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from <a href="/w/index.php?title=Shark_threat_display&redirect=no" class="mw-redirect" title="Shark threat display">Shark threat display</a>)</span></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Animal behavior</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Grey_Reef_Shark_sculpture_Hull_Aug_2019.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/28/Grey_Reef_Shark_sculpture_Hull_Aug_2019.jpg/220px-Grey_Reef_Shark_sculpture_Hull_Aug_2019.jpg" decoding="async" width="220" height="163" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/28/Grey_Reef_Shark_sculpture_Hull_Aug_2019.jpg/330px-Grey_Reef_Shark_sculpture_Hull_Aug_2019.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/28/Grey_Reef_Shark_sculpture_Hull_Aug_2019.jpg/440px-Grey_Reef_Shark_sculpture_Hull_Aug_2019.jpg 2x" data-file-width="3007" data-file-height="2232" /></a><figcaption>Postural configuration of a Gray Reef Shark as it displays agonistic behaviour, in a sculpture</figcaption></figure> <p><a href="/wiki/Agonistic_behaviour" title="Agonistic behaviour">Agonism</a> is a broad term which encompasses many behaviours that result from, or are triggered by biological conflict between competing organisms.<sup id="cite_ref-:0_1-0" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_2-0" class="reference"><a href="#cite_note-:1-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> It is defined as "survivalist animal behaviour that includes aggression, defense, and avoidance<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup>". Approximately 23 shark species are capable of producing such displays when threatened by <a href="/wiki/Intraspecific_competition" title="Intraspecific competition">intraspecific</a> or <a href="/wiki/Interspecific_competition" title="Interspecific competition">interspecific</a> competitors, as an evolutionary strategy to avoid unnecessary combat.<sup id="cite_ref-:0_1-1" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:2_4-0" class="reference"><a href="#cite_note-:2-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_2-1" class="reference"><a href="#cite_note-:1-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> The behavioural, postural, social and kinetic elements which comprise this complex, <a href="/wiki/Ritualization" title="Ritualization">ritualized</a> display can be easily distinguished from normal, or non-display behaviour, considered typical of that species' life history.<sup id="cite_ref-:0_1-2" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The display itself confers <a href="/wiki/Pertinent" class="mw-redirect" title="Pertinent">pertinent</a> information to the foe regarding the displayer's physical fitness, body size, inborn biological weaponry, confidence and determination to fight.<sup id="cite_ref-:0_1-3" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_2-2" class="reference"><a href="#cite_note-:1-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> This behaviour is advantageous because it is much less biologically taxing for an individual to display its intention to fight than the injuries it would sustain during conflict.<sup id="cite_ref-:3_5-0" class="reference"><a href="#cite_note-:3-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_2-3" class="reference"><a href="#cite_note-:1-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_1-4" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Agonistic displays are essential to the social dynamics of many biological taxa, extending far beyond sharks. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Characteristics">Characteristics</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shark_agonistic_display&action=edit&section=1" title="Edit section: Characteristics"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Definition">Definition</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shark_agonistic_display&action=edit&section=2" title="Edit section: Definition"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Attacking_great_white_shark.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/79/Attacking_great_white_shark.jpg/220px-Attacking_great_white_shark.jpg" decoding="async" width="220" height="279" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/79/Attacking_great_white_shark.jpg/330px-Attacking_great_white_shark.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/79/Attacking_great_white_shark.jpg/440px-Attacking_great_white_shark.jpg 2x" data-file-width="606" data-file-height="768" /></a><figcaption>Clear agonistic behaviour observed in Great White Shark</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Shark_threat_display.