171-445: A cephalopod / ˈ s ɛ f ə l ə p ɒ d / is any member of the molluscan class Cephalopoda / s ɛ f ə ˈ l ɒ p ə d ə / ( Greek plural κεφαλόποδες , kephalópodes ; "head-feet") such as a squid , octopus , cuttlefish , or nautilus . These exclusively marine animals are characterized by bilateral body symmetry , a prominent head, and a set of arms or tentacles ( muscular hydrostats ) modified from
342-405: A mantle with a significant cavity used for breathing and excretion , the presence of a radula (except for bivalves ), and the structure of the nervous system . Other than these common elements, molluscs express great morphological diversity, so many textbooks base their descriptions on a "hypothetical ancestral mollusc" (see image below). This has a single, " limpet -like" shell on top, which
513-421: A "shell vestige" or "gladius". The Incirrina have either a pair of rod-shaped stylets or no vestige of an internal shell, and some squid also lack a gladius. The shelled coleoids do not form a clade or even a paraphyletic group. The Spirula shell begins as an organic structure, and is then very rapidly mineralized. Shells that are "lost" may be lost by resorption of the calcium carbonate component. Females of
684-435: A 'rake' to comb up filaments from the sea floor. Others feed on macroscopic 'plants' such as kelp, rasping the plant surface with its radula. To employ this strategy, the plant has to be large enough for the mollusc to 'sit' on, so smaller macroscopic plants are not as often eaten as their larger counterparts. Filter feeders are molluscs that feed by straining suspended matter and food particles from water, typically by passing
855-487: A calming one and a powerful social motivation for remaining in an aggregation. Herring, for instance, will become very agitated if they are isolated from conspecifics. Because of their adaptation to schooling behaviour they are rarely displayed in aquaria . Even with the best facilities aquaria can offer they become fragile and sluggish compared to their quivering energy in wild schools. It has also been proposed that swimming in groups enhances foraging success. This ability
1026-443: A chance to escape or to never join a shoal with larger fish. It has been shown that small fish avoid joining a group with larger fish, although big fish do not avoid joining small conspecifics. This sorting mechanism based on increased quality of perception could have resulted in homogeneity of size of fish in shoals, which would increase the capacity for moving in synchrony. Predators have devised various countermeasures to undermine
1197-535: A disciplined and coordinated school, then shift back to an amorphous shoal within seconds. Such shifts are triggered by changes of activity from feeding, resting, travelling or avoiding predators. When schooling fish stop to feed, they break ranks and become shoals. Shoals are more vulnerable to predator attack. The shape a shoal or school takes depends on the type of fish and what the fish are doing. Schools that are travelling can form long thin lines, or squares or ovals or amoeboid shapes. Fast moving schools usually form
1368-592: A distance of the lab floor, enter another aquarium to feed on captive crabs, and return to their own aquarium. The brain is protected in a cartilaginous cranium. The giant nerve fibers of the cephalopod mantle have been widely used for many years as experimental material in neurophysiology ; their large diameter (due to lack of myelination ) makes them relatively easy to study compared with other animals. Many cephalopods are social creatures; when isolated from their own kind, some species have been observed shoaling with fish. Some cephalopods are able to fly through
1539-552: A diversity of backgrounds. Experiments done in Dwarf chameleons testing these hypotheses showed that chameleon taxa with greater capacity for color change had more visually conspicuous social signals but did not come from more visually diverse habitats, suggesting that color change ability likely evolved to facilitate social signaling, while camouflage is a useful byproduct. Because camouflage is used for multiple adaptive purposes in cephalopods, color change could have evolved for one use and
1710-623: A feeding ground for larger predator fish. Most upwellings are coastal, and many of them support some of the most productive fisheries in the world. Regions of notable upwelling include coastal Peru , Chile , Arabian Sea , western South Africa , eastern New Zealand and the California coast. Copepods , the primary zooplankton , are a major item on the forage fish menu. They are a group of small crustaceans found in ocean and freshwater habitats . Copepods are typically one millimetre (0.04 in) to two millimetres (0.08 in) long, with
1881-522: A flat fan shape with a mucus film between the individual tentacles, while another, Sepioteuthis sepioidea , has been observed putting the tentacles in a circular arrangement. Cephalopods have advanced vision, can detect gravity with statocysts , and have a variety of chemical sense organs. Octopuses use their arms to explore their environment and can use them for depth perception. Most cephalopods rely on vision to detect predators and prey and to communicate with one another. Consequently, cephalopod vision
SECTION 10
#17327717847702052-912: A forage fish of the smelt family found in the Atlantic and Arctic oceans. In summer, they graze on dense swarms of plankton at the edge of the ice shelf. Larger capelin also eat krill and other crustaceans . The capelin move inshore in large schools to spawn and migrate in spring and summer to feed in plankton rich areas between Iceland , Greenland , and Jan Mayen . The migration is affected by ocean currents . Around Iceland maturing capelin make large northward feeding migrations in spring and summer. The return migration takes place in September to November. The spawning migration starts north of Iceland in December or January. This theory states that groups of fish may save energy when swimming together, much in
2223-435: A group comprising cephalopods. Molluscus is used in classical Latin as an adjective only with nux ( nut ) to describe a particular type of soft nut. The use of mollusca in biological taxonomy by Jonston and later Linnaeus may have been influenced by Aristotle 's τὰ μαλάκια ta malákia (the soft ones; < μαλακός malakós "soft"), which he applied inter alia to cuttlefish . The scientific study of molluscs
2394-649: A gunshot-like popping noise, thought to function to frighten away potential predators. Cephalopods employ a similar method of propulsion despite their increasing size (as they grow) changing the dynamics of the water in which they find themselves. Thus their paralarvae do not extensively use their fins (which are less efficient at low Reynolds numbers ) and primarily use their jets to propel themselves upwards, whereas large adult cephalopods tend to swim less efficiently and with more reliance on their fins. Early cephalopods are thought to have produced jets by drawing their body into their shells, as Nautilus does today. Nautilus
2565-418: A hard shell use calcite (sometimes with traces of aragonite) to construct the eggshells. The shell consists of three layers: the outer layer (the periostracum ) made of organic matter, a middle layer made of columnar calcite , and an inner layer consisting of laminated calcite, often nacreous . In some forms the shell contains openings. In abalones there are holes in the shell used for respiration and
2736-430: A herring to eventually snap the copepod. A single juvenile herring could never catch a large copepod. A third proposed benefit of fish groups is that they serve a reproductive function. They provide increased access to potential mates, since finding a mate in a shoal does not take much energy. And for migrating fish that navigate long distances to spawn, it is likely that the navigation of the shoal, with an input from all
2907-409: A hydrodynamic advantage, the leader will be the first to the food. More recent work suggests that, after individuals at the front of the school encounter and ingest more food, they then relocate further back within the school due to the locomotor constraints generated during meal digestion. It is commonly observed that schooling fish are particularly in danger of being eaten if they are separated from
3078-451: A jet as a propulsion mechanism. Squids do not have the longitudinal muscles that octopus do. Instead, they have a tunic. This tunic is made of layers of collagen and it surrounds the top and the bottom of the mantle. Because they are made of collagen and not muscle, the tunics are rigid bodies that are much stronger than the muscle counterparts. This provides the squids some advantages for jet propulsion swimming. The stiffness means that there
3249-400: A length of 8 metres. They may terminate in a broadened, sucker-coated club. The shorter four pairs are termed arms , and are involved in holding and manipulating the captured organism. They too have suckers, on the side closest to the mouth; these help to hold onto the prey. Octopods only have four pairs of sucker-coated arms, as the name suggests, though developmental abnormalities can modify
3420-451: A loose way, with each fish swimming and foraging somewhat independently, they are nonetheless aware of the other members of the group as shown by the way they adjust behaviour such as swimming, so as to remain close to the other fish in the group. Shoaling groups can include fish of disparate sizes and can include mixed-species subgroups. If the shoal becomes more tightly organised, with the fish synchronising their swimming so they all move at
3591-403: A maximum of 120,000 species. The total number of described species is difficult to estimate because of unresolved synonymy . In 1969, David Nicol estimated the probable total number of living mollusc species at 107,000 of which were about 12,000 fresh-water gastropods and 35,000 terrestrial . The Bivalvia would comprise about 14% of the total and the other five classes less than 2% of
