The scaling of locomotor performance in predator-prey encounters: from fish to killer whales.

During predator-prey encounters, a high locomotor performance in unsteady manoeuvres (i.e. acceleration, turning) is desirable for both predators and prey. While speed increases with size in fish and other aquatic vertebrates in continuous swimming, the speed achieved within a given time, a relevant parameter in predator-prey encounters, is size independent. In addition, most parameters indicating high performance in unsteady swimming decrease with size. Both theoretical considerations and data on acceleration suggest a decrease with body size. Small turning radii and high turning rates are indices of maneuverability in space and in time, respectively. Maneuverability decreases with body length, as minimum turning radii and maximum turning rates increase and decrease with body length, respectively. In addition, the scaling of linear performance in fish locomotion may be modulated by turning behaviour, which is an essential component of the escape response. In angelfish, for example, the speed of large fish is inversely related to their turning angle, i.e. fish escaping at large turning angles show lower speed than fish escaping at small turning angles. The scaling of unsteady locomotor performance makes it difficult for large aquatic vertebrates to capture elusive prey by using whole-body attacks, since the overall maneuverability and acceleration of small prey is likely to be superior to that of large predators. Feeding strategies in vertebrate predators can be related to the predator-prey length ratios. At prey-predator ratios higher than approximately 10(-2), vertebrate predators are particulate feeders, while at smaller ratios, they tend to be filter feeders. At intermediate ratios, large aquatic predators may use a variety of feeding methods that aid, or do not involve, whole body attacks. Among these are bubble curtains used by humpback whales to trap fish schools, and tail-slapping of fish by delphinids. Tail slapping by killer whales is discussed as an example of these strategies. The speed and acceleration achieved by the flukes of killer whales during tail slaps are higher and comparable, respectively, to those that can be expected in their prey, making tail-slapping an effective predator behaviour.     

TURNING MANEUVERS IN STELLER SEA LIONS (EUMATOPIAS JUBATUS)

Steller sea lions are highly maneuverable marine mammals (expressed as minimum turning radius). Video recordings of turns ( n = 195) are analyzed from kinematic measurements for three captive animals. Speed-time plots of 180° turns have a typical "V-shape." The sea lions decelerated during the first half of the turn, reached a minimum speed in the middle of the curved trajectory and reaccelerated by adduction of the pectoral flippers. The initial deceleration was greater than that for passive gliding due to pectoral flipper braking and/or change in body contour from a stiff, straight streamlined form. Centripetal force and thrust were determined from the body acceleration. Most thrust was produced during the power phase of the pectoral flipper stroke cycle. Contrary to previous findings on otariids, little or no thrust was generated during initial abduction of the pectoral flippers and during the final drag-based paddling phase of the stroke cycle. Peak thrust force at the center of gravity occurs halfway through the power phase and the centripetal force is maximal at the beginning of the power stroke. Performance is modulated by changes in the duration and intensity of movements without changing their sequence. Turning radius, maximum velocity, maximum acceleration and turning duration were 0.3 body lengths, 3.5 m/s, 5 m/s², and 1.6 s, respectively. The relative maneuverability based on velocity and length specific minimum turning radius is comparable to other otariids, superior to cetaceans but inferior to many fish.     

Omnidirectional sensory and motor volumes in electric fish

Active sensing organisms, such as bats, dolphins, and weakly electric fish, generate a 3-D space for active sensation by emitting self-generated energy into the environment. For a weakly electric fish, we demonstrate that the electrosensory space for prey detection has an unusual, omnidirectional shape. We compare this sensory volume with the animal's motor volume—the volume swept out by the body over selected time intervals and over the time it takes to come to a stop from typical hunting velocities. We find that the motor volume has a similar omnidirectional shape, which can be attributed to the fish's backward-swimming capabilities and body dynamics. We assessed the electrosensory space for prey detection by analyzing simulated changes in spiking activity of primary electrosensory afferents during empirically measured and synthetic prey capture trials. The animal's motor volume was reconstructed from video recordings of body motion during prey capture behavior. Our results suggest that in weakly electric fish, there is a close connection between the shape of the sensory and motor volumes. We consider three general spatial relationships between 3-D sensory and motor volumes in active and passive-sensing animals, and we examine hypotheses about these relationships in the context of the volumes we quantify for weakly electric fish. We propose that the ratio of the sensory volume to the motor volume provides insight into behavioral control strategies across all animals.     

