Article

Backing up from negative stimuli: A back-thrust mechanism during escape-like response in wild sevengill sharks (Notorynchus cepedianus)

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Abstract

Broadnose sevengill sharks (Notorynchus cepedianus) show great interest for bait and display a repertoire of movements while engaging with it. A novel back-thrust mechanism is described in wild sevengill sharks by which individuals back up from a negative stimulus while interacting with baited video stations. This mechanism initiates upon head contact with the device that functions as a negative stimulus eliciting a startle escape-like response. By heavily flipping pectoral fins and curving the body, sharks increase hydrodynamic resistance, backing up from the negative stimulus. Once backed up, sharks performed the common C-shaped double-bend escape maneuver described for sharks. Sharks also used the same back-thrust mechanism as a repositioning maneuver, but not as part of a startle response. The quantification of the turning rate indicated context-dependent variation in velocity and confirmed that the majority of withdrawals corresponded to slow escape-like motions. In general, an elongated body and individual flipping control of pectoral fins allowed for great maneuverability and lateral flexure. Sharks exhibited great tolerance to one another during double and triple encounters. The implications for grouping and social hunting of the species are briefly discussed based on past evidence and the movement behavior, gregarious interactions, and body markings observed in the present study. This work highlights the importance of studies in the natural environment and the use of complementary approaches to investigate the broader range of locomotor aspects of different shark species.

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While most actinopterygian fishes perform C-start or S-start behaviors as their primary startle responses, many elongate species instead use a withdrawal movement. Studies of withdrawal have focused on the response to head-directed or nonspecific stimuli. During withdrawal, the animal moves its head back from the stimulus, often resulting in several tight bends in the body. In contrast to C-start or S-start behaviors, withdrawal to a head stimulus generally does not involve a subsequent propulsive stage of movement. We examined intraspecific diversity in withdrawal behavior and muscle activity patterns of the rope fish, Erpetoichthys calabaricus, in response to stimulation of the head and the tail. In addition, we describe the anatomy of the Mauthner cells and their axon caps, structures that are generally absent in species with a withdrawal startle. We recorded high-speed video (250 Hz) and electromyograms (EMGs) from 12 electrodes in the axial muscle during the behavioral response. We used Bodian silver staining techniques to visualize Mauthner cell and axon cap morphology. We found that E. calabaricus responds with a withdrawal to both head and tail stimulation. Tail stimulation elicits a stronger kinematic and muscle activity response than head stimulation. While withdrawal movement generally constitutes the entire response to head stimuli, withdrawal was followed by propulsive movements when the tail was stimulated, suggesting that withdrawal can both act alone and serve as the first stage of a propulsive startle. Unexpectedly, bilaterality of muscle activity was variable for responses to both head and tail stimuli. In addition, we were surprised to find that E. calabaricus has a distinct axon cap associated with its Mauthner cell. These data suggest that the withdrawal response is a more diverse functional system than has previously been believed.
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Sharks are among the oldest vertebrate lineages in which their success has been attributed to their diversity in body shape and locomotor design. In this study, we investigated the diversity of body forms in extant sharks using landmark-based geometric morphometric analyses on nearly all the known (ca. 470) extant sharks. We ran three different analyses: the ‘full body,' ‘precaudal body,’ and ‘caudal fin' analyses. Our study suggests that there are two basic body forms in sharks, a ‘shallow-bodied’ form (Group A) and ‘deep-bodied’ form (Group B), where all sharks essentially have one basic caudal fin design of a heterocercal tail despite some specializations. We found that swimming modes in sharks are highly correlated with body forms where Group A sharks are predominantly anguilliform swimmers and Group B sharks are represented by carangiform and thunniform swimmers. The majority of Group A sharks are found to be benthic whereas pelagic forms are relatively common among Group B sharks. Each of the two superorders of sharks, Squalomorphii and Galeomorphii, must have gone through complex evolutionary history where each superorder contains both Group A sharks and Group B sharks, possibly involving parallel evolution from one group to the other or at least one episode of evolutionary reversal.
