Phenotypic variation and vulnerability to predation in juvenile bluegill sunfish (<Emphasis Type="Italic">Lepomis macrochirus</Emphasis>)

Bluegill sunfish (Lepomis macrochirus) are known to diversify into two forms specialized for foraging on either limnetic or littoral prey. Because juvenile bluegills seek vegetative cover in the presence of largemouth bass (Micropterus salmoides) predators, natural selection should favor the littoral body design at size ranges most vulnerable to predation. Yet within bluegill populations, both limnetic and littoral forms occur where vegetation and predators are present. While adaptive for foraging in different environments, does habitat-linked phenotypic variation also influence predator evasiveness for juvenile bluegills? We evaluate this question by quantifying susceptibility to predation for two groups of morphologically distinct bluegills; a limnetic form characteristic of bluegills inhabiting open water areas (limnetic bluegill) and a littoral form characteristic of bluegills inhabiting dense vegetation (littoral bluegill). In a series of predation trials, we found that bluegill behaviors differed in open water habitat but not in simulated vegetation. In open water habitat, limnetic bluegills formed more dense shoaling aggregations, maintained a larger distance from the predator, and required longer amounts of time to capture than littoral bluegill. When provided with simulated vegetation, largemouth bass spent longer amounts of time pursuing littoral bluegill and captured significantly fewer littoral bluegills than limnetic fish. Hence, morphological and behavioral variation in bluegills was linked to differential susceptibility to predation in open water and vegetated environments. Combined with previous studies, these findings show that morphological and behavioral adaptations enhance both foraging performance and predator evasiveness in different lake habitats.     

Physiological Plasticity to Water Flow Habitat in the Damselfish,Acanthochromis polyacanthus: Linking Phenotype to Performance

The relationships among animal form, function and performance are complex, and vary across environments. Therefore, it can be difficult to identify morphological and/or physiological traits responsible for enhancing performance in a given habitat. In fishes, differences in swimming performance across water flow gradients are related to morphological variation among and within species. However, physiological traits related to performance have been less well studied. We experimentally reared juvenile damselfish, Acanthochromis polyacanthus, under different water flow regimes to test 1) whether aspects of swimming physiology and morphology show plastic responses to water flow, 2) whether trait divergence correlates with swimming performance and 3) whether flow environment relates to performance differences observed in wild fish. We found that maximum metabolic rate, aerobic scope and blood haematocrit were higher in wave-reared fish compared to fish reared in low water flow. However, pectoral fin shape, which tends to correlate with sustained swimming performance, did not differ between rearing treatments or collection sites. Maximum metabolic rate was the best overall predictor of individual swimming performance; fin shape and fish total length were 3.3 and 3.7 times less likely than maximum metabolic rate to explain differences in critical swimming speed. Performance differences induced in fish reared in different flow environments were less pronounced than in wild fish but similar in direction. Our results suggest that exposure to water motion induces plastic physiological changes which enhance swimming performance in A. polyacanthus. Thus, functional relationships between fish morphology and performance across flow habitats should also consider differences in physiology.     

The Influence of Structural Complexity on Fish–zooplankton Interactions: A Study Using Artificial Submerged Macrophytes

Aquatic macrophytes produce considerable structural variation within the littoral zone and as a result the vegetation provides refuge to prey communities by hindering predator foraging activities. The behavior of planktivorous fish Pseudorasbora parva (Cyprinidae) and their zooplankton prey Daphnia pulex were quantified in a series of laboratory experiments with artificial vegetation at densities of 0, 350, 700, 1400, 2100 and 2800 stemsm^–2. Swimming speeds and foraging rates of the fish were recorded at different prey densities for all stem densities. The foraging efficiency of P. parva decreased significantly with increasing habitat complexity. This decline in feeding efficiency was related to two factors: submerged vegetation impeded swimming behavior and obstructed sight while foraging. This study separated the effects of swimming speed variation and of visual impairment, both due to stems, that led to reduced prey–predator encounters and examined how the reduction of the visual field volume may be predicted using a random encounter model.     

Morphology dependent foraging efficiency in perch: a trade‐off for ecological specialization?

