Fish intelligence is "the resultant of the process of acquiring, storing in memory, retrieving, combining, comparing, and using in new contexts information and conceptual skills" as it applies to
fish
A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...
. Due to a common perception amongst researchers that Teleost fish are "primitive" compared to
mammal
A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s and
bird
Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s, there has been much less research into fish cognition than into those types of animals, and much remains unknown about fish cognition, though evidence of complex navigational skills such as cognitive maps is increasing.
Compared to similarly sized fish, mammals and birds typically have brain sizes fifteen times larger, though some species of fish such as elephantnose fish have very large brain-to-body ratios. However, fish still display intelligence that cannot be explained through Pavlovian and operant conditioning, such as reversal learning, novel obstacle avoidance, and passing simultaneous two-choice tasks. Some fish also match mammals and birds in the executive functioning capability of
inhibitory
An inhibitory postsynaptic potential (IPSP) is a kind of synaptic potential that makes a Chemical synapse, postsynaptic neuron less likely to generate an action potential.Purves et al. Neuroscience. 4th ed. Sunderland (MA): Sinauer Associates, Inc ...
motor control
Motor control is the regulation of movements in organisms that possess a nervous system. Motor control includes conscious voluntary movements, subconscious muscle memory and involuntary reflexes, as well as instinctual taxes.
To control ...
. Australian biologist Culum Brown has argued that fish may give the appearance of being less intelligent than they are due to differences between aquatic and terrestrial environments.
Fish hold records for the relative brain weights of vertebrates. Most vertebrate species have similar brain-to-body mass ratios. The deep sea bathypelagic bony-eared assfish has the smallest ratio of all known vertebrates.Fine ML, Horn MH and Cox B (1987 "''Acanthonus armatus'', a Deep-Sea Teleost Fish with a Minute Brain and Large Ears" ''Proceedings of the Royal Society B'', 230(1259)257-265. At the other extreme, the electrogenic elephantnose fish, an African freshwater fish, has one of the largest brain-to-body weight ratios of all known vertebrates (slightly higher than humans) and the highest brain-to-body oxygen consumption ratio of all known vertebrates (three times that for humans).
Brain
Fish typically have quite small brains relative to body size compared with other vertebrates, typically one-fifteenth the brain mass of a similarly sized bird or mammal. However, some fish have relatively large brains, most notably mormyrids and
shark
Sharks are a group of elasmobranch cartilaginous fish characterized by a ribless endoskeleton, dermal denticles, five to seven gill slits on each side, and pectoral fins that are not fused to the head. Modern sharks are classified within the ...
s, which have brains about as massive relative to body weight as birds and
marsupial
Marsupials are a diverse group of mammals belonging to the infraclass Marsupialia. They are natively found in Australasia, Wallacea, and the Americas. One of marsupials' unique features is their reproductive strategy: the young are born in a r ...
s.
The cerebellum of
cartilaginous
Cartilage is a resilient and smooth type of connective tissue. Semi-transparent and non-porous, it is usually covered by a tough and fibrous membrane called perichondrium. In tetrapods, it covers and protects the ends of long bones at the joints ...
and
bony fish
Osteichthyes ( ; ), also known as osteichthyans or commonly referred to as the bony fish, is a Biodiversity, diverse clade of vertebrate animals that have endoskeletons primarily composed of bone tissue. They can be contrasted with the Chondricht ...
es is large and complex. In at least one important respect, it differs in internal structure from the mammalian cerebellum: The fish cerebellum does not contain discrete
deep cerebellar nuclei
There are four paired deep cerebellar nuclei embedded in the white matter centre of the cerebellum. The nuclei are the fastigial, globose, emboliform, and dentate nuclei.
In lower mammals the emboliform nucleus appears to be continuous with t ...
. Instead, the primary targets of Purkinje cells are a distinct type of cell distributed across the cerebellar cortex, a type not seen in mammals. The circuits in the
cerebellum
The cerebellum (: cerebella or cerebellums; Latin for 'little brain') is a major feature of the hindbrain of all vertebrates. Although usually smaller than the cerebrum, in some animals such as the mormyrid fishes it may be as large as it or eve ...
are similar across all
class
Class, Classes, or The Class may refer to:
Common uses not otherwise categorized
* Class (biology), a taxonomic rank
* Class (knowledge representation), a collection of individuals or objects
* Class (philosophy), an analytical concept used d ...
es of
vertebrate
Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain.
The vertebrates make up the subphylum Vertebra ...
s, including fish, reptiles, birds, and mammals. There is also an analogous brain structure in
cephalopod
A cephalopod is any member of the molluscan Taxonomic rank, class Cephalopoda (Greek language, Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral symm ...
s with well-developed brains, such as
octopus
An octopus (: octopuses or octopodes) is a soft-bodied, eight-limbed mollusc of the order Octopoda (, ). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttlefish, and nautiloids. Like oth ...
es. This has been taken as evidence that the cerebellum performs functions important to all animal
species
A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
with a brain.
