ABSTRACT Unsteady, dynamic flow regimes commonly found in shallow marine ecosystems such as coral reefs pose an energetic challenge for mobile organisms that typically depend on station-holding for fitness-related activities. The majority of experimental studies, however, have measured energetic costs of locomotion at steady speeds, with only a few studies measuring the effects of oscillatory flows. In this study, we used a bidirectional swimming respirometer to create six oscillatory water flow regimes consisting of three frequency and amplitude combinations for both unidirectional and bidirectional oscillatory flows. Using the goldring surgeonfish, Ctenochaetus strigosus , a pectoral-fin (labriform) swimmer, we quantified the net cost of swimming (swimming metabolic rate minus standard metabolic rate) associated with station-holding under these various conditions. We determined that the swimming costs of station-holding in the bidirectional flow regime increased by 2-fold compared with costs based on swimming over the same range of speeds at steady velocities. Furthermore, as we found minimal differences in energetic costs associated with station-holding in the unidirectional, oscillating flow compared with that predicted from steady swimming costs, we conclude that the added acceleration costs are minimal, while the act of turning is an energetically expensive endeavor for this reef fish species.

ABSTRACT Oxygen consumption rates were measured for coral reef fishes during swimming in a bidirectional, oscillatory pattern to simulate station-holding in wave-induced, shallow-water flows. For all species examined, increases in wave intensity, as simulated by increases in frequency and amplitude of oscillation, yielded increased metabolic rates and net costs of swimming (NCOS; swimming metabolic rate minus standard metabolic rate). Comparing species with different swimming modes, the caudal fin swimming Kuhlia spp. (Kuhliidae) and simultaneous pectoral–caudal fin swimming Amphiprion ocellaris (Pomacentridae) turned around to face the direction of swimming most of the time, whereas the median–paired fin (MPF) swimmers, the pectoral fin swimming Ctenochaetus strigosus (Acanthuridae) and dorsal–anal fin swimming Sufflamen bursa (Balistidae), more frequently swam in reverse for one half of the oscillation to avoid turning. Contrary to expectations, the body–caudal fin (BCF) swimming Kuhlia spp. had the lowest overall NCOS in the oscillatory swimming regime compared with the MPF swimmers. However, when examining the effect of increasing frequency of oscillation at similar average velocities, Ku hlia spp. showed a 24% increase in NCOS with a 50% increase in direction changes and accelerations. The two strict MPF swimmers had lower increases on average, suggestive of reduced added costs with increasing frequency of direction changes with this swimming mode. Further studies are needed on the costs of unsteady swimming to determine whether these differences can explain the observed prevalence of fishes using the MPF pectoral fin swimming mode in reef habitats exposed to high, wave-surge-induced water flows.

SUMMARY To determine the energetic costs of rigid-body, median or paired-fin (MPF)swimming versus undulatory, body-caudal fin (BCF) swimming, we measured oxygen consumption as a function of swimming speed in two MPF swimming specialists, Schlegel's parrotfish and Picasso triggerfish. The parrotfish swam exclusively with the pectoral fins at prolonged swimming speeds up to 3.2 total lengths per second ( L s -1 ; 30 min critical swimming speed, U crit ). At higher speeds, gait transferred to a burst-and-coast BCF swimming mode that resulted in rapid fatigue. The triggerfish swam using undulations of the soft dorsal and anal fins up to 1.5 L s -1 , beyond which BCF undulations were recruited intermittently. BCF swimming was used continuously above 3.5 L s -1 , and was accompanied by synchronous undulations of the dorsal and anal fins. The triggerfish were capable of high, prolonged swimming speeds of up to 4.1 L s -1 (30 min U crit ). In both species, the rates of increase in oxygen consumption with swimming speed were higher during BCF swimming than during rigid-body MPF swimming. Our results indicate that, for these species,undulatory swimming is energetically more costly than rigid-body swimming, and therefore support the hypothesis that MPF swimming is more efficient. In addition, use of the BCF gait at higher swimming speed increased the cost of transport in both species beyond that predicted for MPF swimming at the same speeds. This suggests that, unlike for terrestrial locomotion, gait transition in fishes does not occur to reduce energetic costs, but to increase recruitable muscle mass and propulsive surfaces. The appropriate use of the power and exponential functions to model swimming energetics is also discussed.