Trade-offs in foraging efficiency leading to divergent natural selection between and within populations exploiting different resources are thought to be a primary cause of trophic polymorphism. In this study we focused on the trade-offs in foraging efficiency and growth in a polymorphic perch population. Specifically, we related habitat-specific growth and diet of perch to perch morphology. In a subsequent laboratory study we experimentally tested the trade-off by testing the efficiency of perch with different morphology feeding on pelagic ( Daphnia sp., Chaoborus sp.) and littoral (mayfly larvae) food resources. The feeding performance was tested in different physical environments to see if we could predict growth patterns in the field based on foraging rate and behavior of perch.
In the field study, we found that the perch from the littoral and the pelagic zones differed in both morphology and diet. Within the littoral zone the deeper-bodied individuals grew faster compared to the more streamlined individuals, whereas the opposite pattern was found in the pelagic zone. In the aquarium experiments, perch from the littoral zone had higher capture rates on the pelagic prey types in vegetation trials and on mayfly larvae in both open water and vegetation trials. The pelagic perch had higher capture rates on the pelagic prey types in open water trials. The littoral perch had lower search velocity than the pelagic perch in open water trials whereas the opposite pattern was found in vegetation trials. The attack velocity of the pelagic perch was also higher than that of the littoral perch independent of vegetation structure. Our results suggest that there is a functional trade-off between performance in alternate habitats and general body form in perch. Such trade-offs may promote divergent natural selection and could be the mechanism that give rise to and upholds the pattern in the field.     

Swimming restricted foraging behavior of two zooplanktivorous fishes <Emphasis Type="Italic">Pseudorasbora parva</Emphasis> and <Emphasis Type="Italic">Rasbora daniconius</Emphasis> (Cyprinidae) in a simulated structured environment

In littoral zones of aquatic systems, submerged macrophytes have marked structural variation that can modify the foraging activity of planktivores. Swimming and feeding behavior of Pseudorasbora parva and Rasbora daniconius (Cyprinidae) on their prey Daphnia pulex and Artemia salina, respectively, was studied in a series of laboratory experiments with varying stem densities. A range of stem densities was tested for each of the two species to compare the effect of simulated macrophytes on prey attack rates and swimming speed, average stem distance (D) was measured in fish body lengths for each of the two fish species. We found that, with reducing average stem distance, the attack rate decreased in the similar trend and this trend was similar for both fish species. However, the species differed in the degree to which swimming activity was hindered at increased stem densities, and this was due to species-specific differences in the distance moved with one tail beat. Therefore, we conclude that the reductions in swimming speed with reduced average stem distance are due to the differences in fish movement per tail beat.     

Swimming dynamics and propulsive efficiency of squids throughout ontogeny

Squids encounter vastly different flow regimes throughout ontogeny as they undergo critical morphological changes to their two locomotive systems: the fins and jet. Squid hatchlings (paralarvae) operate at low and intermediate Reynolds numbers (Re) and typically have rounded bodies, small fins, and relatively large funnel apertures, whereas juveniles and adults operate at higher Re and generally have more streamlined bodies, larger fins, and relatively small funnel apertures. These morphological changes and varying flow conditions affect swimming performance in squids. To determine how swimming dynamics and propulsive efficiency change throughout ontogeny, digital particle image velocimetry (DPIV) and kinematic data were collected from an ontogenetic range of long-finned squid Doryteuthis pealeii and brief squid Lolliguncula brevis swimming in a holding chamber or water tunnel (Re = 20-20 000). Jet and fin wake bulk properties were quantified, and propulsive efficiency was computed based on measurements of impulse and excess kinetic energy in the wakes. Paralarvae relied predominantly on a vertically directed, high frequency, low velocity jet as they bobbed up and down in the water column. Although some spherical vortex rings were observed, most paralarval jets consisted of an elongated vortical region of variable length with no clear pinch-off of a vortex ring from the trailing tail component. Compared with paralarvae, juvenile and adult squid exhibited a more diverse range of swimming strategies, involving greater overall locomotive fin reliance and multiple fin and jet wake modes with better defined vortex rings. Despite greater locomotive flexibility, jet propulsive efficiency of juveniles/adults was significantly lower than that of paralarvae, even when juvenile/adults employed their highest efficiency jet mode involving the production of periodic isolated vortex rings with each jet pulse. When the fins were considered together with the jet for several juvenile/adult swimming sequences, overall propulsive efficiency increased, suggesting that fin contributions are important and should not be overlooked in analyses of the swimming performance of squids. The fins produced significant thrust and consistently had higher propulsive efficiency than did the jet. One particularly important area of future study is the determination of coordinated jet/fin wake modes that have the greatest impact on propulsive efficiency. Although such research would be technically challenging, requiring new, powerful, 3D approaches, it is necessary for a more comprehensive assessment of propulsive efficiency of the squid dual-mode locomotive system.     