Article
The study of fish escape responses has provided important insights into the accelerative motions and fast response times of these animals. In addition, the accessibility of the underlying neural circuits has made the escape response a fundamental model in neurobiology. Fish escape responses were originally viewed as highly stereotypic all-or-none behaviours. However, research on a wide variety of species has shown considerable taxon-specific and context-dependent variability in the kinematics and neural control of escape. In addition, escape-like motions have been reported: these resemble escape responses kinematically, but occur in situations that do not involve a response to a threatening stimulus. This Review focuses on the diversity of escape responses in fish by discussing recent work on: (1) the types of escape responses as defined by kinematic analysis (these include C- and S-starts, and single- versus double-bend responses); (2) the diversity of neuromuscular control; (3) the variability of escape responses in terms of behaviour and kinematics within the context of predator-prey interactions; and (4) the main escape-like motions observed in various species. Here, we aim to integrate recent knowledge on escape responses and highlight rich areas for research. Rapidly developing approaches for studying the kinematics of swimming motion both in the lab and within the natural environment provide new avenues for research on these critical and common behaviours.
Article
The habitat use, seasonality and demography of the broadnose sevengill shark Notorynchus cepedianus were investigated in central Patagonia, where a data gap exists for the species. Catch and effort and video‐derived indices indicated high relative abundance of sharks during warm months. Video stations revealed differences in the spatial use by sharks, being more frequently observed in the inner section of the bay. Complementary tagging efforts evidenced both a seasonal residence pattern and site fidelity between consecutive warm seasons. Juveniles outnumbered adults, which suggests that they may use the study area as a secondary nursing ground. Evidence from spontaneous regurgitation further suggests that prey abundance may be driving the seasonal occurrence of sharks in the region. This study allows for a more comprehensive understanding of the population structure and dynamics of sevengill sharks in the Southwest Atlantic. Abstract in Spanish is available with online material.
Article
While post-release mortality estimates have been conducted on a range of shark species, the short-term sub-lethal effects of capture, handling and release are poorly known and have been mostly investigated in controlled conditions. In addition, the widely accepted notion that immediate post-release active swimming is an indicator of shark condition has never been tested. This study assessed the effects of deck exposure by analysing post-release behaviour of two species of shark, the draughtboard (Cephaloscyllium laticeps) and the piked spurdog (Squalus megalops) in-situ using a remote-operated-vehicle and in a replicated experiment in controlled conditions. In total, 67 sharks were caught with demersal fish trawls and commercial longlines and subjected to different deck exposures and release environments. Tailbeat rates of deck-exposed sharks were significantly lower than the control sharks, but this effect differed between in-situ and experimental environments. Results indicate that capture has species-specific effects, that post-release effects may last longer than 5 min, and that controlled experiments may not be reliable indicators of post-release effects. Immediate post-release swimming was not a good predictor of post-release behaviour, suggesting capture and release fisheries may have significant sub-lethal effects on some species of shark, and that limiting capture or handling time may reduce post-release effects.
Article
Gaining insight on how species relate to the environment and other members of the trophic community is critical for their management. This study investigated habitat use, seasonal patterns of abundance and population structure of the broadnose sevengill shark Notorynchus cepedianus in the marine temperate environment of a northern Patagonian bay (Argentina). We expected N. cepedianus to exhibit strong seasonality in the bay driven by the seasonal occurrence of important marine mammal prey. However, an alternative or complementary hypothesis could be related to the possibility that N. cepedianus uses the area seasonally as mating grounds. To explore these hypotheses a suite of baited remote underwater video stations and catch and effort data indices were used to estimate the seasonal relative abundance of the species inside the bay. Reproductive hormone levels were employed to determine reproductive status and maturity stages of the population. Results indicated that N. cepedianus uses the bay year-round, showing a peak of abundance during spring when the species aggregates, likely for feeding and mating purposes. Male and female sizes at maturity occurred at 170 and 190 cm of total length, respectively. Contrary to other studies in the Southwest Atlantic, population composition remained similar throughout the seasons, with adults dominating over juveniles and adult females prevailing in the population at all times. Considering the historical abundance decline recently reported for the species, protection and enforcement within protected areas across the Southwest Atlantic must remain of high priority in government initiatives.