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/83/Shark_threat_display.svg/220px-Shark_threat_display.svg.png" decoding="async" width="220" height="212" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/83/Shark_threat_display.svg/330px-Shark_threat_display.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/83/Shark_threat_display.svg/440px-Shark_threat_display.svg.png 2x" data-file-width="591" data-file-height="570" /></a><figcaption>Postural elements of the agonistic display of the Gray Reef Shark</figcaption></figure> <p>Agonistic displays are ritualized sequences of actions, produced by animals belonging to almost all biological taxa, in response to conflict with other organisms.<sup id="cite_ref-:4_6-0" class="reference"><a href="#cite_note-:4-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_2-4" class="reference"><a href="#cite_note-:1-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_1-5" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> If challenged or threatened, animals may employ a suite of adaptive behaviours, which are used to reinforce the chances of their own survival. Behaviours which arise from agonistic conflict include: </p> <ul><li>fight or flight response</li> <li>threat display to warn competitors and signal honest intentions</li> <li>defence behaviour</li> <li>simulated <a href="/wiki/Paralysis" title="Paralysis">paralysis</a></li> <li>avoidance behaviour</li> <li>withdrawal</li> <li>settling behaviour.<sup id="cite_ref-:0_1-6" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup></li></ul> <p>Each of these listed strategies constitute some manifestation of agonistic behaviour, and have been observed in numerous shark species, among many higher taxa in Kingdom Animalia.<sup id="cite_ref-:0_1-7" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Displays of this nature are influenced and reinforced by natural selection, as an optimal strategy for avoiding physical conflict, and the costs of such interactions. In nature, agonistic behaviour is both species and situation-specific, and is often expressed as a graded response to imposed threats. </p> <div class="mw-heading mw-heading3"><h3 id="Triggers">Triggers</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shark_agonistic_display&action=edit&section=3" title="Edit section: Triggers"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The most effective triggers of agonistic behaviour in sharks include:<sup id="cite_ref-:6_7-0" class="reference"><a href="#cite_note-:6-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:2_4-1" class="reference"><a href="#cite_note-:2-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_1-8" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <ul><li>hunger</li> <li>crowding by human divers (independent or group dives) and submersible machinery</li> <li>sustained targeted pursuit</li> <li>invasion of the shark's idiosphere, without appropriate warning or consent<sup id="cite_ref-:5_8-0" class="reference"><a href="#cite_note-:5-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup></li> <li>natural competition for resources with other organisms</li> <li>obstructions on the <a href="/wiki/Benthic_zone" title="Benthic zone">benthic</a> floor which block escape routes or reduce scope of visibility</li> <li>crowding by boats.</li></ul> <p>The intensity and duration of the resulting display graded in proportion to the shark's perceived degree of endangerment during the conflict.<sup id="cite_ref-:0_1-9" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Physical_characteristics">Physical characteristics</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shark_agonistic_display&action=edit&section=4" title="Edit section: Physical characteristics"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Apart from species which incorporate unique elements into their display, a typical threat display will include the following four postural elements: elevated snout, <a href="/wiki/Fish_fin" title="Fish fin">pectoral fin</a> depression, arching of the back, and lateral flexing of the body to display total size, and two <a href="/wiki/Animal_locomotion" title="Animal locomotion">locomotory</a> elements: jerky side-to-side movements and rolling or spiral looping in the water column.<sup id="cite_ref-:7_9-0" class="reference"><a href="#cite_note-:7-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:4_6-1" class="reference"><a href="#cite_note-:4-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:2_4-2" class="reference"><a href="#cite_note-:2-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_1-10" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> There are always exceptions to this rule. Certain species will display unique visual cues, outside of the displays discussed here, as individual species face their own biological stresses that drive the selection and reinforcement of new adaptive behaviours.