The perciform group Labroidei includes approximately 2600 species and comprises some of the most diverse and successful lineages of teleost fishes. Composed of four major clades, Cichlidae, Labridae (wrasses, parrotfishes, and weed whitings), Pomacentridae (damselfishes), and Embiotocidae (surfperches); labroids have been an icon for studies of biodiversity, adaptive radiation, and sexual selection. The success and diversification of labroids have been largely attributed to the presence of a major innovation in the pharyngeal jaw apparatus, pharyngognathy, which is hypothesized to increase feeding capacity and versatility. We present results of large-scale phylogenetic analyses and a survey of pharyngeal jaw functional morphology that allow us to examine the evolution of pharyngognathy in a historical context. Phylogenetic analyses were based on a sample of 188 acanthomorph (spiny-rayed fish) species, primarily percomorphs (perch-like fishes), and DNA sequence data collected from 10 nuclear loci that have been previously used to resolve higher level ray-finned fish relationships. Phylogenies inferred from this dataset using maximum likelihood, Bayesian, and species tree analyses indicate polyphyly of the traditional Labroidei and clearly separate Labridae from the remainder of the traditional labroid lineages (Cichlidae, Embiotocidae, and Pomacentridae). These three "chromide" families grouped within a newly discovered clade of 40 families and more than 4800 species (>27% of percomorphs and >16% of all ray-finned fishes), which we name Ovalentaria for its characteristic demersal, adhesive eggs with chorionic filaments. This fantastically diverse clade includes some of the most species-rich lineages of marine and freshwater fishes, including all representatives of the Cichlidae, Embiotocidae, Pomacentridae, Ambassidae, Gobiesocidae, Grammatidae, Mugilidae, Opistognathidae, Pholidichthyidae, Plesiopidae (including Notograptus), Polycentridae, Pseudochromidae, Atherinomorpha, and Blennioidei. Beyond the discovery of Ovalentaria, this study provides a surprising, but well-supported, hypothesis for a convict-blenny (Pholidichthys) sister group to the charismatic cichlids and new insights into the evolution of pharyngognathy. Bayesian stochastic mapping ancestral state reconstructions indicate that pharyngognathy has evolved at least six times in percomorphs, including four separate origins in members of the former Labroidei, one origin in the Centrogenyidae, and one origin within Beloniformes. Our analyses indicate that all pharyngognathous fishes have a mechanically efficient biting mechanism enabled by the muscular sling and a single lower jaw element. However, a major distinction exists between Labridae, which lacks the widespread, generalized percomorph pharyngeal biting mechanism, and all other pharyngognathous clades, which possess this generalized biting mechanism in addition to pharyngognathy. Our results reveal a remarkable history of pharyngognathy: far from a single origin, it appears to have evolved at lea...
Researchers strive to understand what makes species different, and what allows them to survive in the time and space that they do. Many models have been advanced which encompass an array of ecological, evolutionary, mathematical, and logical principles. The goal has been to develop ecological theories that can, among other things, make specific and robust predictions about how and where organisms should live and what organisms should utilize. The role of functional morphology is often an under-appreciated parameter of these models. A more complete understanding of how anatomical features work to allow the organism to accomplish certain tasks has allowed us to revisit some of these ideas with a new perspective. We illustrate our view of this role for functional morphology in ecology by considering the issue of specialization: we attempt to align several definitions of specialization based upon shared ecological and evolutionary principles, and we summarize theoretical predictions regarding why an organism might specialize. Kinematic studies of prey capture in several types of fishes are explored with regard to the potential ecological and evolutionary consequences of specialization, most notably in the area of trade-offs. We suggest that a functional morphological perspective can increase our understanding of the ecological concepts of specialization and it consequences. The kinds of data that functional morphologists collect can help us to quantify organismal performance associated with specialization and the union of functional morphology with ecology can help us to better understand not just how but why organisms interact in the manner that they do.
The exceptionally high speed at which syngnathid fishes are able to rotate their snout towards prey and capture it by suction is potentially caused by a catapult mechanism in which the energy previously stored in deformed elastic elements is suddenly released. According to this hypothesis, tension is built up in tendons of the post-cranial muscles before prey capture is initiated. Next, an abrupt elastic recoil generates high-speed dorsal rotation of the head and snout, rapidly bringing the mouth close to the prey, thus enabling the pipefish to be close enough to engulf the prey by suction. However, no experimental evidence exists for such a mechanism of mechanical power amplification during feeding in these fishes. To test this hypothesis, inverse dynamical modelling based upon kinematic data from high-speed videos of prey capture in bay pipefish Syngnathus leptorhynchus, as well as electromyography of the muscle responsible for head rotation (the epaxial muscle) was performed. The remarkably high instantaneous muscle-mass-specific power requirement calculated for the initial phase of head rotation (up to 5795 W kg K1 ), as well as the early onset times of epaxial muscle activity (often observed more than 300 ms before the first externally discernible prey capture motion), support the elastic power enhancement hypothesis.
