A power amplification dyad in seahorses.

Throughout evolution, organisms repeatedly developed elastic elements to power explosive body motions, overcoming ubiquitous limits on the power capacity of fast-contracting muscles. Seahorses evolved such a latch-mediated spring-actuated (LaMSA) mechanism; however, it is unclear how this mechanism powers the two complementary functions necessary for feeding: rapidly swinging the head towards the prey, and sucking water into the mouth to entrain it. Here, we combine flow visualization and hydrodynamic modelling to estimate the net power required for accelerating the suction feeding flows in 13 fish species. We show that the mass-specific power of suction feeding in seahorses is approximately three times higher than the maximum recorded from any vertebrate muscle, resulting in suction flows that are approximately eight times faster than similar-sized fishes. Using material testing, we reveal that the rapid contraction of the sternohyoideus tendons can release approximately 72% of the power needed to accelerate the water into the mouth. We conclude that the LaMSA system in seahorses is powered by two elastic elements, the sternohyoideus and epaxial tendons. These elements jointly actuate the coordinated acceleration of the head and the fluid in front of the mouth. These findings extend the known function, capacity and design of LaMSA systems.

Trophic guilds of suction-feeding fishes are distinguished by their characteristic hydrodynamics of swimming and feeding.

Suction-feeding in fishes is a ubiquitous form of prey capture whose outcome depends both on the movements of the predator and the prey, and on the dynamics of the surrounding fluid, which exerts forces on the two organisms. The inherent complexity of suction-feeding has challenged previous efforts to understand how the feeding strikes are modified when species evolve to feed on different prey types. Here, we use the concept of dynamic similarity, commonly applied to understanding the mechanisms of swimming, flying, walking and aquatic feeding. We characterize the hydrodynamic regimes pertaining to (i) the forward movement of the fish (ram), and (ii) the suction flows for feeding strikes of 71 species of acanthomorph fishes. A discriminant function analysis revealed that feeding strikes of zooplanktivores, generalists and piscivores could be distinguished based on their hydrodynamic regimes. Furthermore, a phylogenetic comparative analysis revealed that there are distinctive hydrodynamic adaptive peaks associated with zooplanktivores, generalists and piscivores. The scaling of dynamic similarity across species, body sizes and feeding guilds in fishes indicates that elementary hydrodynamic principles govern the trophic evolution of suction-feeding in fishes.

A new theoretical performance landscape for suction feeding reveals adaptive kinematics in a natural population of reef damselfish.

Understanding how organismal traits determine performance and, ultimately, fitness is a fundamental goal of evolutionary eco-morphology. However, multiple traits can interact in non-linear and context-dependent ways to affect performance, hindering efforts to place natural populations with respect to performance peaks or valleys. Here, we used an established mechanistic model of suction-feeding performance (SIFF) derived from hydrodynamic principles to estimate a theoretical performance landscape for zooplankton prey capture. This performance space can be used to predict prey capture performance for any combination of six morphological and kinematic trait values. We then mapped in situ high-speed video observations of suction feeding in a natural population of a coral reef zooplanktivore, Chromis viridis, onto the performance space to estimate the population's location with respect to the topography of the performance landscape. Although the kinematics of the natural population closely matched regions of high performance in the landscape, the population was not located on a performance peak. Individuals were furthest from performance peaks on the peak gape, ram speed and mouth opening speed trait axes. Moreover, we found that the trait combinations in the observed population were associated with higher performance than expected by chance, suggesting that these combinations are under selection. Our results provide a framework for assessing whether natural populations occupy performance optima.

Elastic energy storage in seahorses leads to a unique suction flow dynamics compared with other actinopterygians.

Suction feeding is a dominant prey-capture strategy across actinopterygians, consisting of a rapid expansion of the mouth cavity that drives a flow of water containing the prey into the mouth. Suction feeding is a power-hungry behavior, involving the actuation of cranial muscles as well as the anterior third of the fish's swimming muscles. Seahorses, which have reduced swimming muscles, evolved a unique mechanism for elastic energy storage that powers their suction flows. This mechanism allows seahorses to achieve head rotation speeds that are 50 times faster than those of fish lacking such a mechanism. However, it is unclear how the dynamics of suction flows in seahorses differ from the conserved pattern observed across other actinopterygians, or how differences in snout length across seahorses affect these flows. Using flow visualization experiments, we show that seahorses generate suction flows that are 8 times faster than those of similar-sized fish, and that the temporal patterns of cranial kinematics and suction flows in seahorses differ from the conserved pattern observed across other actinopterygians. However, the spatial patterns retain the conserved actinopterygian characteristics, where suction flows impact a radially symmetric region of ∼1 gape diameter outside the mouth. Within seahorses, increases in snout length were associated with slower suction flows and faster head rotation speeds, resulting in a trade-off between pivot feeding and suction feeding. Overall, this study shows how the unique cranial kinematics in seahorses are manifested in their suction-feeding performance, and highlights the trade-offs associated with their unique morphology and mechanics.

