The Labridae (including wrasses, the Odacidae and the Scaridae) is a species-rich group of perciform fishes whose members are prominent inhabitants of warm-temperate and tropical reefs worldwide. We analyse functionally relevant morphometrics for the feeding apparatus of 130 labrid species found on the Great Barrier Reef and use these data to explore the morphological and mechanical basis of trophic diversity found in this assemblage. Morphological measurements were made that characterize the functional and mechanical properties of the oral jaws that are used in prey capture and handling, the hyoid apparatus that is used in expanding the buccal cavity during suction feeding, and the pharyngeal jaw apparatus that is used in breaking through the defences of shelled prey, winnowing edible matter from sand and other debris, and pulverizing the algae, detritus and rock mixture eaten by scarids (parrotfishes). A Principal Components Analysis on the correlation matrix of a reduced set of ten variables revealed complete separation of scarids from wrasses on the basis of the former having a small mouth with limited jaw protrusion, high mechanical advantage in jaw closing, and a small sternohyoideus muscle and high kinematic transmission in the hyoid four-bar linkage. Some scarids also exhibit a novel four-bar linkage conformation in the oral jaw apparatus. Within wrasses a striking lack of strong associations was found among the mechanical elements of the feeding apparatus. These weak associations resulted in a highly diverse system in which functional properties occur in many different combinations and reflect variation in feeding ecology. Among putatively monophyletic groups of labrids, the cheilines showed the highest functional diversity and scarids were moderately diverse, in spite of their reputation for being trophically monomorphic and specialized. We hypothesize that the functional and ecological diversity of labrids is due in part to a history of decoupled evolution of major components of the feeding system (i.e. oral jaws, hyoid and pharyngeal jaw apparatus) as well as among the muscular and skeletal elements of each component.
The feeding mechanisms of two labrid fishes (Cheilinus chlorurus and C. diagrammus: Labridae: Perciformes) are modeled using four-bar linkage theory from mechanical engineering. The actions of the feeding mechanisms are simulated by a computer program that uses morphometric data to calculate the geometry of mechanism structure. The predictions of three different four-bar linkages regarding the kinematics of feeding are compared to the movements observed through hign speed (200 fps) cinematography. A previously unidentified four-bar chain was found to be an accurate model of the mechanism by which upper jaw protrusion, maxillary rotation, and gape increase occur in Cheilinus. This mechanism involves the anterior jaws including the mandible, maxilla, premaxilla, palatine, and suspensorium. The accuracy of two previously described four-bar linkages was also tested by comparison of model predictions and film results. The opercular linkage proposed by Anker ('74) as a mechanism of jaw depression via opercular levation was found to be a poor predictor of feeding movements. This four-bar chain involves the opercle, suspensorium, interopercle, and mandible. Muller ('87) proposed a mechanism of hyoid depression involving cranial elevation due to epaxial muscle contraction as input motion The links in this mechanism include the neurocranium and hyomandibula, hyoid, sternohyoideus muscle, and pectoral girdle. This model was an accurate predictor of hyoid depression in Cheilinus when simultaneous cranial elevation and sternohyoideus contraction were simulated. Quantitative kinematic models involve simplifying assumptions when applied to complex musculoskeletal systems, but such models have a wide range of applications to vertebrate functional morphology.
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