<sup id="cite_ref-:8_10-0" class="reference"><a href="#cite_note-:8-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:9_11-0" class="reference"><a href="#cite_note-:9-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:10_12-0" class="reference"><a href="#cite_note-:10-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:11_13-0" class="reference"><a href="#cite_note-:11-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:3_5-1" class="reference"><a href="#cite_note-:3-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_1-11" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>Outside the typical display, certain species have been reported displaying the following behaviours as part of a species-specific display:<sup id="cite_ref-:0_1-12" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col"> <ul><li>stiff posture</li> <li>tail flexure</li> <li>tail depression</li> <li>head shaking</li> <li>jaw gaping</li> <li><a href="/wiki/Ritual" title="Ritual">ritualistic</a> jaw snapping</li> <li>open jaw tooth raking</li> <li>gill pouch billowing</li> <li>torso thrusting</li> <li>clasper flexion</li> <li>tail slapping</li> <li>flank displaying</li> <li>body shivering</li> <li>reduced swimming efficiency</li> <li>charging</li> <li>ramming with snout</li> <li>rapid withdrawal from threat</li></ul> </div> <p>Regardless of the precise sequence, agonistic displays function as accurate, predictive warnings to competitors which communicate a willingness to engage in more intense behaviour, if further provoked. Displays of this nature will always either result in a fleeing behaviour, or rapid, slashing attack from the displaying individual.<sup id="cite_ref-:0_1-13" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Characteristics_of_normal_behaviour">Characteristics of normal behaviour</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shark_agonistic_display&action=edit&section=5" title="Edit section: Characteristics of normal behaviour"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>To recognize and identify true agonistic behaviour, it is first important to understand what constitutes <i>normal</i> behaviour in sharks, specific to the lifestyle of the species of interest. For sharks, normal behaviour is considerably different than the exaggerated appearance of the agonistic display.<sup id="cite_ref-:4_6-2" class="reference"><a href="#cite_note-:4-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_2-5" class="reference"><a href="#cite_note-:1-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_1-14" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> A shark which is in distress will behave in a manner that is easily recognizable, due to the sheer oddity of the sequence of movements and torsional elements characteristic of agonistic displays - <a href="/wiki/Juxtaposition" title="Juxtaposition">juxtaposed</a> to normal behaviour, which appears visibly more relaxed and natural.<sup id="cite_ref-:0_1-15" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>The following table summarizes the characteristic physical elements of sharks displaying agonistic behaviour and non-displaying sharks: </p> <table class="wikitable"> <caption>Table 1: Comparison of postural and kinematic elements of sharks exhibiting both normal and agonistic behaviour<sup id="cite_ref-:0_1-16" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </caption> <tbody><tr> <th>Behavioural Display </th> <th>Postural Elements </th> <th>Kinetic Elements </th></tr> <tr> <td><b>Non-displaying shark</b> </td> <td> <ul><li>Relaxed</li> <li>Body faces a single direction</li> <li>Pectoral fins positioned perpendicular to the body, and both project horizontally outwards</li></ul> </td> <td> <ul><li>Moving in a single direction at a comfortable pace</li> <li>Pectoral fins can be flexed or depressed, depending on the direction of steering</li></ul> </td></tr> <tr> <td><b>Displaying shark</b> <ul><li>Displays are graded proportionally in response to the perceived severity of the threat or conflict</li></ul> </td> <td> <ul><li>Hunched, tense posture</li> <li>Arched back</li> <li>Raised snout</li> <li>Exaggerated, bilateral depression of both pectoral fins</li> <li>Contorted posture</li> <li>Gill billowing</li> <li>Jaw gaping</li></ul> </td> <td> <ul><li>Jerky, rapid movements</li> <li>Exaggerated swimming patterns</li> <li>Body rolling or spiral looping</li> <li>Stiff movements</li></ul> </td></tr></tbody></table> <div