Coral tentacle elasticity promotes an out-of-phase motion that improves mass transfer.

Corals rely almost exclusively on the ambient flow of water to support their respiration, photosynthesis, prey capture, heat exchange and reproduction. Coral tentacles extend to the flow, interact with it and oscillate under the influence of waves. Such oscillating motions of flexible appendages are considered adaptive for reducing the drag force on flexible animals in wave-swept environments, but their significance under slower flows is unclear. Using in situ and laboratory measurements of the motion of coral tentacles under wave-induced flow, we investigated the dynamics of the tentacle motion and its impact on mass transfer. We found that tentacle velocity preceded the water velocity by approximately one-quarter of a period. This out-of-phase behaviour enhanced mass transfer at the tentacle tip by up to 25% as compared with an in-phase motion. The enhancement was most pronounced under flows slower than 3.2 cm s-1, which are prevalent in many coral-reef environments. We found that the out-of-phase motion results from the tentacles' elasticity, which can presumably be modified by the animal. Our results suggest that the mechanical properties of coral tentacles may represent an adaptive advantage that improves mass transfer under the limiting conditions of slow ambient flows. Because the mechanism we describe operates by enhancing convective processes, it is expected to enhance other fitness-determining transport phenomena such as heat exchange and particle capture.

Work that body: fin and body movements determine herbivore feeding performance within the natural reef environment.

Herbivorous fishes form a keystone component of reef ecosystems, yet the functional mechanisms underlying their feeding performance are poorly understood. In water, gravity is counter-balanced by buoyancy, hence fish are recoiled backwards after every bite they take from the substrate. To overcome this recoil and maintain contact with the algae covered substrate, fish need to generate thrust while feeding. However, the locomotory performance of reef herbivores in the context of feeding has hitherto been ignored. We used a three-dimensional high-speed video system to track mouth and body kinematics during in situ feeding strikes of fishes in the genus Zebrasoma, while synchronously recording the forces exerted on the substrate. These herbivores committed stereotypic and coordinated body and fin movements when feeding off the substrate and these movements determined algal biomass removed. Specifically, the speed of rapidly backing away from the substrate was associated with the magnitude of the pull force and the biomass of algae removed from the substrate per feeding bout. Our new framework for measuring biting performance in situ demonstrates that coordinated movements of the body and fins play a crucial role in herbivore foraging performance and may explain major axes of body and fin shape diversification across reef herbivore guilds.

Rapid adaptive evolution of scale-eating kinematics to a novel ecological niche.

The origins of novel trophic specialization, in which organisms begin to exploit resources for the first time, may be explained by shifts in behavior such as foraging preferences or feeding kinematics. One way to investigate behavioral mechanisms underlying ecological novelty is by comparing prey capture kinematics among species. We investigated the contribution of kinematics to the origins of a novel ecological niche for scale-eating within a microendemic adaptive radiation of pupfishes on San Salvador Island, Bahamas. We compared prey capture kinematics across three species of pupfish while they consumed shrimp and scales in the lab, and found that scale-eating pupfish exhibited peak gape sizes twice as large as in other species, but also attacked prey with a more obtuse angle between their lower jaw and suspensorium. We then investigated how this variation in feeding kinematics could explain scale-biting performance by measuring bite size (surface area removed) from standardized gelatin cubes. We found that a combination of larger peak gape and more obtuse lower jaw and suspensorium angles resulted in approximately 40% more surface area removed per strike, indicating that scale-eaters may reside on a performance optimum for scale biting. To test whether feeding performance could contribute to reproductive isolation between species, we also measured F1 hybrids and found that their kinematics and performance more closely resembled generalists, suggesting that F1 hybrids may have low fitness in the scale-eating niche. Ultimately, our results suggest that the evolution of strike kinematics in this radiation is an adaptation to the novel niche of scale eating.