Dolphin Bait‐Balling Behaviors in Relation to Prey Ball Escape Behaviors

Delphinids frequently coordinate behaviors to contain or capture schooling fishes, but we know little about how these behaviors relate to prey escape behaviors, and corresponding costs and benefits. In this study, we described prey ball escape behaviors and investigated how prey ball behaviors related to dusky dolphin, Lagenorhynchus obscurus, prey herding and capturing behaviors using above‐water and underwater video methods. Prey balls exhibited horizontal and vertical movements, both of which would have increased feeding costs for dolphins compared with feeding on stationary prey at the surface. The only prey ball behavior that we observed to precede escape was ‘funneling’ or the brief formation of a prey ball shape where the height was at least twice the width. Funneling was observed most often for large prey balls, immediately before they descended. When prey balls ascended, there was an insignificant trend for dolphins to do a greater proportion of herding passes that did not include a prey capture (p = 0.094), and dolphins did a larger proportion of prey‐capture attempts on the sides of prey balls (p = 0.017) and a smaller proportion through the edges of prey balls (p = 0.037). When prey balls ascended, dolphins also did a smaller proportion of attempts with a dorsal body orientation (p = 0.037); there were insignificant trends for them to do a smaller proportion with a side orientation (p = 0.100) but a larger proportion with a ventral body orientation (p = 0.081). Thus, funneling is an effective behavior by which large prey balls are able to vertically escape, and dolphins may counter vertical prey escape behaviors by doing herding passes that do not include a prey‐attempt, and changing location or body orientation of prey‐capture behaviors.     

Size-scaling of zooplankton foraging in Arctic charr

Prey capture rate (number of prey s⁻¹) and the mode of feeding of Arctic charr Salvelinus alpinus were studied by performing foraging experiments with two sizes (1·1 and 1·8 mm) of Daphnia longispina prey. Arctic charr were particulate feeders at all densities tested. Adjusted for the effect of prey density, the capture rate showed a hump-shaped relationship with Arctic charr size for both sizes of D. longispina . Estimated attack rates ( a ) also tended to show a hump-shaped relationship with fish size. The estimated size-scaling exponent of the attack rate function, however, was relatively small, implying small changes in attack rate over fish sizes. Simultaneous estimations of a and handling time were used in combination with published data on fish metabolism and dry mass rations of prey to estimate maintenance resource density of prey as a function of Arctic charr mass. Maintenance resource densities increased monotonically with Arctic charr size, and rapidly as optimum fish size relative to attack rate on prey was passed.     

The integration of locomotion and prey capture in vertebrates: Morphology, behavior, and performance

For most vertebrates, locomotion is a fundamental componentof prey capture. Despite this ubiquitous link, few studies havequantified the integration of these complex systems. Severalvariables related to locomotor performance, including maximumspeed, acceleration, deceleration, maneuverability, accuracy,and approach stability, likely influence feeding performancein vertebrates. The relative importance of these measures ofperformance, however, depends on the ecology of the predator.While factors such as morphology and physiology likely definethe limits of these variables, other factors such as motivationof the predator, prey type, and habitat structure can also influenceperformance. Understanding how these variables relate to feedingunder a given suite of ecological conditions is central to understandingpredator–prey interactions, and ultimately how locomotionand feeding have co-evolved. The goals of this article are todiscuss several variables of locomotor performance related toprey capture, present new data on the relationship between locomotorand feeding morphology in fishes, discuss the evolution of preycapture in cichlid fishes, and outline some future directionsfor research. While suction feeding is a primary mechanism ofprey capture in fishes, swimming is vital for accurately positioningthe mouth relative to the prey item. Many fishes decelerateduring prey capture using their body and fins, but the pectoralfins have a dominant role in maintaining approach stability.This suggests that fishes employing high-performance suctionfeeding (relatively small mouth) will have larger pectoral finsto facilitate accurate and stable feeding. I provide new dataon the relationship between pectoral fin morphology and maximumgape in centrarchid fishes. For seven species, pectoral finarea was significantly, and negatively, correlated with maximumgape. This example illustrates that the demands from one complexsystem (feeding) can influence another complex system (locomotion).Future studies that examine the morphological, physiological,and functional evolution of locomotion involved in prey captureby aquatic and terrestrial vertebrates will provide insightinto the origin and consequences of diversity.     