SECTION 20
#17327717847703762-472: A muscle, which is why they can change their skin hue as rapidly as they do. Coloration is typically stronger in near-shore species than those living in the open ocean, whose functions tend to be restricted to disruptive camouflage . These chromatophores are found throughout the body of the octopus, however, they are controlled by the same part of the brain that controls elongation during jet propulsion to reduce drag. As such, jetting octopuses can turn pale because
3933-510: A novel mechanism for spectral discrimination in cephalopods was described. This relies on the exploitation of chromatic aberration (wavelength-dependence of focal length). Numerical modeling shows that chromatic aberration can yield useful chromatic information through the dependence of image acuity on accommodation. The unusual off-axis slit and annular pupil shapes in cephalopods enhance this ability by acting as prisms which are scattering white light in all directions. In 2015, molecular evidence
4104-399: A number of paired ganglia, the visceral cords serving the internal organs and the pedal ones serving the foot. Most pairs of corresponding ganglia on both sides of the body are linked by commissures (relatively large bundles of nerves). The ganglia above the gut are the cerebral, the pleural, and the visceral, which are located above the esophagus (gullet). The pedal ganglia, which control
4275-407: A pair of statocysts , which act as balance sensors. In gastropods, it secretes mucus as a lubricant to aid movement. In forms having only a top shell, such as limpets , the foot acts as a sucker attaching the animal to a hard surface, and the vertical muscles clamp the shell down over it; in other molluscs, the vertical muscles pull the foot and other exposed soft parts into the shell. In bivalves,
4446-461: A part but supporting evidence has not been found so far. The lateral line is a line running along each side of the fish from the gill covers to the base of the tail. In laboratory experiments the lateral lines of schooling fish have been removed. They swam closer, leading to a theory that the lateral lines provide additional stimuli input when the fish get too close. The lateral-line system is very sensitive to changes in water currents and vibration in
4617-678: A rasping "tongue", the radula, and a complex digestive system in which exuded mucus and microscopic, muscle-powered "hairs" called cilia play various important roles. The generalized mollusc has two paired nerve cords , or three in bivalves . The brain , in species that have one, encircles the esophagus . Most molluscs have eyes , and all have sensors to detect chemicals, vibrations, and touch . The simplest type of molluscan reproductive system relies on external fertilization , but more complex variations occur. Nearly all produce eggs , from which may emerge trochophore larvae , more complex veliger larvae, or miniature adults. The coelomic cavity
4788-404: A result, their blood is colorless when deoxygenated and turns blue when bonded to oxygen. In oxygen-rich environments and in acidic water, hemoglobin is more efficient, but in environments with little oxygen and in low temperatures, hemocyanin has the upper hand. The hemocyanin molecule is much larger than the hemoglobin molecule, allowing it to bond with 96 O 2 or CO 2 molecules, instead of
4959-424: A risk of food poisoning, and many jurisdictions have regulations to reduce this risk. Molluscs have, for centuries, also been the source of important luxury goods, notably pearls , mother of pearl , Tyrian purple dye, and sea silk . Their shells have also been used as money in some preindustrial societies. A handful of mollusc species are sometimes considered hazards or pests for human activities. The bite of
5130-496: A shell (secondarily absent in a number of taxonomic groups, such as the nudibranchs ) that consists of mainly chitin and conchiolin (a protein hardened with calcium carbonate ), except the outermost layer, which in almost all cases is all conchiolin (see periostracum ). Molluscs never use phosphate to construct their hard parts, with the questionable exception of Cobcrephora . While most mollusc shells are composed mainly of aragonite , those gastropods that lay eggs with
5301-429: A shell-less subclass of cephalopods (squid, cuttlefish, and octopuses), have complex pigment containing cells called chromatophores which are capable of producing rapidly changing color patterns. These cells store pigment within an elastic sac which produces the color seen from these cells. Coleoids can change the shape of this sac, called the cytoelastic sacculus, which then causes changes in the translucency and opacity of
Cephalopod - Misplaced Pages Continue
5472-683: A short life-cycle, living only two or three years. Adult sardines, about two years old, mass on the Agulhas Bank where they spawn during spring and summer, releasing tens of thousands of eggs into the water. The adult sardines then make their way in hundreds of shoals towards the sub-tropical waters of the Indian Ocean . A larger shoal might be 7 kilometres (4.3 mi) long, 1.5 kilometres (0.93 mi) wide and 30 metres (98 ft) deep. Huge numbers of sharks, dolphins, tuna, sailfish, Cape fur seals and even killer whales congregate and follow
5643-413: A single school. These schools move along coastlines and traverse the open oceans. Herring schools in general have very precise arrangements which allow the school to maintain relatively constant cruising speeds. Herrings have excellent hearing, and their schools react very rapidly to a predator. The herrings keep a certain distance from a moving scuba diver or a cruising predator like a killer whale, forming
5814-456: A singular gill. Generally, the gills are rather like feathers in shape, although some species have gills with filaments on only one side. They divide the mantle cavity so water enters near the bottom and exits near the top. Their filaments have three kinds of cilia, one of which drives the water current through the mantle cavity, while the other two help to keep the gills clean. If the osphradia detect noxious chemicals or possibly sediment entering
5985-615: A startling array of fashions. As well as providing camouflage with their background, some cephalopods bioluminesce, shining light downwards to disguise their shadows from any predators that may lurk below. The bioluminescence is produced by bacterial symbionts; the host cephalopod is able to detect the light produced by these organisms. Bioluminescence may also be used to entice prey, and some species use colorful displays to impress mates, startle predators, or even communicate with one another. Cephalopods can change their colors and patterns in milliseconds, whether for signalling (both within
6156-430: A teardrop shaped body. Some scientists say they form the largest animal biomass on the planet. Copepods are very alert and evasive. They have large antennae (see photo below left). When they spread their antennae they can sense the pressure wave from an approaching fish and jump with great speed over a few centimeters. If copepod concentrations reach high levels, schooling herrings adopt a method called ram feeding . In
6327-418: A vacuole which looks like a doughnut from a spotter plane. Many species of large predatory fish also school, including many highly migratory fish , such as tuna and some oceangoing sharks . Cetaceans such as dolphins, porpoises and whales, operate in organised social groups called pods . "Shoaling behaviour is generally described as a trade-off between the anti-predator benefits of living in groups and
6498-636: A wedge shape, while shoals that are feeding tend to become circular. Forage fish are small fish which are preyed on by larger predators for food. Predators include other larger fish, seabirds and marine mammals . Typical ocean forage fish are small, filter-feeding fish such as herring , anchovies and menhaden . Forage fish compensate for their small size by forming schools. Some swim in synchronised grids with their mouths open so they can efficiently filter feed on plankton . These schools can become huge, moving along coastlines and migrating across open oceans. The shoals are concentrated food resources for
6669-415: Is a hemocoel through which blood and coelomic fluid circulate and which encloses most of the other internal organs. These hemocoelic spaces act as an efficient hydrostatic skeleton . The blood of these molluscs contains the respiratory pigment hemocyanin as an oxygen -carrier. The heart consists of one or more pairs of atria ( auricles ), which receive oxygenated blood from the gills and pump it to
6840-432: Is a notable partial exception in that it tolerates brackish water . Cephalopods are thought to be unable to live in fresh water due to multiple biochemical constraints, and in their >400 million year existence have never ventured into fully freshwater habitats. Cephalopods occupy most of the depth of the ocean, from the abyssal plains to the sea surface, and have also been found in the hadal zone . Their diversity
7011-437: Is accordingly called malacology . The name Molluscoida was formerly used to denote a division of the animal kingdom containing the brachiopods , bryozoans , and tunicates , the members of the three groups having been supposed to somewhat resemble the molluscs. As now known, these groups have no relation to molluscs, and very little to one another, so the name Molluscoida has been abandoned. The most universal features of
Cephalopod - Misplaced Pages Continue
7182-514: Is acute: training experiments have shown that the common octopus can distinguish the brightness, size, shape, and horizontal or vertical orientation of objects. The morphological construction gives cephalopod eyes the same performance as shark eyes; however, their construction differs, as cephalopods lack a cornea and have an everted retina. Cephalopods' eyes are also sensitive to the plane of polarization of light. Unlike many other cephalopods, nautiluses do not have good vision; their eye structure