class="mw-heading mw-heading3"><h3 id="Evolutionary_relevance">Evolutionary relevance</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shark_agonistic_display&action=edit&section=6" title="Edit section: Evolutionary relevance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>As an <a href="/wiki/Apex_predator" title="Apex predator">apex</a> predator, sharks are no stranger to biological conflict. To successfully hunt and kill prey, sharks regularly encounter competing <a href="/wiki/Conspecific" class="mw-redirect" title="Conspecific">conspecific</a> and heterospecific individuals, each willing to fight them to earn dominion over the meal, which in itself is a rare, valuable resource.<sup id="cite_ref-:2_4-3" class="reference"><a href="#cite_note-:2-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_2-6" class="reference"><a href="#cite_note-:1-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_1-17" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> As part of any biological conflict, natural <a href="/wiki/Instinct" title="Instinct">instinct</a> dictates that the shark fights to outcompete their competitors, or to subdue prey. In the past, combat was the sole means to determine which competing organism attained dominion over valuable resources, including access to prey, territory and reproductive partners.<sup id="cite_ref-:0_1-18" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> During combat, however, animals are highly susceptible to injury which significantly reduces overall and reproductive fitness.<sup id="cite_ref-:0_1-19" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Sustained injuries impede each individual's ability to perform necessary biological functions, including <a href="/wiki/Foraging" title="Foraging">foraging</a>, <a href="/wiki/Courtship_display" title="Courtship display">courtship</a> and <a href="/wiki/Mating" title="Mating">mating</a> behaviour. Thus, given the <a href="/wiki/Maladaptive_behavior" class="mw-redirect" title="Maladaptive behavior">maladaptive</a> nature of physical combat, animals have since developed refined methods to communicate their intent and willingness to fight, as a strategy to evade suffering the costs of physical combat.<sup id="cite_ref-:6_7-1" class="reference"><a href="#cite_note-:6-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:3_5-2" class="reference"><a href="#cite_note-:3-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_2-7" class="reference"><a href="#cite_note-:1-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_1-20" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>Through the process of ritualization, the initial sequences of the animal's fighting behaviour, referred to as intention movements, are received by other animals as sign stimuli that the signaller is ready to fight. Through evolutionary time, sharks specifically have developed a suite of <a href="/wiki/Adaptive_behavior" title="Adaptive behavior">adaptive</a> agonistic behaviours to communicate their intentions during such conflict.<sup id="cite_ref-:0_1-21" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Sharks display recognizable sequences of behaviour, which <a href="/wiki/Mimicry" title="Mimicry">mimic</a> the initial phases of their fight sequence, to signal their degree of agitation, along with their intent to fight.<sup id="cite_ref-:12_14-0" class="reference"><a href="#cite_note-:12-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:2_4-4" class="reference"><a href="#cite_note-:2-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_2-8" class="reference"><a href="#cite_note-:1-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_1-22" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/Stereotype" title="Stereotype">stereotyped</a> postures and movements displayed by the shark are subsequently interpreted by competing organisms as honest, predictive signals of their level of commitment.<sup id="cite_ref-:12_14-1" class="reference"><a href="#cite_note-:12-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_2-9" class="reference"><a href="#cite_note-:1-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_1-23" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>Producing the agonistic display incurs very slight costs for the displaying organism, but are performed nevertheless, as they mitigate the potential risks of sustaining a life-threatening injuries in combat.<sup id="cite_ref-:0_1-24" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Instead of fleeing or attacking the competitor, a displaying organism sacrifices its competitive edge to perform the display, to warn and hopefully deter the challenger.