Article
Baited remote underwater video systems (BRUVS) are increasingly used to study fish communities, biomass, and animal behaviour. Due to the abundance of BRUVS data, there are many analysis methods. The most commonly used method for analysis of BRUV data is MaxN which refers to the maximum number of individuals observed of a species in a single frame of a video. Here, we present a novel method for BRUVS analysis that involves identifying and counting distinct individuals (MaxIND) to quantify the accuracy of MaxN. Individual oriental bluespotted maskray (Neotrygon orientalis) and the bluespotted fantail ray (Taeniura lymma) were identified on BRUVS by spot patterns, tail characteristics, and sex at three sites in Malaysian Borneo. We demonstrated that MaxIND gave abundances that were 2.4 and 1.1 times higher than MaxN for N. orientalis and T. lymma, respectively. These differences between methods were consistent for each species between sites regardless of the presence of marine reserves. However, differences in abundance estimates from MaxN to MaxIND were apparent between species, indicating that correction factors need to be developed on a species basis to better estimate true abundance. While identifying individuals is time consuming, it provides improved accuracy and information about populations. We therefore recommend the use of MaxIND when rare and endangered species are present, in high density populations, and for behavioural analyses.
Article
The locomotor performance of dogfish during escape responses was observed by means of high-speed video. Dogfish show C-type escape responses that are comparable with those shown previously in teleosts. Dogfish show high variability of turning rates of the anterior part of the body (head to centre of mass), i.e. with peak values from 434 to 1023 deg. s - 1 . We suggest that this variability may be due to the presence of two types of escape manoeuvres, i.e. responses with high and low turning rates, as previously found in a teleost species. Fast responses (i.e. with high maximum turning rates, ranging between 766 and 1023 deg. s - 1 ) showed significantly higher locomotor performance than slow responses (i.e. with low maximum turning rates, ranging between 434 and 593 deg. s - 1 ) in terms of distance covered, speed and acceleration, although no differences were found in the turning radius of the centre of mass during the escape manoeuvres. The existence of two types of escape responses would have implications in terms of both neural control and muscular activation patterns. When compared with literature data for the locomotor performance of bony fishes, dogfish showed relatively low speed and acceleration, comparable turning rates and a turning radius that is in the low part of the range when compared with teleosts, indicating relatively high manoeuvrability. The locomotor performance observed in dogfish is consistent with their morphological characteristics: (1) low locomotor performance associated with low thrust developed by their relatively small posterior depth of section and (2) relatively high manoeuvrability associated with their high flexibility.
Article
To investigate the biochemical effects of capture and restraint on sharks, 17 serum constituents were measured in three species (bonnethead shark Sphyrna tiburo, blacktip shark Carcharhinus limbatus, and bull shark C. leucas) after gill-net capture. The relative degree of capture effects on each animal was judged using an index of behavioral response devised for use in tag−recapture studies. Serum from each shark was assayed for glucose, creatinine, uric acid, sodium, chloride, potassium, inorganic phosphate, total and ionized calcium, total protein, albumin, globulin, alkaline phosphatase, lactate, lactate dehydrogenase, aspartate aminotransferase, and total iron. In addition, hematocrit was measured from whole blood for each shark. When correlated with the relative degree of capture effects, there were significant intraspecific changes in the concentration of potassium, lactate, inorganic phosphate, uric acid, alkaline phosphatase, aspartate aminotransferase, total and ionized calcium, and glucose. Significant interspecific differences in the concentration of sodium, chloride, potassium, total protein, albumin, globulin, lactate dehydrogenase, aspartate aminotransferase, ionized calcium, alkaline phosphatase, and glucose in minimally stressed animals also were observed. The study suggests that the deleterious effects of gill-net capture and restraint probably involve respiratory and metabolic acidosis and hypoglycemia as well as cellular damage. Species-specific and individual differences in the mortality of sharks caught in gill nets are likely related to an animal's respiratory physiology and degree of struggling upon capture as well as to the extent of net entanglement around the gill area.
Article
SUMMARY Fast-start kinematics and performance were determined for Etheostoma caeruleum, Cottus cognatus, Notropis cornutus, Lepomis macrochirus, Perca flavescens, Salmo gairdneri and a hybrid Esox sp. at an acclimation and test temperature of 15 °C. Normal three-stage kinematic patterns were observed for all species. Fast-start movements were similar in all species, except Lepomis, which had slightly higher amplitudes than expected for its length. The duration of kinematic stages was a major variable among the seven species but was a linear function of length. Acceleration rates were not functions of size. Maximum acceleration rates ranged from 22-7 to 39-5 m. s~s with mean rates from 6-i to 12*3 m.s~2, averaged to the completion of kinematic stage 2. Maximum velocity and distance covered in each fast-start stage varied among species but were related to length. Fast-start performance depended primarily on compromise between muscle mass as a percentage of body mass, and lateral body and fin profile. Optimal profiles provide large depth distant from the centre of mass to maximize thrust, and anterior depth enhancement to minimize recoil. The body form of Lepomis is considered optimal for multiple swimming modes.