Field swimming performance of bluegill sunfish,Lepomis macrochirus: implications for field activity cost estimates and laboratory measures of swimming performance

Mobility is essential to the fitness of many animals, and the costs of locomotion can dominate daily energy budgets. Locomotor costs are determined by the physiological demands of sustaining mechanical performance, yet performance is poorly understood for most animals in the field, particularly aquatic organisms. We have used 3‐D underwater videography to quantify the swimming trajectories and propulsive modes of bluegills sunfish (Lepomis macrochirus, Rafinesque) in the field with high spatial (1–3 mm per pixel) and temporal (60 Hz frame rate) resolution. Although field swimming trajectories were variable and nonlinear in comparison to quasi steady‐state swimming in recirculating flumes, they were much less unsteady than the volitional swimming behaviors that underlie existing predictive models of field swimming cost. Performance analyses suggested that speed and path curvature data could be used to derive reasonable estimates of locomotor cost that fit within measured capacities for sustainable activity. The distinct differences between field swimming behavior and performance measures obtained under steady‐state laboratory conditions suggest that field observations are essential for informing approaches to quantifying locomotor performance in the laboratory.     

Sympatric Divergence and Performance Trade-Offs of Bluegill Ecomorphs

Phenotypic plasticity in response to environmental cues can create distinct morphological types within populations. This variation in form, and potentially function, may be a factor in initiating population divergence and the formation of new species. Here we show the translation of sympatric, habitat-specific morphological divergence into performance differences in energy economy, maneuverability and steady-state locomotion. Littoral and pelagic bluegill sunfish ecomorphs show differences in performance that appear adaptive within their respective habitats: greater maneuverability in the heavily vegetated littoral; greater steady-state swimming speed and economy in the open-water pelagic. This represents a trade-off in unsteady versus steady swimming performance, likely because morphological features associated with maximizing maneuverability are incompatible with enhancing steady-swimming performance. This may constrain the direction of adaptive change, maintaining the divergence created by phenotypic plasticity. The combination of habitat specific sympatric adaptation and constraints imposed by performance trade-offs may be an important factor underlying the high rate of speciation in freshwater fishes from post-glacial lakes.     

Foraging behaviour of planktivorous fish in artificial vegetation: the effects on swimming and feeding

In the littoral zones of lakes, aquatic macrophytes produce considerable structural variation that can provide protection to prey communities by hindering predator foraging activity. The swimming and feeding behaviour of a planktivore, Pseudorasbora parva(Cyprinidae) on its prey (Daphnia pulex) was studied in a series of laboratory experiments with varying densities (0, 350, 700, 1400, 2100 and 2800 stems m^–2) of simulated submerged vegetation. Prey availability was varied from 0.5, 1.0, 2.0, 5.0, 10.0 and 25.0 prey l^–1. As the stem density increased, the predator's swimming speed and the number of prey captured decreased relative to feeding in open water. A good relation existed between the number of successful prey captures and swimming speed with the average stem distance to fish body length ratio (D). An abrupt reduction in feeding and swimming was recorded when D was reduced to values less than one.     

A Flexible Fin with Bio-Inspired Stiffness Profile and Geometry

Biological evidence suggests that fish use mostly anterior muscles for steady swimming while the caudal part of the body is passive and,acting as a carrier of energy,transfers the momentum to the surrounding water.Inspired by those findings we hypothesize that certain swimming patterns can be achieved without copying the distributed actuation mechanism of fish but rather using a single actuator at the anterior part to create the travelling wave.To test the hypothesis a pitching flexible fin made of silicone rubber and silicone foam was designed by copying the stiffness distribution profile and geometry of a rainbow trout.The kinematics of the fin was compared to that of a steadily swimming trout.Fin's propulsive wave length and tail-beat amplitude were determined while it was actuated by a single servo motor.Results showed that the propulsive wave length and tail-beat amplitude of a steadily swimming 50 cm rainbow trout was achieved with our biomimetic fin while stimulated using certain actuation parameters (frequency 2.31 Hz and amplitude 6.6 degrees).The study concluded that fish-like swimming can be achieved by mimicking the stiffness and geometry of a rainbow trout and disregarding the details of the actuation mechanism.     