In mormyrid fish (a family of weakly electrosensitive freshwater fish), the cerebellum is considerably larger than the rest of the brain put together. The largest part of it is a special structure called the ''valvula'', which has an unusually regular architecture and receives much of its input from the electrosensory system.
Memory
Individual
carp
The term carp (: carp) is a generic common name for numerous species of freshwater fish from the family (biology), family Cyprinidae, a very large clade of ray-finned fish mostly native to Eurasia. While carp are prized game fish, quarries and a ...
captured by anglers have been shown to become less catchable thereafter. This suggests that fish use their memory of negative experiences to associate capture with stress and therefore become less easy to catch. This type of
associative learning
Learning is the process of acquiring new understanding, knowledge, behaviors, skills, values, attitudes, and preferences. The ability to learn is possessed by humans, non-human animals, and some machines; there is also evidence for some kin ...
has also been shown in
paradise fish
The fork tailed paradisefish (often just called paradise fish, paradise-fish, paradisefish), or paradise gourami (''Macropodus opercularis'') is a species of gourami found in most types of fresh water in East Asia, ranging from the China to nort ...
(''Macropodus opercularis'') which avoid places where they have experienced a single attack by a predator and continue to avoid for many months.
A number of studies have shown that fish can retain information for months or years. Anecdotally,
channel catfish
The channel catfish (''Ictalurus punctatus''), known informally as the "channel cat", is a species of catfish native to North America. They are North America's most abundant catfish species, and the official state fish of Kansas, Missouri, Nebra ...
(''Ictalurus punctatus'') can remember the human voice call announcing food five years after last hearing that call.Reebs, S.G. (2008 Long-term memory in fishes Retrieved 9 July 2014.
Goldfish
The goldfish (''Carassius auratus'') is a freshwater fish in the family Cyprinidae of the order Cypriniformes. It is commonly kept as a pet in indoor aquariums, and is one of the most popular aquarium fish. Goldfish released into the w ...
remember the colour of a tube dispensing food one year after the last tube presentation.
Sockeye salmon
The sockeye salmon (''Oncorhynchus nerka''), also called red salmon, kokanee salmon, blueback salmon, or simply sockeye, is an anadromous species of salmon found in the Northern Pacific Ocean and rivers discharging into it. This species is a ...
still react to a light signal that precedes food arrival up to eight months since the last
reinforcement
In Behaviorism, behavioral psychology, reinforcement refers to consequences that increase the likelihood of an organism's future behavior, typically in the presence of a particular ''Antecedent (behavioral psychology), antecedent stimulus''. Fo ...
. Some
common rudd
The common rudd (''Scardinius erythrophthalmus'') is a benthos, bentho-pelagic freshwater Actinopterygii, ray-finned fish belonging to the Family (biology), family Leuciscidae. This species is widely spread in Europe and central Asia, around the ...
and
European chub
''Squalius cephalus'', the common chub, European chub or simply chub, is a species of freshwater ray-finned fish belonging to the family Leuciscidae, the daces, Eurasian minnows and related fishes. This species is found in Europe where it freq ...
could remember the person who trained them to feed from the hand, even after a 6-month break. Crimson-spotted rainbowfish can learn how to escape from a trawl by swimming through a small hole in the center and they remember this technique 11 months later.
Rainbow trout
The rainbow trout (''Oncorhynchus mykiss'') is a species of trout native to cold-water tributary, tributaries of the Pacific Ocean in North America and Asia. The steelhead (sometimes called steelhead trout) is an Fish migration#Classification, ...
can be trained to press a bar to get food, and they remember this three months after last seeing the bar. Red Sea clownfish can recognize their mate 30 days after it was experimentally removed from the home anemone.
Several fish species are capable of learning complex spatial relationships and forming cognitive maps. They can orient themselves using multiple landmarks or symbols and they are able to integrate experiences which enable them to generate appropriate avoidance responses. In 2020, a neurological study using wireless neural recording systems measured the neurological activity of
goldfish
The goldfish (''Carassius auratus'') is a freshwater fish in the family Cyprinidae of the order Cypriniformes. It is commonly kept as a pet in indoor aquariums, and is one of the most popular aquarium fish. Goldfish released into the w ...
and found strong evidence supporting the idea that fish form navigational cognitive maps.
Tool-like behavior
Tool use
Tool use by non-humans is a phenomenon in which a non-human animal uses any kind of tool in order to achieve a goal such as acquiring food and water, grooming, combat, defence, communication, recreation or construction. Originally thought to b ...
is sometimes considered as an indication of intelligence in animals. There are few examples of tool use in fishes, perhaps because they have only their mouth in which to hold objects.
