high aspect-ratio fins are correlated with oscillatory locomotion. We also show that it is phylogeny that determines locomotor style. In addition, body- and caudal fin-locomotors are shown to exhibit low aspect-ratio fins, whereas a pelagic lifestyle correlates with high aspect-ratio fins. These results emphasize the importance of phylogeny in determining batoid pectoral fin shape, however, interactions with other constraints, most notably locomotor style, are also highlighted as significant.
\cite{Huang_2017} -
Structure and mechanical implications of the pectoral fin skeleton in the Longnose Skate (Chondrichthyes, Batoidea) -
Animal propulsion systems are believed to show high energy and mechanical efficiency in assisting movement compared to artificial designs. As an example, batoid fishes have very light cartilaginous skeletons that facilitate their elegant swimming via enlarged wing-like pectoral fins. The aim of this work is to illustrate the hierarchical structure of the pectoral fin of a representative batoid, the Longnose Skate (Raja rhina), and explain the mechanical implications of its structural design. At the macro level, the pectoral fins are comprised of radially oriented fin rays, formed by staggered mineralized skeletal elements stacked end-to-end. At the micro level, the midsection of each radial element is composed of three mineralized components, which consist of discrete segments (tesserae) that are mineralized cartilage and embedded in unmineralized cartilage. The radial elements are wrapped with aligned, unmineralized collagen fibers. This is the first report of the detailed structure of the ray elements, including the observation of a 3-chain mineralized tesserae. Structural analyses demonstrate that this configuration enhances stiffness in multiple directions. A two-dimensional numerical model based on the morphological analysis demonstrated that the tessera structure helps distributing shear, tensile and compressive stress more ideally, which can better support both lift and thrust forces when swimming without losing flexibility. \cite{Di_Santo_2016} - Batoid locomotion: effects of speed on pectoral fin deformation in the little skate, Leucoraja erinacea. - Most batoids have a unique swimming mode in which thrust is generated by either oscillating or undulating expanded pectoral fins that form a disc. Only one previous study of the freshwater stingray has quantified three-dimensional motions of the wing, and no comparable data are available for marine batoid species that may differ considerably in their mode of locomotion. Here, we investigate three-dimensional kinematics of the pectoral wing of the little skate, Leucoraja erinacea, swimming steadily at two speeds [1 and 2 body lengths (BL) s-1]. We measured the motion of nine points in three dimensions during wing oscillation and determined that there are significant differences in movement amplitude among wing locations, as well as significant differences as speed increases in body angle, wing beat frequency and speed of the traveling wave on the wing. In addition, we analyzed differences in wing curvature with swimming speed. At 1 BL s-1, the pectoral wing is convex in shape during the downstroke along the medio-lateral fin midline, but at 2 BL s-1 the pectoral fin at this location cups into the flow, indicating active curvature control and fin stiffening. Wing kinematics of the little skate differed considerably from previous work on the freshwater stingray, which does not show active cupping of the whole fin on the downstroke.
\cite{Aschliman_2012} - Body plan convergence in the evolution of skates and rays (Chondrichthyes: Batoidea) - Skates, rays and allies (Batoidea) comprise more than half of the species diversity and much of the morphological disparity among chondrichthyan fishes, the sister group to all other jawed vertebrates. While batoids are morphologically well characterized and have an excellent fossil record, there is currently no consensus on the interrelationships of family-level taxa. Here we construct a resolved, robust and time-calibrated batoid phylogeny using mitochondrial genomes, nuclear genes, and fossils, sampling densely across taxa. Data partitioning schemes, biases in the sequence data, and the relative informativeness of each fossil are explored. The molecular phylogeny is largely congruent with morphology crownward in the tree, but the branching orders of major batoid groups are mostly novel. Body plan convergence appears to be widespread in batoids. A depressed, rounded pectoral disk supported to the snout tip by fin radials, common to skates and stingrays, is indicated to have been derived independently by each group, while the long, spiny rostrum of sawfishes similarly appears to be convergent with that of sawsharks, which are not batoids. The major extant batoid lineages are inferred to have arisen relatively rapidly from the Late Triassic into the Jurassic, with long stems followed by subsequent radiations in each group around the Cretaceous/Tertiary boundary. The fossil record indicates that batoids were affected with disproportionate severity by the end-Cretaceous extinction event.
\cite{LUCIFORA_2002} - Walking in skates (Chondrichthyes, Rajidae): anatomy, behaviour and analogies to tetrapod locomotion - Pelvic fin walking in skates is common. However, the structure and function of pelvic fins have not been analysed. Pelvic fins of skates of the genus Psammobatis and Rioraja agassizi are externally divided into an anterior leg-like lobe and a posterior fin-like lobe. Internally, the anterior lobes are supported by a compound radial, a proximal radial and distal radials that resemble a thigh, a calf and a foot, respectively, and three associated radials arising from the pelvic girdle. A highly developed radial condyle on the pelvic girdle enables broad ‘limb’ movements. The muscular arrangement of the anterior lobes is formed by protractor, retractor, flexor and extensor muscles, clearly departing from the generalized fin muscle arrangement of elasmobranchs. Walking is composed of propulsion and recovery phases. A backward movement of the compound radial in the horizontal plane characterizes the propulsive phase. The proximal radial connects vertically the compound radial with the foot-like distal radials, which are anchored on the bottom. During the recovery phase, the foot-like structure is lifted off the bottom and the entire limb-like anterior lobe is moved forwards for starting a new cycle. Walking in skates resembles the ancestral tetrapod sprawling locomotion seen in many salamanders and lizards. Pelvic fin anatomy and walking behaviour in skates and hemiscylliid sharks are compared. Ecological and evolutionary implications of walking locomotion in skates are also discussed.