Evolution of alternative foraging tactics driven by water temperature and physiological constraints in an amphibious snake

Amphibious predatory ectotherms live and forage in two environments (aquatic and terrestrial) that can drastically differ in temperature means and variance across space and time. The locomotor performance of ectotherms is known to be strongly affected by temperature. However, how differences in water temperature may drive the evolution of alternative foraging tactics in amphibious animals remains poorly understood. Fish‐eating Viperine snakes Natrix maura occur from high altitude cold water streams to warm shallow lakes, and employ two main feeding strategies: sentinel foraging (underwater sit‐and‐wait behaviour) and active foraging (fish chasing). Using 272 juvenile snakes we measured: the performance kinetics of diving and swimming in a wide range of water temperatures; basal metabolic levels in relation to body temperature; and the type of foraging mode expressed in water‐temperature‐acclimated snakes. Individual swimming performances increased with testing temperature (10, 15, 20, 25 or 30 °C). Apnoea time followed an opposite trend however, plausibly reflecting the fact that oxygen demands are related to the metabolic rate of ectotherms. That is, snake heart rates increased with body temperature. Snakes acclimated to 10 °C water mostly displayed sentinel foraging. By contrast, 20 °C and 30 °C water‐acclimated snakes were extremely active fish chasers. Individual apnoea times at the various testing temperatures were all correlated; as were individual swimming speeds. There was however no clear relationship between an individual's ability to hold its breath and its ability to swim, suggesting that both performance traits may be the target of different selective pressures. Fast swimming speed and long breath holding abilities are likely key determinants of both foraging success and predatory evasion, although in a context dependent manner. Active swimming foraging is likely to be advantageous in warm water (> 20 °C), while sentinel foraging appears better suited to cold water (< 14 °C). The physiological aspects of foraging tactics of amphibious snakes combined with field and laboratory observations support the idea that physiological and environmental constraints may generate shifts in habitat use and associated foraging tactics in amphibious ectotherms. Avenues for further research are discussed. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 411–422.     

Intraspecific and interspecific trait variability in tadpole meta‐communities from the Brazilian Atlantic rainforest

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The Relationship Among Swimming Performance, Courtship Behavior, and Carotenoid Pigmentation of Guppies in Four Rivers of Trinidad

In the laboratory, courtship behavior and carotenoid pigmentation of male guppies are condition-dependent traits, since their expression is affected by physical vigor and environmental factors such as water velocity and diet. Whether these relationships exist in guppies under field conditions is yet to be determined. We compared the swimming performance, courtship behavior, and carotenoid pigmentation of guppies from headwater and downstream localities in four rivers of Trinidad. Swimming performance and courtship behavior of males differed among rivers and between headwater and downstream sites. Guppies from headwater sites swam significantly faster and had higher display rates than those from downstream sites. Mean swimming performance across sites was positively correlated with mean water velocity, but was correlated with the number of orange color spots (carotenoid pigment) in only one river. These results indicate that the courtship behavior of Trinidadian guppies is condition-dependent because the amount of display behavior is positively correlated to swimming performance, a measure of physical endurance. The proximal cause for this condition-dependence may be predator induced variation in microhabitat use by guppies in headwater and downstream locations.     

Wave energy and swimming performance shape coral reef fish assemblages

Physical factors often have an overriding influence on the distribution patterns of organisms, and can ultimately shape the long-term structure of communities. Although distribution patterns in sessile marine organisms have frequently been attributed to functional characteristics interacting with wave-induced water motion, similar evidence for mobile organisms is lacking. Links between fin morphology and swimming performance were examined in three diverse coral reef fish families from two major evolutionary lineages. Among-habitat variation in morphology and performance was directly compared with quantitative values of wave-induced water motion from seven coral reef habitats of different depth and wave exposure on the Great Barrier Reef. Fin morphology was strongly correlated with both field and experimental swimming speeds in all three families. The range of observed swimming speeds coincided closely with the magnitude of water velocities commonly found on coral reefs. Distribution patterns in all three families displayed highly congruent relationships between fin morphology and wave-induced water motion. Our findings indicate a general functional relationship between fin morphology and swimming performance in labriform-swimming fishes, and provide quantitative evidence that wave energy may directly influence the assemblage structure of coral reef fishes through interactions with morphology and swimming performance.     