<sup id="cite_ref-:4_6-3" class="reference"><a href="#cite_note-:4-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_1-25" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> If the displaying organism is larger or more robust than its competitor, the display would save it from having to fight to prove its dominance.<sup id="cite_ref-:0_1-26" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The smaller opponent would withdraw after recognizing its own inadequacies during this short display period, and retreat from the dominant opponent.<sup id="cite_ref-:0_1-27" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> In this sense, displays are evolutionarily beneficial as they cost very little in the grand-scheme, but maintain the potential benefit of not sustaining life-threatening injuries during combat, which would negatively impact the organism's fitness.<sup id="cite_ref-:0_1-28" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>The following is a summary of the costs and benefits attributed to shark threat displays in comparison to mounting a physical attack as a primary response to natural conflict: </p> <table class="wikitable"> <caption>Table 2: Summary of costs and benefits incurred by sharks performing agonistic behavioural displays in place of mounting a physical attack<sup id="cite_ref-:0_1-29" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </caption> <tbody><tr> <th>Response to Conflict with Another Organism </th> <th>Costs </th> <th>Benefits </th></tr> <tr> <td>Agonistic display </td> <td> <ul><li>Reduced speed and motility while swimming</li> <li>Reduces energetic efficiency, since the display is costly</li> <li>Displaying might put the shark at risk, if this comes at the cost of a potential escape <ul><li>If opponent is not intimidated by the display, they might stay and harm the shark</li> <li>Shark sacrifices the chance to escape by performing display as an intimidation tactic</li></ul></li> <li>Costs of display are real, while the potential benefits of displaying are only a possibility rather than a guarantee</li></ul> </td> <td> <ul><li>An agonistic display could potentially eliminate the need to engage in physical combat</li> <li>Avoiding conflict is much less risky than engaging in conflict, and is the optimal outcome (having one animal become intimidated to the point of retreat)</li></ul> </td></tr> <tr> <td>Physical attack </td> <td> <ul><li>Shark may sustain life-threatening injuries that have severe health consequences</li> <li>Injuries greatly reduce individual fitness and physical ability</li> <li>Cost of under-estimating the opponent's fighting ability could be lethal</li></ul> </td> <td> <ul><li>Animal may gain independent rights to some valued resource (i.e. territory, mate, food item)</li> <li>Experiencing a victory increases the likelihood of future victories</li> <li>Gained tactical fighting experience</li></ul> </td></tr></tbody></table> <div class="mw-heading mw-heading2"><h2 id="Case_studies">Case studies</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shark_agonistic_display&action=edit&section=7" title="Edit section: Case studies"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Gray_reef_shark">Gray reef shark</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shark_agonistic_display&action=edit&section=8" title="Edit section: Gray reef shark"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Available literature suggests that the standardized threat display has been observed and appropriately documented in many shark species. Additionally, some express dissimilar behavioural patterns, in a species-specific context.<sup id="cite_ref-:11_13-1" class="reference"><a href="#cite_note-:11-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:10_12-1" class="reference"><a href="#cite_note-:10-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:9_11-1" class="reference"><a href="#cite_note-:9-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_2-10" class="reference"><a href="#cite_note-:1-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_1-30" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Of the described incidences where agonistic displays have been observed, the <a href="/wiki/Grey_reef_shark" title="Grey reef shark">gray reef shark</a> is the most commonly reported, as they hold a reputation for having a very exaggerated, recognizable display, which incorporates all of the most commonly reported physical and dynamic elements. Thus, gray reef sharks are often selected as model organisms to describe incidences of agonistic behaviour in sharks, as theirs is the most visually distinctive, providing a near perfect representation of the behaviour.