7353-460: Is also capable of creating a jet by undulations of its funnel; this slower flow of water is more suited to the extraction of oxygen from the water. When motionless, Nautilus can only extract 20% of oxygen from the water. The jet velocity in Nautilus is much slower than in coleoids , but less musculature and energy is involved in its production. Jet thrust in cephalopods is controlled primarily by
7524-443: Is an elaboration of safety in numbers , and interacts with the confusion effect. A given predator attack will eat a smaller proportion of a large shoal than a small shoal. Hamilton proposed that animals aggregate because of a "selfish" avoidance of a predator and was thus a form of cover-seeking. Another formulation of the theory was given by Turner and Pitcher and was viewed as a combination of detection and attack probabilities. In
7695-529: Is an endemic snail species of the Eastern Alps . There is strong evidence for self-fertilization in the easternmost snail populations of this species. The most basic molluscan larva is a trochophore , which is planktonic and feeds on floating food particles by using the two bands of cilia around its "equator" to sweep food into the mouth, which uses more cilia to drive them into the stomach, which uses further cilia to expel undigested remains through
7866-565: Is aragonite. As for other mollusc shells or coral skeletons, the smallest visible units are irregular rounded granules. Cephalopods, as the name implies, have muscular appendages extending from their heads and surrounding their mouths. These are used in feeding, mobility, and even reproduction. In coleoids they number eight or ten. Decapods such as cuttlefish and squid have five pairs. The longer two, termed tentacles , are actively involved in capturing prey; they can lengthen rapidly (in as little as 15 milliseconds). In giant squid they may reach
8037-402: Is associated with the odontophore , a cartilaginous supporting organ. The radula is unique to the molluscs and has no equivalent in any other animal. Molluscs' mouths also contain glands that secrete slimy mucus , to which the food sticks. Beating cilia (tiny "hairs") drive the mucus towards the stomach, so the mucus forms a long string called a "food string". At the tapered rear end of
8208-427: Is constantly washing through their gills, even when the organism is stationary. The water flow is controlled by contractions of the radial and circular mantle cavity muscles. The gills of cephalopods are supported by a skeleton of robust fibrous proteins; the lack of mucopolysaccharides distinguishes this matrix from cartilage. The gills are also thought to be involved in excretion, with NH 4 being swapped with K from
8379-412: Is controversial. Fish can be obligate or facultative (optional) shoalers. Obligate shoalers, such as tunas , herrings and anchovy , spend all of their time shoaling or schooling, and become agitated if separated from the group. Facultative shoalers, such as Atlantic cod , saiths and some carangids , shoal only some of the time, perhaps for reproductive purposes. Shoaling fish can shift into
8550-510: Is essential in the final stages of a predator attack. Electro-receptive animals may localize a field source by using spatial non-uniformities. To produce separate signals, individual prey must be about five body widths apart. If objects are too close together to be distinguished, they will form a blurred image. Based on this it was suggested that schooling may confuse the ESS of predators. A third potential anti-predator effect of animal aggregations
8721-580: Is greatest near the equator (~40 species retrieved in nets at 11°N by a diversity study) and decreases towards the poles (~5 species captured at 60°N). Cephalopods are widely regarded as the most intelligent of the invertebrates and have well developed senses and large brains (larger than those of gastropods ). The nervous system of cephalopods is the most complex of the invertebrates and their brain-to-body-mass ratio falls between that of endothermic and ectothermic vertebrates. Captive cephalopods have also been known to climb out of their aquaria, maneuver
SECTION 50
#17327717847708892-424: Is highly developed, but lacks a solid lens . They have a simple " pinhole " eye through which water can pass. Instead of vision, the animal is thought to use olfaction as the primary sense for foraging , as well as locating or identifying potential mates. All octopuses and most cephalopods are considered to be color blind . Coleoid cephalopods (octopus, squid, cuttlefish) have a single photoreceptor type and lack
9063-663: Is important to schooling. The importance of vision is also indicated by the behaviour of fish who have been temporarily blinded. Schooling species have eyes on the sides of their heads, which means they can easily see their neighbours. Also, schooling species often have "schooling marks" on their shoulders or the base of their tails, or visually prominent stripes, which provide reference marks when schooling, similar in function to passive markers in artificial motion capture. However fish without these markers will still engage in schooling behaviour, though perhaps not as efficiently. Other senses are also used. Pheromones or sound may also play
9234-449: Is made of proteins and chitin reinforced with calcium carbonate , and is secreted by a mantle covering the whole upper surface. The underside of the animal consists of a single muscular "foot". Although molluscs are coelomates , the coelom tends to be small. The main body cavity is a hemocoel through which blood circulates; as such, their circulatory systems are mainly open . The "generalized" mollusc's feeding system consists of
9405-438: Is no necessary muscle flexing to keep the mantle the same size. In addition, tunics take up only 1% of the squid mantle's wall thickness, whereas the longitudinal muscle fibers take up to 20% of the mantle wall thickness in octopuses. Also because of the rigidity of the tunic, the radial muscles in squid can contract more forcefully. The mantle is not the only place where squids have collagen. Collagen fibers are located throughout
9576-433: Is one of the largest invertebrates , surpassed in weight but not in length by the colossal squid . Freshwater and terrestrial molluscs appear exceptionally vulnerable to extinction. Estimates of the numbers of non-marine molluscs vary widely, partly because many regions have not been thoroughly surveyed. There is also a shortage of specialists who can identify all the animals in any one area to species. However, in 2004
9747-667: Is reduced. They have an open circulatory system and kidney-like organs for excretion. Good evidence exists for the appearance of gastropods, cephalopods , and bivalves in the Cambrian period, 541–485.4 million years ago. However, the evolutionary history both of molluscs' emergence from the ancestral Lophotrochozoa and of their diversification into the well-known living and fossil forms are still subjects of vigorous debate among scientists. Molluscs have been and still are an important food source for humans. Toxins that can accumulate in certain molluscs under specific conditions create
9918-420: Is secreted by a mantle covering the upper surface. The underside consists of a single muscular "foot". The visceral mass, or visceropallium, is the soft, nonmuscular metabolic region of the mollusc. It contains the body organs. The mantle cavity, a fold in the mantle, encloses a significant amount of space. It is lined with epidermis, and is exposed, according to habitat , to sea, fresh water or air. The cavity
10089-653: Is seen in the structure of schools of predatory fish. Partridge and others analysed the school structure of Atlantic bluefin tuna from aerial photographs and found that the school assumed a parabolic shape, a fact that was suggestive of cooperative hunting in this species. "The reason for this is the presence of many eyes searching for the food. Fish in shoals "share" information by monitoring each other's behaviour closely. Feeding behaviour in one fish quickly stimulates food-searching behaviour in others. Fertile feeding grounds for forage fish are provided by ocean upwellings. Oceanic gyres are large-scale ocean currents caused by
10260-449: Is the "many eyes" hypothesis. This theory states that as the size of the group increases, the task of scanning the environment for predators can be spread out over many individuals. Not only does this mass collaboration presumably provide a higher level of vigilance, it could also allow more time for individual feeding. A fourth hypothesis for an anti-predatory effect of fish schools is the "encounter dilution" effect. The dilution effect
10431-551: Is the first evidence that cephalopod dermal tissues may possess the required combination of molecules to respond to light. Some squids have been shown to detect sound using their statocysts , but, in general, cephalopods are deaf. Most cephalopods possess an assemblage of skin components that interact with light. These may include iridophores, leucophores , chromatophores and (in some species) photophores . Chromatophores are colored pigment cells that expand and contract in accordance to produce color and pattern which they can use in
SECTION 60
#173277178477010602-471: Is the general term for any collection of fish that have gathered together in some locality. Fish aggregations can be structured or unstructured. An unstructured aggregation might be a group of mixed species and sizes that have gathered randomly near some local resource, such as food or nesting sites. If, in addition, the aggregation comes together in an interactive, social way, they may be said to be shoaling . Although shoaling fish can relate to each other in