Article
Fast-starts are brief, sudden accelerations used by fish during predator-prey encounters. The kinematics and performance of fish during fast-start manoeuvres have received a lot of attention since they may determine the outcome of predator-prey interactions in terms of feeding success or survival. We will discuss recent progress on (1) the kinematics of escape responses and feeding strikes, (2) the fast-start performance of species with different body morphologies and from different habitats, and (3) the functional significance of fast-start kinematics and performance within the context of predator-prey interactions.
Article
Combining movement behaviour with other ecological information of predators and their prey is essential for an adequate understanding of ecosystem dynamics. The movement patterns of broadnose sevengill sharks, Notorynchus cepedianus , were monitored with acoustic and satellite technology in coastal areas of southeast Tasmania, Australia. Individuals were tagged in two habitats (Norfolk Bay and the Derwent Estuary) for which we had ecological information such as diet, population structure and abundance. Notorynchus cepedianus showed seasonal site fidelity in the use of the coastal habitats. The general pattern was for sharks to exit coastal areas over winter and females to return the following spring and males in summer. Their movement into these coastal areas coincided with high seasonal abundance of their known prey species during summer, suggesting feeding site fidelity. Individuals tagged in two coastal areas showed low spatial and dietary overlap, suggesting localized site fidelity and fine spatial scale resource partitioning. This has rarely been reported for large mobile predators. Both satellite and acoustic methods showed that males make northerly migrations during winter to distances of at least 1000 km. The combined use of tracking, diet and abundance information demonstrated that N. cepedianus are likely to exert significant predation pressure on prey inhabiting these areas during summer. Overall, this study highlights the benefit of complementing movement data with other ecological information to understand the habitat use of large mobile predators and their potential influences on ecosystem structure and function.
Article
A bstract Daily counts of Southern sea lions ( Otaria byronia ) made at Punta Norte, Argentina, during four consecutive breeding seasons (1980–1984) yielded similar results in the time of the reproductive events. Males and females began to arrive during the second week of December. Males reached peak numbers (100–110) between 15 and 21 January. Females reached maximum numbers (300–350) at the end of January. About 380–430 pups were born between December 20 and February 2, and 70–80 percent of births occurred between 10 and 25 January. Copulations peaked during the third week of January. The sex ratio of adult males to adult females at peak season fluctuated from 1:3.7 to 1:4.2. By the first week of February, coinciding with the maximum number of young males (25–50), 90 percent of the adult males had abandoned the area and the number of females fluctuated greatly. Since 1980 the number of females and pups has shown a slight increase, particularly during the 1983–1984 breeding season.
Article
Bamboo sharks (Chiloscyllium plagiosum) are primarily benthic and use their relatively flexible pectoral and pelvic fins to rest on and move about the substrate. We examined the morphology of the pectoral fins and investigated their locomotory function to determine if pectoral fin function during both benthic station-holding and pelagic swimming differs from fin function described previously in leopard sharks, Triakis semifasciata. We used three-dimensional kinematics and digital particle image velocimetry (DPIV) to quantify pectoral fin function in five white-spotted bamboo sharks, C. plagiosum, during four behaviors: holding station on the substrate, steady horizontal swimming, and rising and sinking during swimming. During benthic station-holding in current flow, bamboo sharks decrease body angle and adjust pectoral fin angle to shed a clockwise fluid vortex. This vortex generates negative lift more than eight times that produced during open water vertical maneuvering and also results in an upstream flow that pushes against the posterior surface of the pectoral fin to oppose drag. In contrast, there is no evidence of significant lift force in the wake of the pectoral fin during steady horizontal swimming. The pectoral fin is held concave downward and at a negative dihedral angle during steady horizontal swimming, promoting maneuverability rather than stability, although this negative dihedral angle is much less than that observed previously in sturgeon and leopard sharks. During sinking, the pectoral fins are held concave upward and shed a clockwise vortex with a negative lift force, while in rising the pectoral fin is held concave downward and sheds a counterclockwise vortex with a positive lift force. Bamboo sharks appear to sacrifice maneuverability for stability when locomoting in the water column and use their relatively flexible fins to generate strong negative lift forces when holding position on the substrate and to enhance stability when swimming in the water column. J. Morphol. 249:195–209, 2001. © 2001 Wiley-Liss, Inc.
