Agonism is a broad term which encompasses many behaviours that result from, or are triggered by biological conflict between competing organisms. [1] [2] It is defined as "survivalist animal behaviour that includes aggression, defense, and avoidance [3] ". Approximately 23 shark species are capable of producing such displays when threatened by intraspecific or interspecific competitors, as an evolutionary strategy to avoid unnecessary combat. [1] [4] [2] The behavioural, postural, social and kinetic elements which comprise this complex, ritualized display can be easily distinguished from normal, or non-display behaviour, considered typical of that species' life history. [1] The display itself confers pertinent information to the foe regarding the displayer's physical fitness, body size, inborn biological weaponry, confidence and determination to fight. [1] [2] 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. [5] [2] [1] Agonistic displays are essential to the social dynamics of many biological taxa, extending far beyond sharks.
Agonistic displays are ritualized sequences of actions, produced by animals belonging to almost all biological taxa, in response to conflict with other organisms. [6] [2] [1] 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:
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. [1] 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.
The most effective triggers of agonistic behaviour in sharks include: [7] [4] [1]
The intensity and duration of the resulting display graded in proportion to the shark's perceived degree of endangerment during the conflict. [1]
Apart from species which incorporate unique elements into their display, a typical threat display will include the following four postural elements: elevated snout, pectoral fin depression, arching of the back, and lateral flexing of the body to display total size, and two locomotory elements: jerky side-to-side movements and rolling or spiral looping in the water column. [9] [6] [4] [1] 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. [10] [11] [12] [13] [5] [1]
Outside the typical display, certain species have been reported displaying the following behaviours as part of a species-specific display: [1]
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. [1]
To recognize and identify true agonistic behaviour, it is first important to understand what constitutes normal 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. [6] [2] [1] 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 - juxtaposed to normal behaviour, which appears visibly more relaxed and natural. [1]
The following table summarizes the characteristic physical elements of sharks displaying agonistic behaviour and non-displaying sharks:
Behavioural Display | Postural Elements | Kinetic Elements |
---|---|---|
Non-displaying shark |
|
|
Displaying shark
|
|
|
As an apex predator, sharks are no stranger to biological conflict. To successfully hunt and kill prey, sharks regularly encounter competing conspecific and heterospecific individuals, each willing to fight them to earn dominion over the meal, which in itself is a rare, valuable resource. [4] [2] [1] As part of any biological conflict, natural instinct 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. [1] During combat, however, animals are highly susceptible to injury which significantly reduces overall and reproductive fitness. [1] Sustained injuries impede each individual's ability to perform necessary biological functions, including foraging, courtship and mating behaviour. Thus, given the maladaptive 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. [7] [5] [2] [1]
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 adaptive agonistic behaviours to communicate their intentions during such conflict. [1] Sharks display recognizable sequences of behaviour, which mimic the initial phases of their fight sequence, to signal their degree of agitation, along with their intent to fight. [14] [4] [2] [1] The stereotyped postures and movements displayed by the shark are subsequently interpreted by competing organisms as honest, predictive signals of their level of commitment. [14] [2] [1]
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. [1] 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. [6] [1] 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. [1] The smaller opponent would withdraw after recognizing its own inadequacies during this short display period, and retreat from the dominant opponent. [1] 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. [1]
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:
Response to Conflict with Another Organism | Costs | Benefits |
---|---|---|
Agonistic display |
|
|
Physical attack |
|
|
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. [13] [12] [11] [2] [1] Of the described incidences where agonistic displays have been observed, the gray reef shark 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. [11] [1]
By definition, pseudodisplays of aggression are apparent physical displays that closely resemble agonistic behaviour patterns elicited in response to conflict, but are not themselves produced as a result of conflict. [9] [1] 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 caudal fins, jaw gaping, and accelerated, jerky movements. [15] [16] [1] 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.
A significant number of displays are mistaken for the actions produced during attempts to lessen the irritation caused by sharksuckers (Echeneis naucrates) along the length of their body. [15] [16] [1] Sharks are known to have sensitive skin, due to the high concentration of electroreceptors 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.
Sharks are extremely limited by their physiology, 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. [9] [14] [1] As a result, sharks rely on a select few features to accomplish all necessary biological functions. Due to these limitations, the available appendages inherit 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. [14] [9] [1] 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. [1] 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.
In biology, a weapon is a specialized physical trait that is used by animals to compete with other individuals for resources. Most commonly, the term refers to structures that males use to fight other males off for access to mates. They can also be used to defend resources in intraspecific competition, or to ward off predators. Examples of weapons include horns and antlers, both among the most recognizable weapons, though even within those categories, the structure of the specific weaponry is often unique to the species, with a wide variety of designs observed across many genera.