The hydrodynamic regime drives flow reversals in suction-feeding larval fishes during early ontogeny.

Fish larvae are the smallest self-sustaining vertebrates. As such, they face multiple challenges that stem from their minute size, and from the hydrodynamic regime in which they dwell. This regime, of intermediate Reynolds numbers, was shown to affect the swimming of larval fish and impede their ability to capture prey. Prey capture is impeded because smaller larvae produce weaker suction flows, exerting weaker forces on the prey. Previous observations on feeding larvae also showed prey exiting the mouth after initially entering it (hereafter 'in-and-out'), although the mechanism causing such failures had been unclear. In this study, we used numerical simulations to investigate the hydrodynamic mechanisms responsible for the failure to feed caused by this in-and-out prey movement. Detailed kinematics of the expanding mouth during prey capture by larval Sparus aurata were used to parameterize age-specific numerical models of the flows inside the mouth. These models revealed that for small larvae which expand their mouth slowly, fluid entering the mouth cavity is expelled through the mouth before it is closed, resulting in flow reversal at the orifice. This relative efflux of water through the mouth was >8% of the influx through the mouth for younger ages. However, similar effluxes were found when we simulated slow strikes by larger fish. The simulations can explain the observations of larval fish failing to feed because of the in-and-out movement of the prey. These results further highlight the importance of transporting the prey from the gape deeper into the mouth cavity in determining suction-feeding success.

Pleiotropic jaw morphology links the evolution of mechanical modularity and functional feeding convergence in Lake Malawi cichlids.

Complexity in how mechanistic variation translates into ecological novelty could be critical to organismal diversification. For instance, when multiple distinct morphologies can generate the same mechanical or functional phenotype, this could mitigate trade-offs and/or provide alternative ways to meet the same ecological challenge. To investigate how this type of complexity shapes diversity in a classic adaptive radiation, we tested several evolutionary consequences of the anterior jaw four-bar linkage for Lake Malawi cichlid trophic diversification. Using a novel phylogenetic framework, we demonstrated that different mechanical outputs of the same four jaw elements are evolutionarily associated with both jaw protrusion distance and jaw protrusion angle. However, these two functional aspects of jaw protrusion have evolved independently. Additionally, although four-bar morphology showed little evidence for attraction to optima, there was substantial evidence of adaptive peaks for emergent four-bar linkage mechanics and jaw protrusion abilities among Malawi feeding guilds. Finally, we highlighted a clear case of two cichlid species that have -independently evolved to graze algae in less than 2 Myr and have converged on similar jaw protrusion abilities as well as four-bar linkage mechanics, but have evolved these similarities via non-convergent four-bar morphologies.

The interaction between suction feeding performance and prey escape response determines feeding success in larval fish.

The survival of larval marine fishes during early development depends on their ability to feed before depleting their yolk reserves. Most larval fish capture prey by expanding their mouth, generating a 'suction flow' that draws the prey into it. These larvae dwell in a hydrodynamic environment that impedes their ability to capture even non-evasive prey; however, the marine environment is characterized by an abundance of evasive prey, predominantly copepods. Copepods sense the hydrodynamic disturbance created by approaching predators and perform high-acceleration escape maneuvers. Using a 3D high-speed video system, we characterized the interaction between Sparus aurata larvae and prey from a natural zooplankton assemblage that contained evasive prey, and assessed the factors that determine the outcome of these interactions. At 8-33 days post hatching, larvae preferentially attacked large prey that was moving prior to the initialization of the strike; however, feeding success was lower for larger, more evasive prey. Thus, larvae were challenged in capturing their preferred prey. Larval feeding success increased with increasing Reynolds numbers, but decreased sharply when the prey performed an escape maneuver. The kinematics of successful strikes resulted in a shorter response time but higher hydrodynamic signature available for the prey, suggesting that strike success in our experiments was determined by brevity rather than stealth: executing a fast strike eliminated a potential escape response by the prey. Our observations of prey selectivity reveal that larval performance, rather than preferences, determines their diet during early development.

A novel mechanism of mixing by pulsing corals.