Article
The escape behaviour of solitary herring (Clupea harengus L.) startled by a sound stimulus was observed by means of high-speed video-filming. The results were compared with data from a previous study on the escape behaviour of schooling herring. Escape responses were divided into “away responses” and “towards responses” according to the orientation of the C-bend of the body relative to the stimulus. The proportion of away responses was smaller for solitary than for schooling herring. In solitary herring, the subsequent escape trajectories of fish making initial away responses showed a bimodal pattern of distribution, with modes at 130 and 180° from the stimulus. Trajectories following towards responses, however, were mainly within the semicircle directed at the stimulus, and their pattern of distribution differed from that of away responses. This result contrasts with observations on schooling herring, whose trajectories following both initial away and towards responses are directed away from the stimulus. In addition, we measured the response latency, defined as the interval of time between stimulus presentation and the first detectable movement of the fish. Solitary herring showed a higher proportion of short-latency responses (latency <50 ms) than schooling herring. Different behaviours appear to be exhibited by herring depending on whether they are solitary or within a school. We hypothesize that schooling may raise the threshold for initiation of fast escape responses, giving longer latencies and slower responses which are more appropriate in their directionality and reduce the possibility of collisions with neighbours. In addition, we suggest that schooling behaviour enhances the directionality and co-ordination of the escape response of the whole school, possibly increasing the probability of surviving a predator attack.
Article
The large size, high trophic level and wide distribution of Hexanchiformes (cow and frilled sharks) should position this order as important apex predators in coastal and deep-water ecosystems. This review synthesizes available information on Hexanchiformes, including information not yet published, with the purpose of evaluating their conservation status and assessing their ecological roles in the dynamics of marine ecosystems. Comprising six species, this group has a wide global distribution, with members occurring from shallow coastal areas to depths of c. 2500 m. The limited information available on their reproductive biology suggests that they could be vulnerable to overexploitation (e.g. small litter sizes for most species and suspected long gestation periods). Most of the fishing pressure exerted on Hexanchiformes is in the form of commercial by-catch or recreational fishing. Comprehensive stock and impact assessments are unavailable for most species in most regions due to limited information on life history and catch and abundance time series. When hexanchiform species have been commercially harvested, however, they have been unable to sustain targeted fisheries for long periods. The potentially high vulnerability to intense fishing pressure warrants a conservative exploitation of this order until thorough quantitative assessments are conducted. At least some species have been shown to be significant apex predators in the systems they inhabit. Should Hexanchiformes be removed from coastal and deep-water systems, the lack of sympatric shark species that share the same resources suggests no other species would be capable of fulfilling their apex predator role in the short term. This has potential ecosystem consequences such as meso-predator release or trophic cascades. This review proposes some hypotheses on the ecology of Hexanchiformes and their role in ecosystem dynamics, highlighting the areas where critical information is required to stimulate research directions.
Article
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 manoeuvrability in space and in time, respectively. Manoeuvrability 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 manoeuvrability 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.
Article
Escape responses are used by most fish species in order to avoid predation. Escape responses include a number of behavioral and kinematic components, such as responsiveness, reaction distance, escape latency, directionality, and distance-derived performance. All of these components can contribute to escape success. Work on the context-dependent variability has focused on reaction distance, and suggests that this component is largely determined by the relative cost and benefits of escaping (economic hypothesis). For example, reaction distance was found to depend on many factors related to perceived risk and cost of escaping, such as the attack speed and size of the predators, the proximity to refuges, and engagement in other activities (e.g., feeding). Evidence from many behavioral, kinematic, and physiological studies suggest that performance in other components of the escape response is also not always maximized. For example, escape latencies may increase in the presence of schooling neighbors, and escape speed is higher in fish that have been subject to higher predation pressure. In addition, all escape components are further modulated by the effect of environmental factors. Variability in escape components can be interpreted by using both ultimate and proximate explanations, for example, the effect of stimulus strength on escape latency can be interpreted as the triggering neural threshold varying with stimulus strength (proximate explanation) and high intensity stimuli representing higher risk to the prey (ultimate explanation). An integrative approach is suggested for a full, ecologically relevant, assessment of escape performance in fish.