Requiem sharks are sharks of the family Carcharhinidae in the order Carcharhiniformes. They are migratory, live-bearing sharks of warm seas and include such species as the bull shark, lemon shark, blacktip shark, and whitetip reef shark.
The grey reef shark or gray reef shark is a species of requiem shark, in the family Carcharhinidae. One of the most common reef sharks in the Indo-Pacific, it is found as far east as Easter Island and as far west as South Africa. This species is most often seen in shallow water near the drop-offs of coral reefs. It has the typical "reef shark" shape, with a broad, round snout and large eyes. It can be distinguished from similar species by the plain or white-tipped first dorsal fin, the dark tips on the other fins, the broad, black rear margin on the tail fin, and the lack of a ridge between the dorsal fins. Most individuals are less than 1.88 m (6.2 ft) long.
The silky shark, also known by numerous names such as blackspot shark, gray whaler shark, olive shark, ridgeback shark, sickle shark, sickle-shaped shark and sickle silk shark, is a species of requiem shark, in the family Carcharhinidae, named for the smooth texture of its skin. It is one of the most abundant sharks in the pelagic zone, and can be found around the world in tropical waters. Highly mobile and migratory, this shark is most often found over the edge of the continental shelf down to 50 m (164 ft). The silky shark has a slender, streamlined body and typically grows to a length of 2.5 m. It can be distinguished from other large requiem sharks by its relatively small first dorsal fin with a curving rear margin, its tiny second dorsal fin with a long free rear tip, and its long, sickle-shaped pectoral fins. It is a deep, metallic bronze-gray above and white below.
Animal communication is the transfer of information from one or a group of animals to one or more other animals that affects the current or future behavior of the receivers. Information may be sent intentionally, as in a courtship display, or unintentionally, as in the transfer of scent from the predator to prey with kairomones. Information may be transferred to an "audience" of several receivers. Animal communication is a rapidly growing area of study in disciplines including animal behavior, sociology, neurology, and animal cognition. Many aspects of animal behavior, such as symbolic name use, emotional expression, learning, and sexual behavior, are being understood in new ways.
The whitetip reef shark is a species of requiem shark, in the family Carcharhinidae, and the only member of its genus. A small shark that does not usually exceed 1.6 m (5.2 ft) in length, this species is easily recognizable by its slender body and short but broad head, as well as tubular skin flaps beside the nostrils, oval eyes with vertical pupils, and white-tipped dorsal and caudal fins. One of the most common sharks found on Indo-Pacific coral reefs, the whitetip reef shark occurs as far west as South Africa and as far east as Central America. It is typically found on or near the bottom in clear water, at a depth of 8–40 m (26–131 ft).
Apparent death is a behavior in which animals take on the appearance of being dead. It is an immobile state most often triggered by a predatory attack and can be found in a wide range of animals from insects and crustaceans to mammals, birds, reptiles, amphibians, and fish. Apparent death is separate from the freezing behavior seen in some animals.
The silvertip shark is a large species of requiem shark, in the family Carcharhinidae, with a fragmented distribution throughout the tropical Indian and Pacific Oceans. It is often encountered around offshore islands and coral reefs, and has been known to dive to a depth of 800 m (2,600 ft). The silvertip shark resembles a larger and bulkier grey reef shark, but can be easily identified by the prominent white margins on its fins. It attains a maximum length of 3 m (10 ft).
The blacktip shark is a species of requiem shark, and part of the family Carcharhinidae. It is common to coastal tropical and subtropical waters around the world, including brackish habitats. Genetic analyses have revealed substantial variation within this species, with populations from the western Atlantic Ocean isolated and distinct from those in the rest of its range. The blacktip shark has a stout, fusiform body with a pointed snout, long gill slits, and no ridge between the dorsal fins. Most individuals have black tips or edges on the pectoral, dorsal, pelvic, and caudal fins. It usually attains a length of 1.5 m (4.9 ft).
Display behaviour is a set of ritualized behaviours that enable an animal to communicate to other animals about specific stimuli. Such ritualized behaviours can be visual, but many animals depend on a mixture of visual, audio, tactical and chemical signals. Evolution has tailored these stereotyped behaviours to allow animals to communicate both conspecifically and interspecifically which allows for a broader connection in different niches in an ecosystem. It is connected to sexual selection and survival of the species in various ways. Typically, display behaviour is used for courtship between two animals and to signal to the female that a viable male is ready to mate. In other instances, species may make territorial displays, in order to preserve a foraging or hunting territory for its family or group. A third form is exhibited by tournament species in which males will fight in order to gain the 'right' to breed. Animals from a broad range of evolutionary hierarchies avail of display behaviours - from invertebrates such as the simple jumping spider to the more complex vertebrates like the harbour seal.