Analyzing nested variation in the body form of Lepomid sunfishes

Phylogenetic hierarchies are often composed of younger diverging lineages nested within older diverging lineages. Comparing phenotypic variation among several hierarchical levels can be used to test hypotheses about selection, phenotypic evolution and speciation. Such hierarchical comparisons have only been performed in threespine stickleback, and so here we use a hierarchical pattern of divergences between near-shore littoral and off-shore pelagic habitats to test for selection on the evolution of body form in Lepomis sunfish in lakes. We compare variation in external body form between fish from littoral and pelagic habitats at three levels: among ecomorphs within individual lake populations (intrapopulation), among populations of the same species in different lakes (interpopulation), and between bluegill and pumpkinseed sunfish species (interspecifically). Using geometric morphometric methods, we first demonstrate that interpopulation variation in mean body form of pumpkinseed sunfish varies with the presence of pelagic habitat. We then incorporate these results with existing data in order to test the similarity of phenotypic divergence between littoral and pelagic habitats at different hierarchical levels. Parallel relationships between certain body form traits (head length, caudal length and pectoral length) and habitat occur at all three levels suggesting that selection persistently acts at all levels to diversify these traits and so may contribute to species formation. For other traits (caudal depth and pectoral altitude), divergence between habitats is inconsistent at different hierarchical levels. Thus, nested biological variation in Lepomid body form reflects a history of deterministic selection and historical contingency, and also identifies traits that likely have likely influenced fitness and so serve important functions.     

Allometric relationships in morphological traits associated with foraging,swimming ability,and predator defense reveal adaptations toward brackish and freshwater environments in the threespine stickleback

Freshwater colonization by threespine stickleback has led to divergence in morphology between ancestral marine and derived freshwater populations, making them ideal for studying natural selection on phenotypes. In an open brackish–freshwater system, we previously discovered two genetically distinct stickleback populations that also differ in geometric shape: one mainly found in the brackish water lagoon and one throughout the freshwater system. As shape and size are not perfectly correlated, the aim of this study was to identify the morphological trait(s) that separated the populations in geometric shape. We measured 23 phenotypes likely to be important for foraging, swimming capacity, and defense against predation. The lateral plate morphs in freshwater displayed few significant changes in trait sizes, but the low plated expressed feeding traits more associated with benthic habitats. When comparing the completely plated genetically assigned populations, the freshwater, the hybrids, the migrants and the lagoon fish, many of the linear traits had different slopes and intercepts in trait‐size regressions, precluding our ability to directly compare all traits simultaneously, which most likely results from low variation in body length for the lagoon and migrant population. We found the lagoon stickleback population to be more specialized toward the littoral zone, displaying benthic traits such as large, deep bodies with smaller eyes compared to the freshwater completely plated morph. Further, the lagoon and migrant fish had an overall higher body coverage of lateral plates compared to freshwater fish, and the dorsal and pelvic spines were longer. Evolutionary constraints due to allometric scaling relationships could explain the observed, overall restricted, differences in morphology between the sticklebacks in this study, as most traits have diversified in common allometric trajectories. The observed differences in foraging and antipredation traits between the fish with a lagoon and freshwater genetic signature are likely a result of genetic or plastic adaptations toward brackish and freshwater environments.     

Daphnid morphology deters fish predators

Spine and helmet production in zooplankton are thought to provide protection from invertebrate rather than vertebrate predators. We examined selectivity for Daphnia lumholtzi, a species that exhibits extreme cyclomorphosis with a large helmet and long tail spine (total length can exceed 5 mm), by juvenile bluegill (15–80 mm) in the laboratory and field. Bluegill consumed more D. pulex than D. lumholtzi when the species were presented alone. When the daphnids were offered together in equal numbers, bluegill selected against D. lumholtzi. Bluegill foraging behavior helped explain the observed nonrandom feeding. Bluegill capture efficiency foraging on D. pulex was high (85–100%) and handling times were low (usually too short to detect), whereas efficiencies were lower (40–96%) and handling times were longer (1–3 s) when foraging on D. lumholtzi, particularly for fish <50 mm. As they gained experience, bluegill <50 mm that oriented towards D. lumholtzi rejected them more often than striking. In addition, more D. lumholtzi were rejected and expelled than were D. pulex. From these experiments, we conclude that larger bluegill (>50 mm) are able to forage more successfully on D. lumholtzi than smaller fish. Selectivity by bluegill collected from a reservoir infested with D. lumholtzi verified our laboratory conclusions. Smaller bluegill selected against D. lumholtzi, whereas it was a preferred diet item for bluegill >50 mm. These results show that the morphology of D. lumholtzi interferes with predation by small planktivorous fish, posing foraging constraints for these fish more similar to those of piscivores, where handling time is important, than to those of planktivores, where prey density is of primary importance. Received: 13 August 1998 / Accepted 21 August 1998     