<sup id="cite_ref-:9_11-2" class="reference"><a href="#cite_note-:9-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_1-31" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Sharksucker_(Echeneis_naucrates)_(46722837981).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/17/Sharksucker_%28Echeneis_naucrates%29_%2846722837981%29.jpg/220px-Sharksucker_%28Echeneis_naucrates%29_%2846722837981%29.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/17/Sharksucker_%28Echeneis_naucrates%29_%2846722837981%29.jpg/330px-Sharksucker_%28Echeneis_naucrates%29_%2846722837981%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/17/Sharksucker_%28Echeneis_naucrates%29_%2846722837981%29.jpg/440px-Sharksucker_%28Echeneis_naucrates%29_%2846722837981%29.jpg 2x" data-file-width="3900" data-file-height="2925" /></a><figcaption>Sharksucker (Echeneis naucrates)</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Sharksucker-induced_agonistic_pseudodisplays">Sharksucker-induced agonistic pseudodisplays</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shark_agonistic_display&action=edit&section=9" title="Edit section: Sharksucker-induced agonistic pseudodisplays"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>By definition, pseudodisplays of aggression are <i>apparent</i> physical displays that closely resemble agonistic behaviour patterns elicited in response to conflict, but are not themselves produced as a result of conflict.<sup id="cite_ref-:7_9-1" class="reference"><a href="#cite_note-:7-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_1-32" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> While the visual cues performed during the pseudodisplay appear visually similar to the genuine agonistic display, sharks only exhibit the true elements of this display in response to biological conflict. The overlapping kinetic and postural elements of the pseudodisplay and the typical agonistic display include twitching, shaking, body rolling, flexure or depression of pectoral and <a href="/wiki/Fish_fin" title="Fish fin">caudal fins</a>, jaw gaping, and accelerated, jerky movements.<sup id="cite_ref-:13_15-0" class="reference"><a href="#cite_note-:13-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:14_16-0" class="reference"><a href="#cite_note-:14-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_1-33" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Biologists and behavioural scientists often mistake one or more of these elements as part of their stereotyped threat display, when only a proportion of the observed instances truly are. </p><p>A significant number of displays are mistaken for the actions produced during attempts to lessen the irritation caused by <a href="/wiki/Sharksucker" class="mw-redirect" title="Sharksucker">sharksuckers</a> (<i>Echeneis naucrates)</i> along the length of their body.<sup id="cite_ref-:13_15-1" class="reference"><a href="#cite_note-:13-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:14_16-1" class="reference"><a href="#cite_note-:14-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_1-34" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Sharks are known to have sensitive skin, due to the high concentration of <a href="/wiki/Electroreception" class="mw-redirect" title="Electroreception">electroreceptors</a> distributed across their face and body. For sharks, irritation is a regular occurrence when they spend as much time as they do being cleaned by sharsuckers. In recognition that the majority of behavioural elements exhibited during a genuine agonistic encounter overlap with those performed as part of the pseudodisplay, it is easy to understand how and why accurately differentiation between the two is a challenge. </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Electroreceptors_in_a_sharks_head_fr.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e0/Electroreceptors_in_a_sharks_head_fr.svg/220px-Electroreceptors_in_a_sharks_head_fr.svg.png" decoding="async" width="220" height="135" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e0/Electroreceptors_in_a_sharks_head_fr.svg/330px-Electroreceptors_in_a_sharks_head_fr.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e0/Electroreceptors_in_a_sharks_head_fr.svg/440px-Electroreceptors_in_a_sharks_head_fr.svg.png 2x" data-file-width="1012" data-file-height="622" /></a><figcaption>Distribution of highly sensitive ampullae of Lorenzini across the shark's head and rostrum.</figcaption></figure> <p>Sharks are extremely limited by their <a href="/wiki/Physiology" title="Physiology">physiology</a>, in the sense that the distribution of key appendages or structures limit overall flexibility and maneuverability, based solely on their apparent location on the body.