The Caribbean reef shark is a species of requiem shark, belonging to the family Carcharhinidae. It is found in the tropical waters of the western Atlantic Ocean from Florida to Brazil, and is the most commonly encountered reef shark in the Caribbean Sea. With a robust, streamlined body typical of the requiem sharks, this species is difficult to tell apart from other large members of its family such as the dusky shark and the silky shark. Distinguishing characteristics include dusky-colored fins without prominent markings, a short free rear tip on the second dorsal fin, and tooth shape and number.
The Galapagos shark is a species of requiem shark, in the family Carcharhinidae, found worldwide. It favors clear reef environments around oceanic islands, where it is often the most abundant shark species. A large species that often reaches 3.0 m (9.8 ft), the Galapagos reef shark has a typical fusiform "reef shark" shape and is very difficult to distinguish from the dusky shark and the grey reef shark. An identifying character of this species is its tall first dorsal fin, which has a slightly rounded tip and originates over the rear tips of the pectoral fins.
The blacknose shark is a species of requiem shark, belonging to the family Carcharhinidae, common in the tropical and subtropical waters of the western Atlantic Ocean. This species generally inhabits coastal seagrass, sand, or rubble habitats, with adults preferring deeper water than juveniles. A small shark typically measuring 1.3 m (4.3 ft) long, the blacknose has a typical streamlined "requiem shark" shape with a long, rounded snout, large eyes, and a small first dorsal fin. Its common name comes from a characteristic black blotch on the tip of its snout, though this may be indistinct in older individuals.
One of the first shark species to be protected was the grey nurse shark. The biology, distribution and conservation of this species are dealt with in the following paragraphs with a main focus on Australia as it was here it first became protected.
Cleaner fish are fish that show a specialist feeding strategy by providing a service to other species, referred to as clients, by removing dead skin, ectoparasites, and infected tissue from the surface or gill chambers. This example of cleaning symbiosis represents mutualism and cooperation behaviour, an ecological interaction that benefits both parties involved. However, the cleaner fish may consume mucus or tissue, thus creating a form of parasitism called cheating. The client animals are typically fish of a different species, but can also be aquatic reptiles, mammals, or octopuses. A wide variety of fish including wrasse, cichlids, catfish, pipefish, lumpsuckers, and gobies display cleaning behaviors across the globe in fresh, brackish, and marine waters but specifically concentrated in the tropics due to high parasite density. Similar behaviour is found in other groups of animals, such as cleaner shrimps.
Agonistic behaviour is any social behaviour related to fighting. The term has broader meaning than aggressive behaviour because it includes threats, displays, retreats, placation, and conciliation. The term "agonistic behaviour" was first defined and used by J.P Scott and Emil Fredericson in 1951 in their paper "The Causes of Fighting in Mice and Rats" in Physiological Zoology.Agonistic behaviour is seen in many animal species because resources including food, shelter, and mates are often limited.
A courtship display is a set of display behaviors in which an animal, usually a male, attempts to attract a mate; the mate exercises choice, so sexual selection acts on the display. These behaviors often include ritualized movement ("dances"), vocalizations, mechanical sound production, or displays of beauty, strength, or agonistic ability.
Ritualized aggression or ritualized fighting is when animals use a range of behaviours as posture or warning but without engaging in serious aggression or fighting, which would be expensive in terms of energy and the risk of injury. Ritualized aggression involves a graded series of behaviours or displays that include threatening gestures and occasionally posturing physical actions such as inhibited (non-injurious) bites.
Communication occurs when an animal produces a signal and uses it to influences the behaviour of another animal. A signal can be any behavioural, structural or physiological trait that has evolved specifically to carry information about the sender and/or the external environment and to stimulate the sensory system of the receiver to change their behaviour. A signal is different from a cue in that cues are informational traits that have not been selected for communication purposes. For example, if an alerted bird gives a warning call to a predator and causes the predator to give up the hunt, the bird is using the sound as a signal to communicate its awareness to the predator. On the other hand, if a rat forages in the leaves and makes a sound that attracts a predator, the sound itself is a cue and the interaction is not considered a communication attempt.
Gaping is a common form of behavior in the animal kingdom, in which an animal opens its mouth widely and displays the interior of its mouth, for any of various purposes. This may be a form of deimatic behaviour, colloquially known as a startle display or threat display, as it enlarges the appearance of the animal, and for those with teeth it shows the threat that these represent. Animals may also use gaping as part of a courtship display, or to otherwise communicate with each other. Some animals have evolved features which make gaping behavior more visually effective. For example, "[i]n many species of reptile, the oral mucosa may be a bright color that serves to distract the predator". Gaping is part of the shark agonistic display, and is also found in snakes such as the cottonmouth, and in birds ranging from seagulls to puffins to roosters.
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