10773-554: Is transmitted to humans by water snail hosts, and affects about 200 million people. Snails and slugs can also be serious agricultural pests, and accidental or deliberate introduction of some snail species into new environments has seriously damaged some ecosystems . The words mollusc and mollusk are both derived from the French mollusque , which originated from the post-classical Latin mollusca , from mollis , soft, first used by J. Jonston (Historiæ Naturalis, 1650) to describe
10944-426: Is unknown, but chromatophores are under the control of neural pathways, allowing the cephalopod to coordinate elaborate displays. Together, chromatophores and iridophores are able to produce a large range of colors and pattern displays. Cephalopods utilize chromatophores' color changing ability in order to camouflage themselves. Chromatophores allow Coleoids to blend into many different environments, from coral reefs to
11115-469: The Coriolis effect . Wind-driven surface currents interact with these gyres and the underwater topography, such as seamounts , fishing banks , and the edge of continental shelves , to produce downwellings and upwellings . These can transport nutrients which plankton thrive on. The result can be rich feeding grounds attractive to the plankton feeding forage fish. In turn, the forage fish themselves become
11286-487: The IUCN Red List of Threatened Species included nearly 2,000 endangered non-marine molluscs. For comparison, the great majority of mollusc species are marine, but only 41 of these appeared on the 2004 Red List. About 42% of recorded extinctions since the year 1500 are of molluscs, consisting almost entirely of non-marine species. Because of the great range of anatomical diversity among molluscs, many textbooks start
11457-526: The Paleozoic era , as competition with fish produced an environment where efficient motion was crucial to survival, jet propulsion has taken a back role, with fins and tentacles used to maintain a steady velocity. Whilst jet propulsion is never the sole mode of locomotion, the stop-start motion provided by the jets continues to be useful for providing bursts of high speed – not least when capturing prey or avoiding predators . Indeed, it makes cephalopods
11628-405: The blue-ringed octopus is often fatal, and that of Octopus apollyon causes inflammation that can last over a month. Stings from a few species of large tropical cone shells of the family Conidae can also kill, but their sophisticated, though easily produced, venoms have become important tools in neurological research. Schistosomiasis (also known as bilharzia, bilharziosis, or snail fever)
11799-462: The coelom , a small cavity that surrounds the heart, into which they shed ova or sperm . The nephridia extract the gametes from the coelom and emit them into the mantle cavity. Molluscs that use such a system remain of one sex all their lives and rely on external fertilization . Some molluscs use internal fertilization and/or are hermaphrodites , functioning as both sexes; both of these methods require more complex reproductive systems. C. obtusus
11970-516: The hyponome , created by a fold in the mantle. The size difference between the posterior and anterior ends of this organ control the speed of the jet the organism can produce. The velocity of the organism can be accurately predicted for a given mass and morphology of animal. Motion of the cephalopods is usually backward as water is forced out anteriorly through the hyponome, but direction can be controlled somewhat by pointing it in different directions. Some cephalopods accompany this expulsion of water with
12141-411: The lateral line organ (LLO) as well as the electrosensory system (ESS) of predators. Fin movements of a single fish act as a point-shaped wave source, emitting a gradient by which predators might localize it. Since fields of many fish will overlap, schooling should obscure this gradient, perhaps mimicking pressure waves of a larger animal, and more likely confuse the lateral line perception. The LLO
12312-580: The sparkling enope squid ( Watasenia scintillans ). It achieves color vision with three photoreceptors , which are based on the same opsin , but use distinct retinal molecules as chromophores: A1 (retinal), A3 (3-dehydroretinal), and A4 (4-hydroxyretinal). The A1-photoreceptor is most sensitive to green-blue (484 nm), the A2-photoreceptor to blue-green (500 nm), and the A4-photoreceptor to blue (470 nm) light. In 2015,
12483-400: The ventricle , which pumps it into the aorta (main artery ), which is fairly short and opens into the hemocoel. The atria of the heart also function as part of the excretory system by filtering waste products out of the blood and dumping it into the coelom as urine . A pair of metanephridia ("little kidneys") to the rear of and connected to the coelom extracts any re-usable materials from
12654-463: The Atlantic bottlenose dolphin takes this one step further with what has become known as strand feeding, where the fish are driven onto mud banks and retrieved from there. Common bottlenose dolphins have been observed using another technique. One dolphin acts as a "driver" and herds a school of fish towards several other dolphins who form a barrier. The driver dolphin slaps its fluke which makes
12825-611: The Coleoidea, the molluscan shell has been internalized or is absent, whereas in the Nautiloidea, the external shell remains. About 800 living species of cephalopods have been identified. Two important extinct taxa are the Ammonoidea (ammonites) and Belemnoidea (belemnites). Extant cephalopods range in size from the 10 mm (0.3 in) Idiosepius thailandicus to the 700 kilograms (1,500 lb) heavy Colossal squid ,
12996-554: The ability to determine color by comparing detected photon intensity across multiple spectral channels. When camouflaging themselves, they use their chromatophores to change brightness and pattern according to the background they see, but their ability to match the specific color of a background may come from cells such as iridophores and leucophores that reflect light from the environment. They also produce visual pigments throughout their body and may sense light levels directly from their body. Evidence of color vision has been found in
13167-450: The acidity of the organic shell matrix (see Mollusc shell ); shell-forming cephalopods have an acidic matrix, whereas the gladius of squid has a basic matrix. The basic arrangement of the cephalopod outer wall is: an outer (spherulitic) prismatic layer, a laminar (nacreous) layer and an inner prismatic layer. The thickness of every layer depends on the taxa. In modern cephalopods, the Ca carbonate
13338-432: The air for distances of up to 50 metres (160 ft). While cephalopods are not particularly aerodynamic, they achieve these impressive ranges by jet-propulsion; water continues to be expelled from the funnel while the organism is in the air. The animals spread their fins and tentacles to form wings and actively control lift force with body posture. One species, Todarodes pacificus , has been observed spreading tentacles in
13509-412: The anus. New tissue grows in the bands of mesoderm in the interior, so the apical tuft and anus are pushed further apart as the animal grows. The trochophore stage is often succeeded by a veliger stage in which the prototroch , the "equatorial" band of cilia nearest the apical tuft, develops into the velum ("veil"), a pair of cilia-bearing lobes with which the larva swims. Eventually, the larva sinks to
13680-454: The appearance of their surroundings is notable given that cephalopods' vision is monochromatic. Cephalopods also use their fine control of body coloration and patterning to perform complex signaling displays for both conspecific and intraspecific communication. Coloration is used in concert with locomotion and texture to send signals to other organisms. Intraspecifically this can serve as a warning display to potential predators. For example, when
13851-426: The base of the funnel means the ink can be distributed by ejected water as the cephalopod uses its jet propulsion. The ejected cloud of melanin is usually mixed, upon expulsion, with mucus , produced elsewhere in the mantle, and therefore forms a thick cloud, resulting in visual (and possibly chemosensory) impairment of the predator, like a smokescreen . However, a more sophisticated behavior has been observed, in which
14022-514: The body cavity; others, like some fish, accumulate oils in the liver; and some octopuses have a gelatinous body with lighter chloride ions replacing sulfate in the body chemistry. Squids are the primary sufferers of negative buoyancy in cephalopods. The negative buoyancy means that some squids, especially those whose habitat depths are rather shallow, have to actively regulate their vertical positions. This means that they must expend energy, often through jetting or undulations, in order to maintain
14193-672: The body structure of molluscs are a mantle with a significant body cavity used for breathing and excretion , and the organization of the nervous system. Many have a calcareous shell. Molluscs have developed such a varied range of body structures, finding synapomorphies (defining characteristics) to apply to all modern groups is difficult. The most general characteristic of molluscs is they are unsegmented and bilaterally symmetrical. The following are present in all modern molluscs: Other characteristics that commonly appear in textbooks have significant exceptions: Estimates of accepted described living species of molluscs vary from 50,000 to
14364-578: The brain is unable to achieve both controlling elongation and controlling the chromatophores. Most octopuses mimic select structures in their field of view rather than becoming a composite color of their full background. Evidence of original coloration has been detected in cephalopod fossils dating as far back as the Silurian ; these orthoconic individuals bore concentric stripes, which are thought to have served as camouflage. Devonian cephalopods bear more complex color patterns, of unknown function. Coleoids,