Diet and foraging opportunism of the Guiana dolphin (Sotalia guianensis) in the Abrolhos Bank,Brazil

Diet analysis allows exploring how coastal dolphins interact with the environment and their role in the marine food webs. We studied the diet and feeding ecology of the Guiana dolphin, Sotalia guianensis, through analysis of stomach content from 42 animals stranded on the eastern coast of Brazil. A total of 1,336 semidigested prey items (fish, otoliths, cephalopod beaks, and crustaceans) were identified. Teleost fish comprised the most frequent food item (92% of the total), followed by cephalopods, and crustaceans. Prey belonged to 34 taxa and richness in individual stomachs varied from 1 to 15 prey taxa. Prey were generally small, but showed a significant trend to increase in size with dolphin length. The main prey of Guiana dolphins were demersal, estuarine, and sound-making fish such as catfish and sciaenids. No sex-related differences in diet were found. Diet composition varied seasonally and occurrence of prey items was coherent with breeding or high abundance periods of some fish species and squids. Our study shows the importance of demersal prey from estuarine and soft-sediment habitats to Guiana dolphin in the Abrolhos Bank and reveals that feeding habits are generalist and opportunistic, with diet reflecting the seasonal abundance and availability of prey.     

Forelimb indicators of prey‐size preference in the Felidae

The forelimbs, along with the crania, are an essential part of the prey‐killing apparatus in cats. Linear morphometrics of the forelimbs were used to determine the morphological differences between felids that specialize on large prey, small prey, or mixed prey. We also compared the scaling of felid forelimbs to those of canids to test whether prey capture strategies affect forelimb scaling. Results suggest that large prey specialists have relatively robust forelimbs when compared with smaller prey specialists. This includes relatively more robust humeri and radii, relatively larger distal ends of the humerus, and relatively larger articular areas of the humerus and radius. Large prey specialists also had relatively longer olecranon processes of the ulna and wider proximal paws. These characters are all important for subduing large prey while the cat positions itself for the killing bite. Small prey specialists have relatively longer distal limb elements for swift prey capture, and mixed prey specialists had intermediate values with relatively more robust metacarpals. Arboreal felids also had more robust limbs. They had relatively longer proximal phalanges for better grip while climbing, and a relatively short brachial index (radius to humerus ratio). Additionally, we found that felids and canids differ in forelimb scaling, which emphasizes the dual use of forelimbs for locomotion and prey capture in felids. This morphometric technique worked well to separate prey‐size preference in felids, but did not work as well to separate locomotor groups, as scansorial and terrestrial felids were not clearly distinguished. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Balancing requirements for stability and maneuverability in cetaceans

The morphological designs of animals represent a balance betweenstability for efficient locomotion and instability associatedwith maneuverability. Morphologies that deviate from designsassociated with stability are highly maneuverable. Major featuresaffecting maneuverability are positions of control surfacesand flexibility of the body. Within odontocete cetaceans (i.e.,toothed whales), variation in body design affects stabilityand turning performance. Position of control surfaces (i.e.,flippers, fin, flukes, peduncle) provides a generally stabledesign with respect to an arrow model. Destabilizing forcesgenerated during swimming are balanced by dynamic stabilizationdue to the phase relationships of various body components. Cetaceanswith flexible bodies and mobile flippers are able to turn tightlyat low turning rates, whereas fast-swimming cetaceans with lessflexibility and relatively immobile flippers sacrifice smallturn radii for higher turning rates. In cetaceans, body andcontrol surface mobility and placement appear to be associatedwith prey type and habitat. Flexibility and slow, precise maneuveringare found in cetaceans that inhabit more complex habitats, whereashigh-speed maneuvers are used by cetaceans in the pelagic environment.     