The head of ray-finned fishes is structurally complex and is composed of numerous bony, muscular, and ligamentous elements capable of intricate movement. Nearly two centuries of research have been devoted to understanding the function of this cranial musculoskeletal system during prey capture in the dense and viscous aquatic medium. Most fishes generate some amount of inertial suction to capture prey in water. In this overview we trace the history of functional morphological analyses of suction feeding in ray-finned fishes, with a particular focus on the mechanisms by which suction is generated, and present new data using a novel flow imaging technique that enables quantification of the water flow field into the mouth. We begin with a brief overview of studies of cranial anatomy and then summarize progress on understanding function as new information was brought to light by the application of various forms of technology, including high-speed cinematography and video, pressure, impedance, and bone strain measurement. We also provide data from a new technique, digital particle image velocimetry (DPIV) that allows us to quantify patterns of flow into the mouth. We believe that there are three general areas in which future progress needs to occur. First, quantitative three-dimensional studies of buccal and opercular cavity dimensions during prey capture are needed; sonomicrometry and endoscopy are techniques likely to yield these data. Second, a thorough quantitative analysis of the flow field into the mouth during prey capture is necessary to understand the effect of head movement on water in the vicinity of the prey; three-dimensional DPIV analyses will help to provide these data. Third, a more precise understanding of the fitness effects of structural and functional variables in the head coupled with rigorous statistical analyses will allow us to better understand the evolutionary consequences of intra- and interspecific variation in cranial morphology and function.
An important aim of fish ecology is to understand and predict patterns of distribution and abundance in marine communities. Such patterns were examined at four kelp-dominated sites along the northern coast of Chile (19° through 30° S) over 1 year. Fish species richness, diversity and abundance estimates obtained via observational and destructive sampling methods were compared among sites as were habitat and environmental variables that characterised the sites; including sea water temperature below the surface, nutrients, productivity, visibility, density of macroalgae stands, and percentage cover of observed microhabitats (including understorey algae and faunal assemblages). In total, 19 fish species belonging to 14 families were observed from all sites. Species richness and diversity were highest in sites where kelp canopy was composed of two species and where kelp was densest, although only species diversity was significantly different among sites. The sites with high kelp density, in turn, sustained abundant habitat-forming species in the kelp understorey. Principal coordinate analysis indicated that the composition and structure of the fish assemblages varied significantly with depth at all study sites. The depth distribution of fishes was correlated with the arrangement of site-specific biological microhabitats, defined by the algae or invertebrate species that form the microhabitat. Temperature, productivity, and nutrients did not vary consistently across study sites, but did vary within individual sites. We suggest that kelp cover and composition strongly affects the diversity and distribution of fishes at shallow coastal habitats in northern Chile through the availability of microhabitats.
The rate and magnitude of buccal expansion are thought to determine the pattern of water flow and the change in buccal pressure during suction feeding. Feeding events that generate higher flow rates should induce stronger suction pressure and allow predators to draw prey from further away. We tested these expectations by measuring the effects of prey capture kinematics on suction pressure and the effects of the latter on the distance from which prey were drawn-termed suction distance. We simultaneously, but not synchronously, recorded 500-Hz video and buccal pressure from 199 sequences of four largemouth bass, Micropterus salmoides, feeding on goldfish. From the video, we quantified several kinematic variables associated with the head and jaws of the feeding bass that were hypothesized to affect pressure. In a multiple regression, kinematic data accounted for 79.7% of the variation among strikes in minimum pressure. Faster mouth opening and hyoid depression were correlated with lower pressures, a larger area under the pressure curve, and a faster rate of pressure reduction. In contrast, buccal pressure variables explained only 16.5% of the variation in suction distance, and no single pressure variable had a significant relationship with suction distance. Thus, although expected relationships between head kinematics and buccal pressure were confirmed, suction distance was only weakly related to buccal pressure. Three explanations are considered. First, bass may not attempt to maximize the distance from which prey are drawn. Second, the response of prey items to suction-induced flow depends on prey behavior and orientation and is, therefore, subject to considerable variation. Third, previous theoretical work indicates that water velocity decays exponentially with distance from the predator's mouth, indicating that variation among strikes in flow at the mouth opening is compressed away from the mouth. These findings are consistent with other recent data and suggest that suction distance is a poor metric of suction feeding performance.
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