14535-433: The cavity by entering not only through the orifices, but also through the funnel. Squid can expel up to 94% of the fluid within their cavity in a single jet thrust. To accommodate the rapid changes in water intake and expulsion, the orifices are highly flexible and can change their size by a factor of twenty; the funnel radius, conversely, changes only by a factor of around 1.5. Some octopus species are also able to walk along
14706-559: The cell. By rapidly changing multiple chromatophores of different colors, cephalopods are able to change the color of their skin at astonishing speeds, an adaptation that is especially notable in an organism that sees in black and white. Chromatophores are known to only contain three pigments, red, yellow, and brown, which cannot create the full color spectrum. However, cephalopods also have cells called iridophores, thin, layered protein cells that reflect light in ways that can produce colors chromatophores cannot. The mechanism of iridophore control
14877-405: The cephalopod releases a cloud, with a greater mucus content, that approximately resembles the cephalopod that released it (this decoy is referred to as a pseudomorph ). This strategy often results in the predator attacking the pseudomorph, rather than its rapidly departing prey. For more information, see Inking behaviors . The ink sac of cephalopods has led to a common name of "inkfish", formerly
15048-707: The chance of individual capture), enhanced foraging success, and higher success in finding a mate. It is also likely that fish benefit from shoal membership through increased hydrodynamic efficiency. Fish use many traits to choose shoalmates. Generally they prefer larger shoals, shoalmates of their own species, shoalmates similar in size and appearance to themselves, healthy fish, and kin (when recognized). The oddity effect posits that any shoal member that stands out in appearance will be preferentially targeted by predators. This may explain why fish prefer to shoal with individuals that resemble themselves. The oddity effect thus tends to homogenize shoals. An aggregation of fish
15219-417: The confusion of casting nets, the dolphins catch a large number of fish as well. Intraspecific cooperative foraging techniques have also been observed, and some propose that these behaviours are transmitted through cultural means. Rendell & Whitehead have proposed a structure for the study of culture in cetaceans. Some whales lunge feed on bait balls. Lunge feeding is an extreme feeding method, in which
15390-439: The costs of increased foraging competition." Landa (1998) argues that the cumulative advantages of shoaling, as elaborated below, are strong selective inducements for fish to join shoals. Parrish et al. (2002) argue similarly that schooling is a classic example of emergence , where there are properties that are possessed by the school but not by the individual fish. Emergent properties give an evolutionary advantage to members of
15561-410: The defensive shoaling and schooling manoeuvres of forage fish. The sailfish raises its sail to make it appear much larger so it can herd a school of fish or squid. Swordfish charge at high speed through forage fish schools, slashing with their swords to kill or stun prey. They then turn and return to consume their "catch". Thresher sharks use their long tails to stun shoaling fishes. Before striking,
15732-435: The detection component of the theory, it was suggested that potential prey might benefit by living together since a predator is less likely to chance upon a single group than a scattered distribution. In the attack component, it was thought that an attacking predator is less likely to eat a particular fish when a greater number of fish are present. In sum, a fish has an advantage if it is in the larger of two groups, assuming that
15903-500: The end of these spiralling runs often carries it into the air. Some predators, such as dolphins, hunt in groups of their own. One technique employed by many dolphin species is herding , where a pod will control a school of fish while individual members take turns ploughing through and feeding on the more tightly packed school (a formation commonly known as a bait ball ). Corralling is a method where fish are chased to shallow water where they are more easily captured. In South Carolina ,
16074-478: The exception of the Nautilidae and the species of octopus belonging to the suborder Cirrina , all known cephalopods have an ink sac, which can be used to expel a cloud of dark ink to confuse predators . This sac is a muscular bag which originated as an extension of the hindgut. It lies beneath the gut and opens into the anus, into which its contents – almost pure melanin – can be squirted; its proximity to
16245-467: The expansion of the mantle at the end of the jet. In some tests, the collagen has been shown to be able to begin raising mantle pressure up to 50ms before muscle activity is initiated. These anatomical differences between squid and octopuses can help explain why squid can be found swimming comparably to fish while octopuses usually rely on other forms of locomotion on the sea floor such as bipedal walking, crawling, and non-jetting swimming. Nautiluses are
16416-416: The fastest marine invertebrates, and they can out-accelerate most fish. The jet is supplemented with fin motion; in the squid, the fins flap each time that a jet is released, amplifying the thrust; they are then extended between jets (presumably to avoid sinking). Oxygenated water is taken into the mantle cavity to the gills and through muscular contraction of this cavity, the spent water is expelled through
16587-440: The fish leap into the air. As the fish leap, the driver dolphin moves with the barrier dolphins and catches the fish in the air. This type of cooperative role specialization seems to be more common in marine animals than in terrestrial animals , perhaps because the oceans have more variability in prey diversity, biomass , and predator mobility. During the sardine run , as many as 18,000 dolphins, behaving like sheepdogs, herd
16758-421: The foot is adapted for burrowing into the sediment; in cephalopods it is used for jet propulsion, and the tentacles and arms are derived from the foot. Most molluscs' circulatory systems are mainly open , except for cephalopods , whose circulatory systems are closed . Although molluscs are coelomates , their coeloms are reduced to fairly small spaces enclosing the heart and gonads. The main body cavity
16929-422: The foot, are below the esophagus and their commissure and connectives to the cerebral and pleural ganglia surround the esophagus in a circumesophageal nerve ring or nerve collar . The acephalic molluscs (i.e., bivalves) also have this ring but it is less obvious and less important. The bivalves have only three pairs of ganglia— cerebral, pedal, and visceral— with the visceral as the largest and most important of
17100-413: The form of jetting. The composition of these mantles differs between the two families, however. In octopuses, the mantle is made up of three muscle types: longitudinal, radial, and circular. The longitudinal muscles run parallel to the length of the octopus and they are used in order to keep the mantle the same length throughout the jetting process. Given that they are muscles, it can be noted that this means
17271-555: The great marine predators. These sometimes immense gatherings fuel the ocean food web . Most forage fish are pelagic fish , which means they form their schools in open water, and not on or near the bottom ( demersal fish ). Forage fish are short-lived, and go mostly unnoticed by humans. The predators are keenly focused on the shoals, acutely aware of their numbers and whereabouts, and make migrations themselves, often in schools of their own, that can span thousands of miles to connect with, or stay connected with them. Herring are among
17442-401: The group is swimming in the same direction in a coordinated manner, they are schooling . In common usage, the terms are sometimes used rather loosely. About one quarter of fish species shoal all their lives, and about one half shoal for part of their lives. Fish derive many benefits from shoaling behaviour including defence against predators (through better predator detection and by diluting
17613-422: The hemoglobin's just four. But unlike hemoglobin, which are attached in millions on the surface of a single red blood cell, hemocyanin molecules float freely in the bloodstream. Cephalopods exchange gases with the seawater by forcing water through their gills, which are attached to the roof of the organism. Water enters the mantle cavity on the outside of the gills, and the entrance of the mantle cavity closes. When
17784-432: The individual animals to follow three rules: An example of such a simulation is the boids program created by Craig Reynolds in 1986. Another is the self-propelled particle model introduced by Vicsek et al. in 1995 Many current models use variations on these rules. For instance, many models implement these three rules through layered zones around each fish. The shape of these zones will necessarily be affected by
17955-548: The kind observed in cephalopod lineages, is typically the result of the movement of pigment within the chromatophore, changing where different pigments are localized within the cell. This physiological change typically occurs on much shorter timescales compared to morphological change. Cephalopods have a rare form of physiological color change which utilizes neural control of muscles to change the morphology of their chromatophores. This neural control of chromatophores has evolved convergently in both cephalopods and teleosts fishes. With
18126-504: The largest extant invertebrate . There are over 800 extant species of cephalopod, although new species continue to be described. An estimated 11,000 extinct taxa have been described, although the soft-bodied nature of cephalopods means they are not easily fossilised. Cephalopods are found in all the oceans of Earth. None of them can tolerate fresh water , but the brief squid, Lolliguncula brevis , found in Chesapeake Bay ,