<sup id="cite_ref-:7_9-2" class="reference"><a href="#cite_note-:7-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:12_14-2" class="reference"><a href="#cite_note-:12-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_1-35" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> As a result, sharks rely on a select few features to accomplish all necessary biological functions. Due to these limitations, the available appendages <a href="/wiki/Heredity" title="Heredity">inherit</a> multifunctional capabilities, and the shark can perform tasks this include signalling, swimming and removal of irritants, all with the same appendages and sequences of movement.<sup id="cite_ref-:12_14-3" class="reference"><a href="#cite_note-:12-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:7_9-3" class="reference"><a href="#cite_note-:7-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_1-36" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Despite the inherent redundancy of their body plan, sharks are able to effectively communicate their intentions to nearby organisms, such as when they produce their agonistic display. However, due to the nature of this overlap in function, agonistic behaviour is often mistaken/misidentified as pseudodisplays, resulting from sharksucker-induced irritation.<sup id="cite_ref-:0_1-37" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The very slight distinction between true agonistic behaviour and the irritation-borne pseudodisplay has historically, and will continue to confound scientists dedicated to this area of research, since the two are nearly indistinguishable from each other, at first glance. </p> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shark_agonistic_display&action=edit&section=10" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width"> <ol class="references"> <li id="cite_note-:0-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-:0_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:0_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-:0_1-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-:0_1-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-:0_1-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-:0_1-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-:0_1-6"><sup><i><b>g</b></i></sup></a> <a href="#cite_ref-:0_1-7"><sup><i><b>h</b></i></sup></a> <a href="#cite_ref-:0_1-8"><sup><i><b>i</b></i></sup></a> <a href="#cite_ref-:0_1-9"><sup><i><b>j</b></i></sup></a> <a href="#cite_ref-:0_1-10"><sup><i><b>k</b></i></sup></a> <a href="#cite_ref-:0_1-11"><sup><i><b>l</b></i></sup></a> <a href="#cite_ref-:0_1-12"><sup><i><b>m</b></i></sup></a> <a href="#cite_ref-:0_1-13"><sup><i><b>n</b></i></sup></a> <a href="#cite_ref-:0_1-14"><sup><i><b>o</b></i></sup></a> <a href="#cite_ref-:0_1-15"><sup><i><b>p</b></i></sup></a> <a href="#cite_ref-:0_1-16"><sup><i><b>q</b></i></sup></a> <a href="#cite_ref-:0_1-17"><sup><i><b>r</b></i></sup></a> <a href="#cite_ref-:0_1-18"><sup><i><b>s</b></i></sup></a> <a href="#cite_ref-:0_1-19"><sup><i><b>t</b></i></sup></a> <a href="#cite_ref-:0_1-20"><sup><i><b>u</b></i></sup></a> <a href="#cite_ref-:0_1-21"><sup><i><b>v</b></i></sup></a> <a href="#cite_ref-:0_1-22"><sup><i><b>w</b></i></sup></a> <a href="#cite_ref-:0_1-23"><sup><i><b>x</b></i></sup></a> <a href="#cite_ref-:0_1-24"><sup><i><b>y</b></i></sup></a> <a href="#cite_ref-:0_1-25"><sup><i><b>z</b></i></sup></a> <a href="#cite_ref-:0_1-26"><sup><i><b>aa</b></i></sup></a> <a href="#cite_ref-:0_1-27"><sup><i><b>ab</b></i></sup></a> <a href="#cite_ref-:0_1-28"><sup><i><b>ac</b></i></sup></a> <a href="#cite_ref-:0_1-29"><sup><i><b>ad</b></i></sup></a> <a href="#cite_ref-:0_1-30"><sup><i><b>ae</b></i></sup></a> <a href="#cite_ref-:0_1-31"><sup><i><b>af</b></i></sup></a> <a href="#cite_ref-:0_1-32"><sup><i><b>ag</b></i></sup></a> <a href="#cite_ref-:0_1-33"><sup><i><b>ah</b></i></sup></a> <a href="#cite_ref-:0_1-34"><sup><i><b>ai</b></i></sup></a> <a href="#cite_ref-:0_1-35"><sup><i><b>aj</b></i></sup></a> <a href="#cite_ref-:0_1-36"><sup><i><b>ak</b></i></sup></a> <a href="#cite_ref-:0_1-37"><sup><i><b>al</b></i></sup></a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFMartin2007" class="citation journal cs1">Martin, R. Aidan (March 2007). <a rel="nofollow" class="external text" href="https://doi.org/10.1080%2F10236240601154872">"A review of shark agonistic displays: comparison of display features and implications for shark–human interactions"</a>. <i>Marine and Freshwater Behaviour and Physiology</i>. <b>40</b> (1): 3–34. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2007MFBP...40....3M">2007MFBP...40....3M</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1080%2F10236240601154872">10.1080/10236240601154872</a></span>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/1023-6244">1023-6244</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Marine+and+Freshwater+Behaviour+and+Physiology&rft.atitle=A+review+of+shark+agonistic+displays%3A+comparison+of+display+features+and+implications+for+shark%E2%80%93human+interactions&rft.volume=40&rft.issue=1&rft.pages=3-34&rft.date=2007-03&rft.issn=1023-6244&rft_id=info%3Adoi%2F10.1080%2F10236240601154872&rft_id=info%3Abibcode%2F2007MFBP...40....3M&rft.aulast=Martin&rft.aufirst=R.+Aidan&rft_id=https%3A%2F%2Fdoi.org%2F10.1080%252F10236240601154872&rfr_id=info%3Asid%2Fen.wikipedia.org%3AShark+agonistic+display" class="Z3988"></span></span> </li> <li id="cite_note-:1-2"><span class="mw-cite-backlink">^ <a href="#cite_ref-:1_2-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:1_2-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-:1_2-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-:1_2-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-:1_2-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-:1_2-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-:1_2-6"><sup><i><b>g</b></i></sup></a> <a href="#cite_ref-:1_2-7"><sup><i><b>h</b></i></sup></a> <a href="#cite_ref-:1_2-8"><sup><i><b>i</b></i></sup></a> <a href="#cite_ref-:1_2-9"><sup><i><b>j</b></i></sup></a> <a href="#cite_ref-:1_2-10"><sup><i><b>k</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGRUBERMYRBERG1977" class="citation journal cs1">GRUBER, SAMUEL H.; MYRBERG, ARTHUR A. 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"Do Sharksuckers, Echeneis Naucrates , Induce Jump Behaviour in Blacktip Sharks, Carcharhinus Limbatus ?". <i>Marine and Freshwater Behaviour and Physiology</i>. <b>36</b> (2): 111–113. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1080%2F10236240306697">10.1080/10236240306697</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/1023-6244">1023-6244</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Marine+and+Freshwater+Behaviour+and+Physiology&rft.atitle=Do+Sharksuckers%2C+Echeneis+Naucrates+%2C+Induce+Jump+Behaviour+in+Blacktip+Sharks%2C+Carcharhinus+Limbatus+%3F&rft.volume=36&rft.issue=2&rft.pages=111-113&rft.date=2003-01-01&rft_id=info%3Adoi%2F10.1080%2F10236240306697&rft.issn=1023-6244&rft.aulast=Ritter&rft.aufirst=Erich+K.&rft.au=Brunnschweiler%2C+Juerg+M.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AShark+agonistic+display" class="Z3988"></span></span> </li> <li id="cite_note-:14-16"><span class="mw-cite-backlink">^ <a href="#cite_ref-:14_16-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:14_16-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBrunnschweiler2006" class="citation journal cs1">Brunnschweiler, Juerg M. 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href="/wiki/List_of_threatened_sharks" title="List of threatened sharks">Threatened sharks</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Taxonomy</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Carcharhiniformes" title="Carcharhiniformes">Carcharhiniformes (Ground sharks)</a></li> <li><a href="/wiki/Bullhead_shark" title="Bullhead shark">Heterodontiformes (Bullhead sharks)</a></li> <li><a href="/wiki/Hexanchiformes" title="Hexanchiformes">Hexanchiformes</a></li> <li><a href="/wiki/Lamniformes" title="Lamniformes">Lamniformes (Mackerel sharks)</a></li> <li><a href="/wiki/Carpet_shark" title="Carpet shark">Orectolobiformes (Carpet sharks)</a></li> <li><a href="/wiki/Sawshark" title="Sawshark">Pristiophoriformes (Sawsharks)</a></li> <li><a href="/wiki/Squaliformes" title="Squaliformes">Squaliformes</a></li> <li><a href="/wiki/Angelshark" title="Angelshark">Squatiniformes (Angelsharks)</a></li> <li>† <a href="/wiki/Cladoselachiformes" class="mw-redirect" title="Cladoselachiformes">Cladoselachiformes</a></li> <li>† <a href="/wiki/Hybodontiformes" title="Hybodontiformes">Hybodontiformes</a></li> <li>† <a href="/wiki/Symmoriida" class="mw-redirect" title="Symmoriida">Symmoriida</a></li> <li>† <a href="/wiki/Xenacanthida" title="Xenacanthida">Xenacanthida (Xenacantiformes)</a></li> <li><a href="/wiki/List_of_sharks" title="List of sharks">List of sharks</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Human<br />interaction</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Shark_attack" title="Shark attack">Shark attack</a> <ul><li><a href="/wiki/Shark_attacks_in_Australia" title="Shark attacks in Australia">Australia</a></li> <li><a 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