The dynamic pulsation of xeniid corals is one of the most fascinating phenomena observed in coral reefs. We quantify for the first time the flow near the tentacles of these soft corals, the active pulsations of which are thought to enhance their symbionts' photosynthetic rates by up to an order of magnitude. These polyps are approximately 1 cm in diameter and pulse at frequencies between approximately 0.5 and 1 Hz. As a result, the frequency-based Reynolds number calculated using the tentacle length and pulse frequency is on the order of 10 and rapidly decays as with distance from the polyp. This introduces the question of how these corals minimize the reversibility of the flow and bring in new volumes of fluid during each pulse. We estimate the Péclet number of the bulk flow generated by the coral as being on the order of 100-1000 whereas the flow between the bristles of the tentacles is on the order of 10. This illustrates the importance of advective transport in removing oxygen waste. Flow measurements using particle image velocimetry reveal that the individual polyps generate a jet of water with positive vertical velocities that do not go below 0.1 cm s-1 and with average volumetric flow rates of approximately 0.71 cm3 s-1 Our results show that there is nearly continual flow in the radial direction towards the polyp with only approximately 3.3% back flow. 3D numerical simulations uncover a region of slow mixing between the tentacles during expansion. We estimate that the average flow that moves through the bristles of the tentacles is approximately 0.03 cm s-1 The combination of nearly continual flow towards the polyp, slow mixing between the bristles, and the subsequent ejection of this fluid volume into an upward jet ensures the polyp continually samples new water with sufficient time for exchange to occur.

Phylogenomics of a putatively convergent novelty: did hypertrophied lips evolve once or repeatedly in Lake Malawi cichlid fishes?

BACKGROUND: Phylogenies provide critical information about convergence during adaptive radiation. To test whether there have been multiple origins of a distinctive trophic phenotype in one of the most rapidly radiating groups known, we used ultra-conserved elements (UCEs) to examine the evolutionary affinities of Lake Malawi cichlids lineages exhibiting greatly hypertrophied lips. RESULTS: The hypertrophied lip cichlids Cheilochromis euchilus, Eclectochromis ornatus, Placidochromis "Mbenji fatlip", and Placidochromis milomo are all nested within the non-mbuna clade of Malawi cichlids based on both concatenated sequence and single nucleotide polymorphism (SNP) inferred phylogenies. Lichnochromis acuticeps that exhibits slightly hypertrophied lips also appears to have evolutionary affinities to this group. However, Chilotilapia rhoadesii that lacks hypertrophied lips was recovered as nested within the species Cheilochromis euchilus. Species tree reconstructions and analyses of introgression provided largely ambiguous patterns of Malawi cichlid evolution. CONCLUSIONS: Contrary to mitochondrial DNA phylogenies, bifurcating trees based on our 1024 UCE loci supported close affinities of Lake Malawi lineages with hypertrophied lips. However, incomplete lineage sorting in Malawi tends to render these inferences more tenuous. Phylogenomic analyses will continue to provide powerful inferences about whether phenotypic novelties arose once or multiple times during adaptive radiation.

Conserved spatio-temporal patterns of suction-feeding flows across aquatic vertebrates: a comparative flow visualization study.

Suction feeding is a widespread prey capture strategy among aquatic vertebrates. It is almost omnipresent across fishes, and has repeatedly evolved in other aquatic vertebrates. By rapidly expanding the mouth cavity, suction feeders generate a fluid flow outside of their mouth, drawing prey inside. Fish and other suction-feeding organisms display remarkable trophic diversity, echoed in the diversity of their skull and mouth morphologies. Yet, it is unclear how variable suction flows are across species, and whether variation in suction flows supports trophic diversity. Using a high-speed flow visualization technique, we characterized the spatio-temporal patterns in the flow fields produced during feeding in 14 species of aquatic suction feeders. We found that suction-feeding hydrodynamics are highly conserved across species. Suction flows affected only a limited volume of ∼1 gape diameter away from the mouth, and peaked around the timing of maximal mouth opening. The magnitude of flow speed increased with increasing mouth diameter and, to a lesser extent, with decreasing time to peak gape opening. Other morphological, kinematic and behavioral variables played a minor role in shaping suction-feeding dynamics. We conclude that the trophic diversity within fishes, and likely other aquatic vertebrates, is not supported by a diversity of mechanisms that modify the characteristics of suction flow. Rather, we suggest that suction feeding supports such trophic diversity owing to the general lack of strong trade-offs with other mechanisms that contribute to prey capture.

Hydrodynamic regime determines the feeding success of larval fish through the modulation of strike kinematics.