Largemouth bass (Micropterus salmoides) switch feeding modalities in response to sensory deprivation

Animals use a suite of sensory modalities to precisely locate and capture prey. While numerous studies have examined the effects of sensory deprivation on the behaviors leading to prey capture and while it is generally believed that information in the pre-strike period determines the way fish capture prey, this study is the first to examine the contribution of sensory information to jaw kinematics during capture. Largemouth bass were filmed using high-speed videography while capturing live mosquitofish. Bass were examined intact, with visual deprivation under infrared light, and with lateral line deprivation following treatment with cobalt chloride. Deprived of visual cues, this visual ram-feeding predator switches towards suction-based feeding to successfully capture prey. They approach prey slowly but open their mouths more rapidly, which has been shown to result in greater buccal pressure, causing their prey to move a greater distance at a more rapid velocity as they are being drawn into the predators' mouths. Deprived of lateral line cues, bass have higher forward velocities during capture and capture prey earlier in the gape cycle. This study demonstrates that sensory pre-strike information directly affects the capture modality employed by fishes and that fish can modulate between ram and suction not only by adjusting the amount of ram by increasing or decreasing their movements, but also by actively increasing the amount of suction used. These results suggest that the ability to modulate feeding behavior may allow animals to not only exploit a broader breadth of prey items, but also to be capable of doing so in a wider variety of environments.     

HABITAT USE PATTERNS AND RANGES OF THE BOTTLENOSE DOLPHIN IN THE GULF OF CALIFORNIA, MEXICO

Abstract: I studied behavior and range patterns of individual bottlenose dolphins during 1984 in the mid-eastern Gulf of California, Mexico. Dolphin sighting rate was significantly higher in areas close to estuary mouths, 0.306 sightings per hour compared with 0.155 sightings per hour in areas distant from estuary mouths. Dolphins used these estuarine areas to feed; 61% of all behavior observed near estuaries was feeding as compared with 23% elsewhere. Traveling comprised 61% of all behavior observed in areas distant from estuary mouths. Estuaries are sites of large concentrations of nutrients which support great numbers of filter-feeding zooplankton and fish. Bottlenose dolphins may specialize on esmarine prey, or they may feed in estuarine areas simply because of the abundance of potential prey that these systems support. In either case, data on relative numbers, distribution patterns, behavior and diet indicate that this is a general trend in habitat use for many coastal populations of this species in the Pacific and Atlantic.
Ranges of a few individuals spanned a minimum of 65 km of coastline, and animals were not permanent residents of a monitored bay. In contrast, dolphins off the coast of Sarasota, Florida, have been reported to be year-round residents with smaller ranges. This difference in degree of site fidelity may be related to habitat differences. The west coast of Florida is dotted with numerous and large estuarine systems which may host permanent prey populations and support resident groups of dolphins. The Gulf of California coastline contains few estuaries; most are small and perhaps support prey resources which are ephemeral, requiring dolphins to range over larger distances in search of food.     

Optimal foraging as the criteria of prey selection by two centrarchid fishes

The nature of prey selection by two centrarchids (white crappie and bluegill) is presented as a model incorporating optimal foraging strategies. The visual field of the foraging fish as represented by the reactive distance is analysed in detail to estimate the number of prey encounters per search bout. The predicted reactive distances are compared with experimental data. The energetic cost associated with fish foraging behaviour is calculated based on the sequence of events that takes place for each prey consumed. Comparisons of the relative abundance of prey species and size categories in the stomach to the lake environment indicated that both white crappie and bluegill (length < 100 mm) strongly select prey utilising an energy optimization strategy. In most cases, the fish exclusively selected large Daphnia ignoring evasive prey types (Cyclops, Diaptomids) and small cladocera. This selectivity is the result of fish actively avoiding prey with high evasion capabilities even though they appear to be high in energetic content and having translated this into optimal selectivity through capture success rates. The energy consideration and visual system, apart from the forager's ability to capture prey, are the major determinants of prey selectivity for large-sized bluegill and white crappie still at planktivorous stages.     