18297-503: The larvae develop into juvenile fish. When they are old enough, they aggregate into dense shoals and migrate southwards, returning to the Agulhas banks to restart the cycle. The development of schooling behavior was probably associated with an increased quality of perception, predatory lifestyle and size sorting mechanisms to avoid cannibalism. In filter-feeding ancestors, before vision and the octavolateralis system (OLS) had developed,
18468-475: The living molluscs. In 2009, Chapman estimated the number of described living mollusc species at 85,000. Haszprunar in 2001 estimated about 93,000 named species, which include 23% of all named marine organisms. Molluscs are second only to arthropods in numbers of living animal species — far behind the arthropods' 1,113,000 but well ahead of chordates ' 52,000. About 200,000 living species in total are estimated, and 70,000 fossil species, although
18639-468: The mantle cavity, the gills' cilia may stop beating until the unwelcome intrusions have ceased. Each gill has an incoming blood vessel connected to the hemocoel and an outgoing one to the heart. Molluscs use intracellular digestion . Most molluscs have muscular mouths with radulae , "tongues", bearing many rows of chitinous teeth, which are replaced from the rear as they wear out. The radula primarily functions to scrape bacteria and algae off rocks, and
18810-452: The mantle cavity. There is a trade-off with gill size regarding lifestyle. To achieve fast speeds, gills need to be small – water will be passed through them quickly when energy is needed, compensating for their small size. However, organisms which spend most of their time moving slowly along the bottom do not naturally pass much water through their cavity for locomotion; thus they have larger gills, along with complex systems to ensure that water
18981-413: The mantle contracts, water is forced through the gills, which lie between the mantle cavity and the funnel. The water's expulsion through the funnel can be used to power jet propulsion. If respiration is used concurrently with jet propulsion, large losses in speed or oxygen generation can be expected. The gills, which are much more efficient than those of other mollusks, are attached to the ventral surface of
19152-467: The maximum diameter of the funnel orifice (or, perhaps, the average diameter of the funnel) and the diameter of the mantle cavity. Changes in the size of the orifice are used most at intermediate velocities. The absolute velocity achieved is limited by the cephalopod's requirement to inhale water for expulsion; this intake limits the maximum velocity to eight body-lengths per second, a speed which most cephalopods can attain after two funnel-blows. Water refills
19323-676: The more spectacular schooling fish. They aggregate together in huge numbers. The largest schools are often formed during migrations by merging with smaller schools. "Chains" of schools one hundred kilometres (60 miles) long have been observed of mullet migrating in the Caspian Sea . Radakov estimated herring schools in the North Atlantic can occupy up to 4.8 cubic kilometres (1.2 cubic miles) with fish densities between 0.5 and 1.0 fish/cubic metre ( 3 ⁄ 8 to 3 ⁄ 4 fish per cubic yard), totalling about three billion fish in
19494-422: The most neurologically advanced of all invertebrates —and either the giant squid or the colossal squid is the largest known extant invertebrate species. The gastropods ( snails , slugs and abalone ) are by far the most diverse class and account for 80% of the total classified molluscan species. The four most universal features defining modern molluscs are a soft body composed almost entirely of muscle ,
19665-411: The neighbours of a fish in a school, it is thought that efficiency gains do occur in the wild. More recent experiments with groups of fish swimming in flumes support this, with fish reducing their swimming costs by as much as 20% as compared to when the same fish are swimming in isolation. Landa (1998) argued that the leader of a school constantly changes, because while being in the body of a school gives
19836-403: The non threatening herbivorous parrotfish to approach unaware prey. The octopus Thaumoctopus mimicus is known to mimic a number of different venomous organisms it cohabitates with to deter predators. While background matching, a cephalopod changes its appearance to resemble its surroundings, hiding from its predators or concealing itself from prey. The ability to both mimic other organisms and match
20007-981: The number of classes of molluscs; for example, the table below shows seven living classes, and two extinct ones. Although they are unlikely to form a clade, some older works combine the Caudofoveata and Solenogasters into one class, the Aplacophora . Two of the commonly recognized "classes" are known only from fossils. Brachiopods Bivalves Monoplacophorans ("limpet-like", "living fossils") Scaphopods (tusk shells) Gastropods ( snails , slugs , limpets , sea hares ) Cephalopods ( nautiloids , ammonites , octopus , squid , etc.) Aplacophorans (spicule-covered, worm-like) Polyplacophorans (chitons) Wiwaxia Halkieria † Orthrozanclus † Odontogriphus Shoaling and schooling In biology, any group of fish that stay together for social reasons are shoaling , and if
20178-427: The number of arms expressed. Mollusca Mollusca is a phylum of protostomic invertebrate animals , whose members are known as molluscs or mollusks ( / ˈ m ɒ l ə s k s / ). Around 76,000 extant species of molluscs are recognized, making it the second-largest animal phylum after Arthropoda . The number of additional fossil species is estimated between 60,000 and 100,000, and
20349-524: The oceans, from the seashores to the abyssal zone , but some form a significant part of the freshwater fauna and the terrestrial ecosystems . Molluscs are extremely diverse in tropical and temperate regions, but can be found at all latitudes . About 80% of all known mollusc species are gastropods. Cephalopoda such as squid , cuttlefish , and octopuses are among the most neurologically advanced of all invertebrates. The giant squid , which until recently had not been observed alive in its adult form,
20520-413: The octopus Callistoctopus macropus is threatened, it will turn a bright red brown color speckled with white dots as a high contrast display to startle predators. Conspecifically, color change is used for both mating displays and social communication. Cuttlefish have intricate mating displays from males to females. There is also male to male signaling that occurs during competition over mates, all of which are
20691-443: The octopus genus Argonauta secrete a specialized paper-thin egg case in which they reside, and this is popularly regarded as a "shell", although it is not attached to the body of the animal and has a separate evolutionary origin. The largest group of shelled cephalopods, the ammonites , are extinct, but their shells are very common as fossils . The deposition of carbonate, leading to a mineralized shell, appears to be related to
20862-444: The octopus must actively flex the longitudinal muscles during jetting in order to keep the mantle at a constant length. The radial muscles run perpendicular to the longitudinal muscles and are used to thicken and thin the wall of the mantle. Finally, the circular muscles are used as the main activators in jetting. They are muscle bands that surround the mantle and expand/contract the cavity. All three muscle types work in unison to produce
21033-493: The only extant cephalopods with a true external shell. However, all molluscan shells are formed from the ectoderm (outer layer of the embryo); in cuttlefish ( Sepia spp.), for example, an invagination of the ectoderm forms during the embryonic period, resulting in a shell ( cuttlebone ) that is internal in the adult. The same is true of the chitinous gladius of squid and octopuses. Cirrate octopods have arch-shaped cartilaginous fin supports , which are sometimes referred to as
21204-433: The other developed later, or it evolved to regulate trade offs within both. Color change is widespread in ectotherms including anoles, frogs, mollusks, many fish, insects, and spiders. The mechanism behind this color change can be either morphological or physiological. Morphological change is the result of a change in the density of pigment containing cells and tends to change over longer periods of time. Physiological change,
21375-439: The other muscle fibers in the mantle. These collagen fibers act as elastics and are sometimes named "collagen springs". As the name implies, these fibers act as springs. When the radial and circular muscles in the mantle contract, they reach a point where the contraction is no longer efficient to the forward motion of the creature. In such cases, the excess contraction is stored in the collagen which then efficiently begins or aids in
21546-411: The part of the mantle cavity, is swept by the outgoing "lane" of the current created by the gills. Carnivorous molluscs usually have simpler digestive systems. As the head has largely disappeared in bivalves, the mouth has been equipped with labial palps (two on each side of the mouth) to collect the detritus from its mucus. The cephalic molluscs have two pairs of main nerve cords organized around
21717-433: The pen-and-ink fish. Cephalopods are the only molluscs with a closed circulatory system. Coleoids have two gill hearts (also known as branchial hearts ) that move blood through the capillaries of the gills . A single systemic heart then pumps the oxygenated blood through the rest of the body. Like most molluscs, cephalopods use hemocyanin , a copper-containing protein, rather than hemoglobin , to transport oxygen. As