Reductions in prolonged swimming capacity following freshwater colonization in multiple threespine stickleback populations

We compared ancestral anadromous-marine and nonmigratory, stream-resident threespine stickleback (Gasterosteus aculeatus) populations to examine the outcome of relaxed selection on prolonged swimming performance. We reared marine and stream-resident fish from two locations in a common environment and found that both stream-resident populations had lower critical swimming speeds (U(crits) ) than marine populations. F1 hybrids from the two locations displayed significant differences in dominance, suggesting that the genetic basis for variation in U(crit) differs between locations. To determine which traits evolved in conjunction with, and may underlie, differences in performance capacity we measured a suite of traits known to affect prolonged swimming performance in fish. Although some candidate traits did not evolve (standard metabolic rate and two body shape traits), multiple morphological (pectoral fin size, shape, and four body shape measures) and physiological (maximum metabolic rate; MMR) traits evolved in the predicted direction in both stream-resident populations. However, data from F1 hybrids suggested that only one of these traits (MMR) had dominance effects similar to those of U(crit) in both locations. Overall, our data suggest that reductions in prolonged swimming performance were selected for in nonmigratory populations of threespine stickleback, and that decreases in MMR may mediate these reductions in performance.     

Intraspecific phenotypic variation among alewife populations drives parallel phenotypic shifts in bluegill

Evolutionary diversification within consumer species may generate selection on local ecological communities, affecting prey community structure. However, the extent to which this niche construction can propagate across food webs and shape trait variation in competing species is unknown. Here, we tested whether niche construction by different life-history variants of the planktivorous fish alewife (Alosa pseudoharengus) can drive phenotypic divergence and resource use in the competing species bluegill (Lepomis macrochirus). Using a combination of common garden experiments and a comparative field study, we found that bluegill from landlocked alewife lakes grew relatively better when fed small than large zooplankton, had gill rakers better adapted for feeding on small-bodied prey and selected smaller zooplankton compared with bluegill from lakes with anadromous or no alewife. Observed shifts in bluegill foraging traits in lakes with landlocked alewife parallel those in alewife, suggesting interspecific competition leading to parallel phenotypic changes rather than to divergence (which is commonly predicted). Our findings suggest that species may be locally adapted to prey communities structured by different life-history variants of a competing dominant species.     

Function of the dorsal fin in bluegill sunfish: Motor patterns during four distinct locomotor behaviors

The median fins of fishes are key features of locomotor morphology which function as complex control surfaces during a variety of behaviors. However, very few studies have experimentally assessed median fin function, as most workers focus on axial structures. In particular, the dorsal fin of many teleost fishes possesses both spiny anterior and soft posterior portions which may function separately during locomotion. We analyzed the function of the soft region of the dorsal fin and of the dorsal inclinator (Di) muscles which are the primary muscles responsible for lateral flexion. We used electromyography to measure in vivo Di activity, as well as activity of the red myomeric muscles located at a similar longitudinal position. We quantified motor patterns during four locomotor behaviors: braking and three propulsive behaviors (steady swimming, kick and glide swimming, and C-starts). During the three propulsive swimming behaviors, the timing of Di activity was more similar to that of ipsilateral red myomeric muscle rather than to contralateral myomeric activity, whereas during braking the timing of activity of the Di muscles was similar to that of the contralateral myomeric musculature. During the three propulsive behaviors, when the Di muscles had activity, it was consistent with the function of stiffening the soft dorsal fin to oppose its tendency to bend as a result of the body being swept laterally through the water. In contrast, activity of the Di muscles during braking was consistent with the function of actively flexing the soft dorsal fin towards the side of the fish that had Di activity. Activity of the Di muscles during steady speed swimming was generally sufficient to resist lateral bending of the soft dorsal fin, whereas during high speed kick and glide swimming and C-starts, Di activity was not sufficient to resist the bending caused by resistive forces imposed by the water. Cumulative data from all four behaviors suggest that the Di muscles can be activated independently relative to the myomeric musculature rather than having a single phase relationship with the myomeric muscle common to all of the observed behaviors. © 1996 Wiley-Liss, Inc.