Article
Given the diversity of vertebral morphologies among fishes, it is tempting to propose causal links between axial morphology and body curvature. We propose that shape and size of the vertebrae, intervertebral joints, and the body will more accurately predict differences in body curvature during swimming rather than a single meristic such as total vertebral number alone. We examined the correlation between morphological features and maximum body curvature seen during routine turns in five species of shark: Triakis semifasciata, Heterodontus francisci, Chiloscyllium plagiosum, Chiloscyllium punctatum, and Hemiscyllium ocellatum. We quantified overall body curvature using three different metrics. From a separate group of size-matched individuals, we measured 16 morphological features from precaudal vertebrae and the body. As predicted, a larger pool of morphological features yielded a more robust prediction of maximal body curvature than vertebral number alone. Stepwise linear regression showed that up to 11 features were significant predictors of the three measures of body curvature, yielding highly significant multiple regressions with r(2) values of 0.523, 0.537, and 0.584. The second moment of area of the centrum was always the best predictor, followed by either centrum length or transverse height. Ranking as the fifth most important variable in three different models, the body's total length, fineness ratio, and width were the most important non-vertebral morphologies. Without considering the effects of muscle activity, these correlations suggest a dominant role for the vertebral column in providing the passive mechanical properties of the body that control, in part, body curvature during swimming.
Article
Fast-starts are brief, sudden accelerations used by fish during predator­prey encounters. The kinematics and performance of fish during fast-start manoeuvres have received a lot of attention since they may determine the outcome of predator­prey interactions in terms of feeding success or survival. We will discuss recent progress on (1) the kinematics of escape responses and feeding strikes, (2) the fast-start performance of species with different body morphologies and from different habitats, and (3) the functional significance of fast-start kinematics and performance within the context of predator­prey interactions.
Article
The startle response is a model system for examining the neural basis of behavior because of its relatively simple neural circuit organization and kinematic pattern. In fishes, the two primary types of startle behavior differ in their initial movements. In the C-start type of startle, the fish bends into a C shape, while the S-start involves an S-shaped body bend. Although considerable research has focused on determining how the C-start is generated neurally, S-start neurobiology has not been examined. I quantify the kinematics and electromyographic patterns of the initial movements of the C-start and S-start behaviors of the muskellunge (Esox masquinongy) to test three hypotheses for how the S-start is generated. (i) The S-start is generated by the same motor neural circuit as the C-start, but passive bending of the tail causes the body to take on an S shape. (ii) The S-start is generated by the same motor neural circuit as undulatory swimming. (iii) The S-start is generated by an independent neural mechanism from that used either in the C-start or in undulatory swimming. Results from kinematics and muscle activity patterns support the third hypothesis. In the muskellunge, the S-start is a high-performance startle behavior with peak angular velocity and peak angular acceleration of its initial bending comparable with those of the C-start and higher than would be expected for undulatory swimming. The S-start motor pattern, however, is distinct from the C-start motor pattern in having simultaneous muscle activity anteriorly on one side of the body and posteriorly on the opposite side. In contrast, the C-start is characterized by simultaneous unilateral muscle activity along the full length of the body. Alternative models are proposed for S-start neural circuit organization involving reticulospinal and local control of muscle activity.
Article
The peculiar head morphology of hammerhead sharks has spawned a variety of untested functional hypotheses. One of the most intuitively appealing ideas is that the anterior foil acts, as in canard-winged aircraft, to increase maneuverability. We tested this hypothesis by determining whether juveniles of two hammerhead species (Sphyrna tiburo and S. lewini) turn more sharply, more often, and with greater velocity than a juvenile carcharhinid shark (Carcharhinus plumbeus). Although the hammerheads were more maneuverable, further investigation revealed that they do not roll their body during turns, suggesting that the cephalofoil does not act as a steering wing. We also show that hammerhead sharks demonstrate greater lateral flexure in a turn than carcharhinids, and that this flexibility may be due to cross sectional shape rather than number of vertebrae.
Elasmobranch phylogeny: a mitochondrial estimate based on 595 species
  • G J Naylor
  • J N Caira
  • K Jensen
  • K A Rosana
  • N Straube
  • C Lakner
Naylor, G. J., Caira, J. N., Jensen, K., Rosana, K. A., Straube, N., & Lakner, C. (2012). Elasmobranch phylogeny: a mitochondrial estimate based on 595 species. In J. Carrier, J. Musick, & M. Heithaus (Eds.), Biology of sharks and their relatives (pp. 125-151). CRC Press.
Biology of sharks and their relatives
  • G. J. Naylor
  • J. N. Caira
  • K. Jensen
  • K. A. Rosana
  • N. Straube
  • C. Lakner