Maneuvering hydrodynamics of fish and small underwater vehicles

The understanding of fish maneuvering and its application tounderwater rigid bodies are considered. The goal is to gaininsight into stealth. The recent progress made in NUWC is reviewed.Fish morphology suggests that control fins for maneuverabilityhave unique scalar relationships irrespective of their speedtype. Maneuvering experiments are carried out with fish thatare fast yet maneuverable. The gap in maneuverability betweenfish and small underwater vehicles is quantified. The hydrodynamicsof a dorsal fin based brisk maneuvering device and a dual flappingfoil device, as applied to rigid cylindrical bodies, are described.The role of pectoral wings in maneuvering and station keepingnear surface waves is discussed. A pendulum model of dolphinswimming is presented to show that body length and tail flappingfrequency are related. For nearly neutrally buoyant bodies,Froude number and maneuverability are related. Analysis of measurementsindicates that the Strouhal number of dolphins is a constant.The mechanism of discrete and deterministic vortex sheddingfrom oscillating control surfaces has the property of largeamplitude unsteady forcing and an exquisite phase dependence,which makes it inherently amenable to active control for precisionmaneuvering. Theoretical control studies are carried out todemonstrate the feasibility of maneuverability of biologicallyinspired bodies under surface waves. The application of fishhydrodynamics to the silencing of propulsors is considered.Two strategies for the reduction of radiated noise are developed.The effects of a reduction of rotational rate are modeled. Theactive cambering of blades made of digitally programmable artificialmuscles, and their thrust enhancement, are demonstrated. Next,wake momentum filling is carried out by artificial muscles atthe trailing edge of a stator blade of an upstream stator propulsor,and articulating them like a fish tail. A reduction of radiatednoise, called blade tonals, is demonstrated theoretically.     

The swimming endurance of threespine sticklebacks, Gasterosteus aculeatus L., from the Afon Rheidol, Wales

The endurance of threespine sticklebacks, Gasterosteus aculeatus , swimming with pectoral fin locomotion at 20° C in a laboratory flume was measured. Each trial lasted a maximum of 480 min. At a speed of 4 body lengths per sec (L s⁻¹) all fish were still swimming at the end of the trial, but endurance decreased at higher speeds. At speeds of 5 or 6 L s⁻¹ (20–30 cm s⁻¹) a few fish still maintained labriform locomotion for the 480 min. However, at a speed of 7 L s⁻¹ all fish furled their pectoral fins and used body and caudal fin propulsion but fatigued rapidly. During sustained swimming, fish could cover distances of 6 km or more. No significant differences between males and females were found.     

How feeding performance and energy intake change with a small increase in the body size of the three-spined stickleback

Changes in the foraging behaviour due to variation in the body size of the three-spined stickleback Gasterosteus aculeatus were investigated. All sizes of fish had a high probability of attacking prey whenever encountered. The probability of eating the prey increased with the size of the fish, as the larger fish had larger jaws and a greater stomach capacity. Therefore, as fish increased in size there was an increase in the probability of successful prey capture. The level of satiation did not have an effect on the prey handling time, which is contrary to other studies and is probably a result of the large prey sizes. The physical size of the prey meant that the handling times were long regardless of the motivational level of the fish. The larger fish took in more energy and at a faster rate, although the time to reach satiation was similar for all fish sizes. The advantage that large fish appear to have in successfully gaining large prey is negated by their greater metabolic requirement. The changes in feeding performance induced by small increases in body size could have important consequences for intraspecific competition, habitat Use and risk of predation.     

