SUMMARYCetaceans are thought to display a diversity of feeding modes that are often described as convergent with other more basal aquatic vertebrates (i.e. actinopterygians). However, the biomechanics of feeding in cetaceans has been relatively ignored by functional biologists. This study investigated the feeding behavior, kinematics and pressure generation of three odontocetes with varying feeding modes (belugas, Delphinapterus leucas; Pacific white-sided dolphins, Lagenorhynchus obliquidens; and longfinned pilot whales, Globicephala melas). Four feeding phases were recognized in all odontocetes: (I) preparatory, (II) jaw opening, (III) gular depression, and (IV) jaw closing. Belugas relied on a feeding mode that was composed of discrete ram and suction components. Pacific white-sided dolphins fed using ram, with some suction for compensation or manipulation of prey. Pilot whales were kinematically similar to belugas but relied on a combination of ram and suction that was less discrete than belugas. Belugas were able to purse the anterior lips to occlude lateral gape and form a small, circular anterior aperture that is convergent with feeding behaviors observed in more basal vertebrates. Suction generation in odontocetes is a function of hyolingual displacement and rapid jaw opening, and is likely to be significantly enhanced by lip pursing behaviors. Some degree of subambient pressure was measured in all species, with belugas reaching 126kPa. Functional variations of suction generation during feeding demonstrate a wider diversity of feeding behaviors in odontocetes than previously thought. However, odontocete suction generation is convergent with that of more basal aquatic vertebrates.
SUMMARYOntogenetic studies of vertebrate feeding performance can help address questions relevant to foraging ecology. Feeding morphology and performance can either limit access to food resources or open up new trophic niches in both aquatic and terrestrial systems. Loggerhead sea turtles are long-lived vertebrates with complex life histories that are marked by an ontogenetic shift from an oceanic habitat to a coastal neritic habitat, and a transition from soft oceanic prey to hard, benthic prey. Although considered durophagous and strong biters, bite performance has not been measured in loggerheads, nor has the ontogeny of bite performance been characterized. In the present study, we collected measurements of bite force in loggerhead turtles from hatchlings to adults. When subadults reach the body size at which the ontogenetic shift occurs, their crushing capability is great enough for them to consume numerous species of hard benthic prey of small sizes. As loggerheads mature and bite performance increases, larger and harder benthic prey become accessible. Loggerhead bite performance eventually surpasses the crushing capability of other durophagous carnivores, thereby potentially reducing competition for hard benthic prey. The increasing bite performance and accompanying changes in morphology of the head and jaws are likely an effective mechanism for resource partitioning and decreasing trophic competition. Simultaneous measurements of body and head size and the use of non-linear reduced major axis regression show that bite force increases with significant positive allometry relative to body size (straight carapace length, straight carapace width and mass) and head size (head width, height and length). Simple correlation showed that all recorded morphometrics were good predictors of measured bite performance, but an AICc-based weighted regression showed that body size (straight carapace width followed by straight carapace length and mass, respectively) were more likely predictors of bite force than head size morphometrics (head width and head length).
Prey capture is critical for survival, and differences in correctly positioning and timing a strike (accuracy) are likely related to variation in capture success. However, an ability to quantify accuracy under natural conditions, particularly for fishes, is lacking. We developed a predictive model of suction hydrodynamics and applied it to natural behaviours using three-dimensional kinematics of three centrarchid fishes capturing evasive and non-evasive prey. A spheroid ingested volume of water (IVW) with dimensions predicted by peak gape and ram speed was verified with known hydrodynamics for two species. Differences in capture success occurred primarily with evasive prey (64-96% success). Micropterus salmoides had the greatest ram and gape when capturing evasive prey, resulting in the largest and most elongate IVW. Accuracy predicted capture success, although other factors may also be important. The lower accuracy previously observed in M. salmoides was not replicated, but this is likely due to more natural conditions in our study. Additionally, we discuss the role of modulation and integrated behaviours in shaping the IVW and determining accuracy. With our model, accuracy is a more accessible performance measure for suction-feeding fishes, which can be used to explore macroevolutionary patterns of prey capture evolution.
Organisms are comprised of many interacting parts, and an increased number or specialization of those parts leads to greater complexity and the necessity for increased integration (the ability of those parts to perform together and maintain a functioning organism). Although this idea is widely recognized among biologists, organisms are more tangibly studied when those parts are considered independently. This reductionist approach has successfully advanced our understanding of organisms' performance. However, performance of one system might (or might not) be dependent on performance of another system to achieve a relevant outcome, and the mechanism of this dependence is poorly understood. We synthesize the concepts of complexity and integration and discuss their application in a biomechanical context. Capture of prey by predatory fishes is used as an example to highlight the application of these ideas. We provide a theoretical framework for future hypotheses of integration and predict an "integration space" for fishes that is then populated with data extracted from the literature. Additionally, using the kinematics of prey-capture in two species of sculpin (Scorpaeniformes: Cottidae), we show that species exhibit multivariate integration in distinct ways, and that these differences add additional insight into ecological divergence that would not be apparent by considering systems independently. Finally, we discuss new insights into organismal performance gained through the study of integration as an emergent property of kinematic systems working together during a common task. Integration is rarely the trait of interest, but we show that future work should adopt a more holistic approach to understand why and how animals perform complex behaviors.
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