Larval fishes experience extreme mortality rates, with 99% of a cohort perishing within days after starting to actively feed. While recent evidence suggests that hydrodynamic factors contribute to constraining larval feeding during early ontogeny, feeding is a complex process that involves numerous interacting behavioural and biomechanical components. How these components change throughout ontogeny and how they contribute to feeding remain unclear. Using 339 observations of larval feeding attempts, we quantified the effects of morphological and behavioural traits on feeding success of Sparus aurata larvae during early ontogeny. Feeding success was determined using high-speed videography, under both natural and increased water viscosity treatments. Successful strikes were characterized by Reynolds numbers that were an order of magnitude higher than those of failed strikes. The pattern of increasing strike success with increasing age was driven by the ontogeny of traits that facilitate the transition to higher Reynolds numbers. Hence, the physical growth of a larva plays an important role in its transition to a hydrodynamic regime of higher Reynolds numbers, in which suction feeding is more effective.

Hydrodynamic starvation in first-feeding larval fishes.

Larval fishes suffer prodigious mortality rates, eliminating 99% of the brood within a few days after first feeding. Hjort (1914) famously attributed this "critical period" of low survival to the larvae's inability to obtain sufficient food [Hjort (1914) Rapp P-v Réun Cons Int Explor Mer 20:1-228]. However, the cause of this poor feeding success remains to be identified. Here, we show that hydrodynamic constraints on the ubiquitous suction mechanism in first-feeding larvae limit their ability to capture prey, thereby reducing their feeding rates. Dynamic-scaling experiments revealed that larval size is the primary determinant of feeding rate, independent of other ontogenetic effects. We conclude that first-feeding larvae experience "hydrodynamic starvation," in which low Reynolds numbers mechanistically limit their feeding performance even under high prey densities. Our results provide a hydrodynamic perspective on feeding of larval fishes that focuses on the physical properties of the larvae and prey, rather than on prey concentration and the rate of encounters.

Automated detection of feeding strikes by larval fish using continuous high-speed digital video: a novel method to extract quantitative data from fast, sparse kinematic events.

Using videography to extract quantitative data on animal movement and kinematics constitutes a major tool in biomechanics and behavioral ecology. Advanced recording technologies now enable acquisition of long video sequences encompassing sparse and unpredictable events. Although such events may be ecologically important, analysis of sparse data can be extremely time-consuming and potentially biased; data quality is often strongly dependent on the training level of the observer and subject to contamination by observer-dependent biases. These constraints often limit our ability to study animal performance and fitness. Using long videos of foraging fish larvae, we provide a framework for the automated detection of prey acquisition strikes, a behavior that is infrequent yet critical for larval survival. We compared the performance of four video descriptors and their combinations against manually identified feeding events. For our data, the best single descriptor provided a classification accuracy of 77-95% and detection accuracy of 88-98%, depending on fish species and size. Using a combination of descriptors improved the accuracy of classification by ∼2%, but did not improve detection accuracy. Our results indicate that the effort required by an expert to manually label videos can be greatly reduced to examining only the potential feeding detections in order to filter false detections. Thus, using automated descriptors reduces the amount of manual work needed to identify events of interest from weeks to hours, enabling the assembly of an unbiased large dataset of ecologically relevant behaviors.

Suction feeding across fish life stages: flow dynamics from larvae to adults and implications for prey capture.

Suction feeding is thought to be the primary mode of prey capture in most larval fishes. Similar to adult suction feeders, larvae swim towards their prey while rapidly expanding their mouth cavity to generate an inward flow of water that draws the prey into the mouth. Although larvae are known to experience flows with lower Reynolds numbers than adults, it is unclear how the suction-induced flow field changes throughout ontogeny, and how such changes relate to prey capture performance. To address these questions, we determined mouth dimensions and opening speeds in Sparus aurata from first-feeding larvae to adults. We proceeded to develop a computational model of mouth expansion in order to analyze the scaling of suction flows under the observed parameters. Larval fish produced suction flows that were around two orders of magnitude slower than those of adults. Compared with adult fish, in which flow speed decays steeply with distance in front of the mouth, flow speed decayed more gradually in larval fish. This difference indicates that viscous forces in low Reynolds number flows modify the spatial distribution flow speed in front of the mouth. Consequently, simulated predator-prey encounters showed that larval fish could capture inert prey from a greater distance compared with adults. However, if prey attempted to escape then larval fish performed poorly: simulations inferred capture success in only weakly escaping prey immediately in front of the mouth. These ontogenetic changes in Reynolds number, suction-induced flow field and feeding performance could explain a widespread ontogenetic diet shift from passive prey at early life stages to evasive prey as larvae mature.