Effect of locomotor approach on feeding kinematics in the green anole (Anolis carolinensis)

Squamates are well-known models for studying to examine locomotor and feeding behaviors in tetrapods, but studies that integrate both behavioral activities remain scarce. Anolis lizards are a classical lineage to study the evolutionary relationships between locomotor behavior and complex structural features of the habitat. Here, we analyzed prey-capture behavior in one representative arboreal predator, Anolis carolinensis, to demonstrate the functional links between locomotor strategies and the kinematics of feeding. A. carolinensis uses two strategies to catch living insects on perches: Head-Up Capture and Jump Capture. In both cases, lizards use lingual prehension to capture the prey and the kinematic patterns of the trophic apparatus are not significantly influenced by the selected strategies. Therefore, to capture one prey type, movements of the trophic structures are highly fixed and A. carolinensis modulates the locomotor pattern to exploit the environment. Predation behavior in A. carolinensis integrates two different behavioral patterns: locomotor plasticity of prey-approach and biomechanical stereotypy of tongue prehension to successfully capture the prey.     

Prey capture efficiency in brown capuchin monkeys (Cebus apella) is influenced by sex and corpus callosum morphology

The diet of capuchin monkeys consists largely of fruits, but these monkeys commonly prey upon insects and other invertebrates as well as vertebrates such as lizards, birds, and fish. Capturing small fast‐moving prey requires the ability to process complex visuospatial information such as motion detection, shape, and pursuit. Here we report the results of an experimental investigation into whether capuchins display sex differences in prey capture efficiency, and whether these differences are associated with the morphology of regions of the corpus callosum (CC) involved in visuospatial ability. We examined the prey capture behavior of seven capuchin subjects (four female, three male) in the laboratory by providing subjects opportunities to fish. Additionally, we obtained structural magnetic resonance images from these subjects to determine if spatial‐ability was related to CC anatomy. Over 30 fishing trials, we recorded the number of prey capture attempts, success rate in capturing fish, and hand techniques used in these attempts. Males were significantly faster and more successful than females at capturing prey. In addition, males had smaller total CC:brain ratios than females. Males displayed a left hand bias, as well as significant unimanual usage, whereas females displayed no significant preference for hand usage. Individual capture times were correlated with total CC:brain ratio. Taken together, our results suggest a relationship between prey capture efficiency, sex, and the degree of brain lateralization. Am. J. Primatol. 72:502–508, 2010. © 2010 Wiley‐Liss, Inc.     

Diet of common dolphins (Delphinus delphis) off the south-east coast of southern Africa: opportunism or specialization?

A total of 297 common dolphin (Delphinus delphis) stomachs was used for these diet analyses. All originated from common dolphins incidentally captured in Natal, where their occurrence is strongly associated with an annual, northward fish migration, the sardine run, along the east coast. Thirty-six fish and four cephalopod prey species were identified. Though 86.9% by weight of the diet was made up of only five prey species, common dolphins appear to feed opportunistically, their diet reflecting local prey abundance and availability. Prey were primarily small, easily-caught, pelagic shoaling species, with the main prey being South African pilchard (Sardinops ocellatus). Although pilchard dominated the diet between 1974 and 1992, there were marked annual and seasonal fluctuations in dietary composition, reflecting changes in prey availability and abundance. There were clear differences in the contribution of particular prey species to the diet of different sex and size groups, with strong evidence for resource partitioning between groups. The influence of the annual 'sardine run', and the fish predators associated with it, on the diet, feeding biology and distribution of common dolphins is discussed.     

Does prey capture induce area-restricted search? A fine-scale study using GPS in a marine predator, the wandering albatross

In a patchy environment, predators are expected to increase turning rate and start an area-restricted search (ARS) when prey have been encountered, but few empirical data exist for large predators. By using GPS loggers with devices measuring prey capture, we studied how a marine predator adjusts foraging movements at various scales in relation to prey capture. Wandering albatrosses use two tactics, sit and wait and foraging in flight, the former tactic being three times less efficient than the latter. During flight foraging, birds caught large isolated prey and used ARS at scales varying from 5 to 90 km, with large-scale ARS being used only by young animals. Birds did not show strong responses to prey capture at a large scale, few ARS events occurred after prey capture, and birds did not have high rates of prey capture in ARS. Only at small scales did birds increase sinuosity after prey captures for a limited time period, and this occurred only after they had caught a large prey item within an ARS zone. When this species searches over a large scale, the most effective search rule was to follow a nearly straight path. ARS may be used to restrict search to a particular environment where prey capture is more predictable and profitable.     

Foraging and feeding ecology of Platanista: an integrative review

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