21888-403: The photo below, herring ram feed on a school of copepods . They swim with their mouth wide open and their opercula fully expanded. The fish swim in a grid where the distance between them is the same as the jump length of their prey, as indicated in the animation above right. In the animation, juvenile herring hunt the copepods in this synchronised way. The copepods sense with their antennae
22059-408: The predator's visual channel. Milinski and Heller's findings have been corroborated both in experiment and computer simulations. "Shoaling fish are the same size and silvery, so it is difficult for a visually oriented predator to pick an individual out of a mass of twisting, flashing fish and then have enough time to grab its prey before it disappears into the shoal." Schooling behaviour confuses
22230-407: The pressure-wave of an approaching herring and react with a fast escape jump. The length of the jump is fairly constant. The fish align themselves in a grid with this characteristic jump length. A copepod can dart about 80 times before it tires. After a jump, it takes it 60 milliseconds to spread its antennae again, and this time delay becomes its undoing, as the almost endless stream of herrings allows
22401-576: The primitive molluscan foot. Fishers sometimes call cephalopods " inkfish ", referring to their common ability to squirt ink . The study of cephalopods is a branch of malacology known as teuthology . Cephalopods became dominant during the Ordovician period, represented by primitive nautiloids . The class now contains two, only distantly related, extant subclasses: Coleoidea , which includes octopuses , squid , and cuttlefish ; and Nautiloidea , represented by Nautilus and Allonautilus . In
22572-542: The probability of detection and attack does not increase disproportionately with the size of the group. Schooling forage fish are subject to constant attacks by predators. An example is the attacks that take place during the African sardine run . The African sardine run is a spectacular migration by millions of silvery sardines along the southern coastline of Africa. In terms of biomass, the sardine run could rival East Africa's great wildebeest migration . Sardines have
22743-407: The product of chromatophore coloration displays. There are two hypotheses about the evolution of color change in cephalopods. One hypothesis is that the ability to change color may have evolved for social, sexual, and signaling functions. Another explanation is that it first evolved because of selective pressures encouraging predator avoidance and stealth hunting. For color change to have evolved as
22914-548: The proportion of undescribed species is very high. Many taxa remain poorly studied. Molluscs are the largest marine phylum, comprising about 23% of all the named marine organisms . They are highly diverse, not just in size and anatomical structure, but also in behaviour and habitat , as numerous groups are freshwater and even terrestrial species. The phylum is typically divided into 7 or 8 taxonomic classes , of which two are entirely extinct . Cephalopod molluscs, such as squid , cuttlefish , and octopuses , are among
23085-440: The release of egg and sperm, in the nautilus a string of tissue called the siphuncle goes through all the chambers, and the eight plates that make up the shell of chitons are penetrated with living tissue with nerves and sensory structures. The body of a mollusc has a ventral muscular foot, which is adapted to different purposes (locomotion, grasping the substratum, burrowing or feeding) in different classes. The foot carries
23256-439: The result of natural selection different parameters would have to be met. For one, you would need some phenotypic diversity in body patterning among the population. The species would also need to cohabitate with predators which rely on vision for prey identification. These predators should have a high range of visual sensitivity, detecting not just motion or contrast but also colors. The habitats they occupy would also need to display
23427-424: The result of social selection the environment of cephalopods' ancestors would have to fit a number of criteria. One, there would need to be some kind of mating ritual that involved signaling. Two, they would have to experience demonstrably high levels of sexual selection. And three, the ancestor would need to communicate using sexual signals that are visible to a conspecific receiver. For color change to have evolved as
23598-494: The risk of predation would have been limited and mainly due to invertebrate predators. Hence, at that time, safety in numbers was probably not a major incentive for gathering in shoals or schools. The development of vision and the OLS would have permitted detection of potential prey. This could have led to an increased potential for cannibalism within the shoal. On the other hand, increased quality of perception would give small individuals
23769-405: The same as the flounders as well as move with the same speed and movements. Females of two species, Ocythoe tuberculata and Haliphron atlanticus , have evolved a true swim bladder . Two of the categories of cephalopods, octopus and squid, are vastly different in their movements despite being of the same class. Octopuses are generally not seen as active swimmers; they are often found scavenging
23940-543: The same depth. As such, the cost of transport of many squids are quite high. That being said, squid and other cephalopod that dwell in deep waters tend to be more neutrally buoyant which removes the need to regulate depth and increases their locomotory efficiency. The Macrotritopus defilippi , or the sand-dwelling octopus, was seen mimicking both the coloration and the swimming movements of the sand-dwelling flounder Bothus lunatus to avoid predators. The octopuses were able to flatten their bodies and put their arms back to appear
24111-406: The same speed and in the same direction, then the fish may be said to be schooling . Schooling fish are usually of the same species and the same age/size. Fish schools move with the individual members precisely spaced from each other. The schools undertake complicated manoeuvres, as though the schools have minds of their own. The intricacies of schooling are far from fully understood, especially
24282-446: The sandy sea floor. The color change of chromatophores works in concert with papillae, epithelial tissue which grows and deforms through hydrostatic motion to change skin texture. Chromatophores are able to perform two types of camouflage, mimicry and color matching. Mimicry is when an organism changes its appearance to appear like a different organism. The squid Sepioteuthis sepioide has been documented changing its appearance to appear as
24453-490: The sardines into bait balls, or corral them in shallow water. Once the bait balls are rounded up, the dolphins and other predators take turns ploughing through them, gorging on the fish as they sweep through. Seabirds also attack them from above, flocks of gannets , cormorants , terns and gulls . Some of these seabirds plummet from heights of 30 metres (100 feet), plunging through the water leaving vapour-like trails, similar to that of fighter planes. Gannets plunge into
24624-450: The school which non members do not receive. Support for the social and genetic function of aggregations, especially those formed by fish, can be seen in several aspects of their behaviour. For instance, experiments have shown that individual fish removed from a school will have a higher respiratory rate than those found in the school. This effect has been attributed to stress, and the effect of being with conspecifics therefore appears to be
24795-417: The school. Several anti-predator functions of fish schools have been proposed. One potential method by which fish schools might thwart predators is the "predator confusion effect" proposed and demonstrated by Milinski and Heller (1978). This theory is based on the idea that it becomes difficult for predators to choose individual prey from groups because the many moving targets create a sensory overload of
24966-447: The sea floor instead of swimming long distances through the water. Squids, on the other hand, can be found to travel vast distances, with some moving as much as 2000 km in 2.5 months at an average pace of 0.9 body lengths per second. There is a major reason for the difference in movement type and efficiency: anatomy. Both octopuses and squids have mantles (referenced above) which function towards respiration and locomotion in
25137-487: The seabed. Squids and cuttlefish can move short distances in any direction by rippling of a flap of muscle around the mantle. While most cephalopods float (i.e. are neutrally buoyant or nearly so; in fact most cephalopods are about 2–3% denser than seawater), they achieve this in different ways. Some, such as Nautilus , allow gas to diffuse into the gap between the mantle and the shell; others allow purer water to ooze from their kidneys, forcing out denser salt water from
25308-613: The seafloor and metamorphoses into the adult form. While metamorphosis is the usual state in molluscs, the cephalopods differ in exhibiting direct development: the hatchling is a 'miniaturized' form of the adult. The development of molluscs is of particular interest in the field of ocean acidification as environmental stress is recognized to affect the settlement, metamorphosis, and survival of larvae. Most molluscs are herbivorous, grazing on algae or filter feeders. For those grazing, two feeding strategies are predominant. Some feed on microscopic, filamentous algae, often using their radula as
25479-432: The seawater. While most cephalopods can move by jet propulsion, this is a very energy-consuming way to travel compared to the tail propulsion used by fish. The efficiency of a propeller -driven waterjet (i.e. Froude efficiency ) is greater than a rocket . The relative efficiency of jet propulsion decreases further as animal size increases; paralarvae are far more efficient than juvenile and adult individuals. Since