Mexican blind cavefish use mouth suction to detect obstacles.

Fish commonly use their lateral line system to detect moving bodies such as prey and predators. A remarkable case is the Mexican blind cavefish Astyanax fasciatus, which evolved the ability to detect non-moving obstacles. The swimming body of A. fasciatus generates fluid disturbances, the alteration of which by an obstacle can be sensed by the fish's lateral line system. It is generally accepted that these alterations can provide information on the distance to the obstacle. We observed that A. fasciatus swimming in an unfamiliar environment open and close their mouths at high frequency (0.7-4.5 Hz) in order to generate suction flows. We hypothesized that repeated mouth suction generates a hydrodynamic velocity field, which is altered by an obstacle, inducing pressure gradients in the neuromasts of the lateral line and corresponding strong lateral line stimuli. We observed that the frequency and rate of mouth-opening events varied with the fish's distance to obstacles, a hallmark of pulse-based navigation mechanisms such as echolocation. We formulated a mathematical model of this hitherto unrecognized mechanism of obstacle detection and parameterized it experimentally. This model suggests that suction flows induce lateral line stimuli that are weakly dependent on the fish's speed, and may be an order of magnitude stronger than the correspondent stimuli induced by the fish's gliding body. We illustrate that A. fasciatus can navigate non-visually using a combination of two deeply ancestral and highly conserved mechanisms of ray-finned fishes: the mechanism of sensing water motion by the lateral line system and the mechanism of generating water motion by mouth suction.

Multiple performance peaks for scale-biting in an adaptive radiation of pupfishes.

The physical interactions between organisms and their environment ultimately shape their rate of speciation and adaptive radiation, but the contributions of biomechanics to evolutionary divergence are frequently overlooked. Here we investigated an adaptive radiation of Cyprinodon pupfishes to measure the relationship between feeding kinematics and performance during adaptation to a novel trophic niche, lepidophagy, in which a predator removes only the scales, mucus, and sometimes tissue from their prey using scraping and biting attacks. We used high-speed video to film scale-biting strikes on gelatin cubes by scale-eater, molluscivore, generalist, and hybrid pupfishes and subsequently measured the dimensions of each bite. We then trained the SLEAP machine-learning animal tracking model to measure kinematic landmarks and automatically scored over 100,000 frames from 227 recorded strikes. Scale-eaters exhibited increased peak gape and greater bite length; however, substantial within-individual kinematic variation resulted in poor discrimination of strikes by species or strike type. Nonetheless, a complex performance landscape with two distinct peaks best predicted gel-biting performance, corresponding to a significant nonlinear interaction between peak gape and peak jaw protrusion in which scale-eaters and their hybrids occupied a second performance peak requiring larger peak gape and greater jaw protrusion. A bite performance valley separating scale-eaters from other species may have contributed to their rapid evolution and is consistent with multiple estimates of a multi-peak fitness landscape in the wild. We thus present an efficient deep-learning automated pipeline for kinematic analyses of feeding strikes and a new biomechanical model for understanding the performance and rapid evolution of a rare trophic niche.

Biomechanical trade-offs bias rates of evolution in the feeding apparatus of fishes.

Morphological diversification does not proceed evenly across the organism. Some body parts tend to evolve at higher rates than others, and these rate biases are often attributed to sexual and natural selection or to genetic constraints. We hypothesized that variation in the rates of morphological evolution among body parts could also be related to the performance consequences of the functional systems that make up the body. Specifically, we tested the widely held expectation that the rate of evolution for a trait is negatively correlated with the strength of biomechanical trade-offs to which it is exposed. We quantified the magnitude of trade-offs acting on the morphological components of three feeding-related functional systems in four radiations of teleost fishes. After accounting for differences in the rates of morphological evolution between radiations, we found that traits that contribute more to performance trade-offs tend to evolve more rapidly, contrary to the prediction. While ecological and genetic factors are known to have strong effects on rates of phenotypic evolution, this study highlights the role of the biomechanical architecture of functional systems in biasing the rates and direction of trait evolution.