25650-634: The shape of the school, without collisions. It is as if their motions are choreographed, though they are not. There must be very fast response systems to allow the fish to do this. Young fish practice schooling techniques in pairs, and then in larger groups as their techniques and senses mature. The schooling behaviour develops instinctively and is not learned from older fish. To school the way they do, fish require sensory systems which can respond with great speed to small changes in their position relative to their neighbour. Most schools lose their schooling abilities after dark, and just shoal. This indicates that vision
25821-430: The sharks compact schools of prey by swimming around them and splashing the water with their tails, often in pairs or small groups. Threshers swim in circles to drive schooling prey into a compact mass, before striking them sharply with the upper lobe of its tail to stun them. Spinner sharks charge vertically through the school, spinning on their axis with their mouths open and snapping all around. The shark's momentum at
25992-619: The shoal members, will be better than that taken by an individual fish. Forage fish often make great migrations between their spawning, feeding and nursery grounds. Schools of a particular stock usually travel in a triangle between these grounds. For example, one stock of herrings have their spawning ground in southern Norway , their feeding ground in Iceland , and their nursery ground in northern Norway. Wide triangular journeys such as these may be important because forage fish, when feeding, cannot distinguish their own offspring. Capelin are
26163-480: The shoals, creating a feeding frenzy along the coastline. When threatened, sardines (and other forage fish) instinctively group together and create massive bait balls . Bait balls can be up to 20 metres (66 ft) in diameter. They are short lived, seldom lasting longer than 20 minutes. The fish eggs, left behind at the Agulhas Banks, drift north west with the current into waters off the west coast, where
26334-415: The smaller particles, mainly minerals, to the prostyle so eventually they are excreted, while the larger ones, mainly food, are sent to the stomach's cecum (a pouch with no other exit) to be digested. The sorting process is by no means perfect. Periodically, circular muscles at the hindgut's entrance pinch off and excrete a piece of the prostyle, preventing the prostyle from growing too large. The anus, in
26505-436: The species and for warning ) or active camouflage , as their chromatophores are expanded or contracted. Although color changes appear to rely primarily on vision input, there is evidence that skin cells, specifically chromatophores , can detect light and adjust to light conditions independently of the eyes. The octopus changes skin color and texture during quiet and active sleep cycles. Cephalopods can use chromatophores like
26676-412: The stomach and projecting slightly into the hindgut is the prostyle, a backward-pointing cone of feces and mucus, which is rotated by further cilia so it acts as a bobbin, winding the mucus string onto itself. Before the mucus string reaches the prostyle, the acidity of the stomach makes the mucus less sticky and frees particles from it. The particles are sorted by yet another group of cilia, which send
26847-435: The subject of molluscan anatomy by describing what is called an archi-mollusc , hypothetical generalized mollusc , or hypothetical ancestral mollusc ( HAM ) to illustrate the most common features found within the phylum. The depiction is visually rather similar to modern monoplacophorans . The generalized mollusc is an unsegmented, bilaterally symmetrical animal and has a single, " limpet -like" shell on top. The shell
27018-418: The swimming and feeding energetics. Many hypotheses to explain the function of schooling have been suggested, such as better orientation, synchronized hunting, predator confusion and reduced risk of being found. Schooling also has disadvantages, such as excretion buildup in the breathing media and oxygen and food depletion. The way the fish array in the school probably gives energy saving advantages, though this
27189-414: The three dimensional structure of real world fish shoals because of the large number of fish involved. Techniques include the use of recent advances in fisheries acoustics . Parameters defining a fish shoal include: [REDACTED] Boids simulation – needs Java The observational approach is complemented by the mathematical modelling of schools. The most common mathematical models of schools instruct
27360-499: The three functioning as the principal center of "thinking". Some such as the scallops have eyes around the edges of their shells which connect to a pair of looped nerves and which provide the ability to distinguish between light and shadow. The simplest molluscan reproductive system relies on external fertilization , but with more complex variations. All produce eggs, from which may emerge trochophore larvae, more complex veliger larvae, or miniature adults. Two gonads sit next to
27531-423: The total number of mollusc species ever to have existed, whether or not preserved, must be many times greater than the number alive today. Molluscs have more varied forms than any other animal phylum . They include snails , slugs and other gastropods ; clams and other bivalves ; squids and other cephalopods ; and other lesser-known but similarly distinctive subgroups. The majority of species still live in
27702-402: The urine and dumps additional waste products into it, and then ejects it via tubes that discharge into the mantle cavity. Exceptions to the above are the molluscs Planorbidae or ram's horn snails, which are air-breathing snails that use iron-based hemoglobin instead of the copper-based hemocyanin to carry oxygen through their blood. Most molluscs have only one pair of gills, or even only
27873-520: The usual fashion, but its diet includes protists such as the xenophyophore Stannophyllum . Sacoglossan sea-slugs suck the sap from algae, using their one-row radula to pierce the cell walls, whereas dorid nudibranchs and some Vetigastropoda feed on sponges and others feed on hydroids . (An extensive list of molluscs with unusual feeding habits is available in the appendix of GRAHAM, A. (1955). "Molluscan diets" . Journal of Molluscan Studies . 31 (3–4): 144. .) Opinions vary about
28044-477: The water at up to 100 kilometres per hour (60 mph). They have air sacs under their skin in their face and chest which act like bubble-wrap , cushioning the impact with the water. Subsets of bottlenose dolphin populations in Mauritania are known to engage in interspecific cooperative fishing with human fishermen. The dolphins drive a school of fish towards the shore where humans await with their nets. In
28215-417: The water over their gills. Most bivalves are filter feeders, which can be measured through clearance rates. Research has demonstrated that environmental stress can affect the feeding of bivalves by altering the energy budget of organisms. Cephalopods are primarily predatory, and the radula takes a secondary role to the jaws and tentacles in food acquisition. The monoplacophoran Neopilina uses its radula in
28386-413: The water. It uses receptors called neuromasts , each of which is composed of a group of hair cells. The hairs are surrounded by a protruding jelly-like cupula , typically 0.1 to 0.2 mm long. The hair cells in the lateral line are similar to the hair cells inside the vertebrate inner ear, indicating that the lateral line and the inner ear share a common origin. It is difficult to observe and describe
28557-572: The way that bicyclists may draft one another in a peloton . Geese flying in a Vee formation are also thought to save energy by flying in the updraft of the wingtip vortex generated by the previous animal in the formation. Increased efficiencies in swimming in groups have been proposed for schools of fish and Antarctic krill . It would seem reasonable to think that the regular spacing and size uniformity of fish in schools would result in hydrodynamic efficiencies. While early laboratory-based experiments failed to detect hydrodynamic benefits created by
28728-517: The whale accelerates from below a bait ball to a high velocity and then opens its mouth to a large gape angle. This generates the water pressure required to expand its mouth and engulf and filter a huge amount of water and fish. Lunge feeding by the huge rorquals is said to be the largest biomechanical event on Earth. Fish schools swim in disciplined phalanxes, with some species, such as herrings, able to stream up and down at impressive speeds, twisting this way and that, and making startling changes in
28899-437: Was at the rear in the earliest molluscs, but its position now varies from group to group. The anus , a pair of osphradia (chemical sensors) in the incoming "lane", the hindmost pair of gills and the exit openings of the nephridia (kidneys) known as "Organs of bojanus" and gonads (reproductive organs) are in the mantle cavity. The whole soft body of bivalves lies within an enlarged mantle cavity. The mantle edge secretes
29070-441: Was demonstrated by Pitcher and others in their study of foraging behaviour in shoaling cyprinids . In this study, the time it took for groups of minnows and goldfish to find a patch of food was quantified. The number of fishes in the groups was varied, and a statistically significant decrease in the amount of time necessary for larger groups to find food was established. Further support for an enhanced foraging capability of schools
29241-401: Was published indicating that cephalopod chromatophores are photosensitive; reverse transcription polymerase chain reactions (RT-PCR) revealed transcripts encoding rhodopsin and retinochrome within the retinas and skin of the longfin inshore squid ( Doryteuthis pealeii ), and the common cuttlefish ( Sepia officinalis ) and broadclub cuttlefish ( Sepia latimanus ). The authors claim this
#769230