Several species of
wrasse
The wrasses are a family, Labridae, of marine ray-finned fish, many of which are brightly colored. The family is large and diverse, with over 600 species in 81 genera, which are divided into nine subgroups or tribes.
They are typically small, ...
hold
bivalves
Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed by a calcified exoskeleton consis ...
(scallops and clams) or
sea urchin
Sea urchins or urchins () are echinoderms in the class (biology), class Echinoidea. About 950 species live on the seabed, inhabiting all oceans and depth zones from the intertidal zone to deep seas of . They typically have a globular body cove ...
s in their mouth and smash them against the surface of a rock (an "anvil") to break them up. This behaviour in an orange-dotted tuskfish (''Choerodon anchorago'') has been filmed; the fish fans sand to unearth the bivalve, takes it into its mouth, swims several metres to a rock which it uses as an anvil by smashing the mollusc apart with sideward thrashes of the head.
Archerfish
The archerfish (also known as spinner fish or archer fish) or Toxotidae are a monotypic family (although some include a second genus) of perciform tropical fish known for their unique predation technique of "shooting down" land-based insects a ...
(
family
Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Toxotidae) squirt jets of water at insects on plants above the surface to knock them into the water. They can adjust the size of the squirts to the size of the insect prey. They can even learn to shoot at moving targets.
Whitetail damselfish clean the rock face where they intend to lay eggs by sucking up and blowing sand grains onto the surface.
Triggerfish
Triggerfish are about 40 species of often brightly colored marine ray-finned fishes belonging to the family Balistidae. Often marked by lines and spots, they inhabit tropical and subtropical oceans throughout the world, with the greatest speci ...
blow water at
sea urchin
Sea urchins or urchins () are echinoderms in the class (biology), class Echinoidea. About 950 species live on the seabed, inhabiting all oceans and depth zones from the intertidal zone to deep seas of . They typically have a globular body cove ...
s to turn them over, thereby exposing their more vulnerable underside. River stingrays create water currents with their fins to suck food out of a PVC pipe. Banded acaras (''Bujurquina vittata'') lay their eggs on a loose leaf and carry the leaf away when a predator approaches.
In one laboratory study,
Atlantic cod
The Atlantic cod (: cod; ''Gadus morhua'') is a fish of the family Gadidae, widely consumed by humans. It is also commercially known as '' cod'' or ''codling''.operant feeding machine learned to pull a string to get food. The researchers had also tagged the fish by threading a bead in front of their dorsal fin. Some fish snagged the string with their bead, resulting in food delivery. These fish eventually learned to swim in a particular way to repeatedly make the bead snag the string and get food. Because the fish used an object external to their body in a goal-oriented way, this satisfies some definitions of tool use - though in a more rudimentary fashion, a human blowing air wouldn't qualify as tool usage just like a fish using water in the same fashion, or even in the case of certain wrasse, the bivalve is not a tool but the prey itself. If the wrasse had used a rock to access the bivalve, then it would fall under that classification. There appear to be very few to no observed behaviors of fish using tools in a fashion similar to humans, but instead altering water flows in ways that benefits their survival.
Construction
As for tool use, construction behaviour may be mostly innate. Yet it can be sophisticated, and the fact that fish can make judicious repairs to their creation suggests intelligence. Construction methods in fishes can be divided into three categories: excavations, pile-ups, and gluing.Reebs, S.G. (2009-2013 Can fish build things? Retrieved 10 July 2014.
Excavations may be simple depressions dug up in the substrate, such as the nests of
bowfin
The ruddy bowfin (''Amia calva'') is a ray-finned fish native to North America. Common names include mudfish, mud pike, dogfish, grindle, grinnel, swamp trout, and choupique. It is regarded as a relict, being one of only two surviving species ...
,
smallmouth bass
The smallmouth bass (''Micropterus dolomieu'') is a species of freshwater fish in the Centrarchidae, sunfish family (biology), family (Centrarchidae) of the order (biology), order Centrarchiformes. It is the type species of its genus ''Micropterus ...
, and
Pacific salmon
''Oncorhynchus'', from Ancient Greek ὄγκος (''ónkos''), meaning "bend", and ῥύγχος (''rhúnkhos''), meaning "snout", is a genus of ray-finned fish in the subfamily Salmoninae of the family Salmonidae, native to coldwater tributarie ...
, but it can also consist of fairly large burrows used for shelter and for nesting. Burrowing species include the mudskippers, the red band-fish (''Cepola rubescens)'' (burrows up to 1 m deep, often with a side branch), the yellowhead jawfish (''Opistognathus aurifrons)'' (chambers up to 22 cm deep, lined with coral fragments to solidify it), the convict blenny (''Pholidichthys leucotaenia)'', whose burrow is a maze of tunnels and chambers thought to be as much as 6 m long, and the Nicaragua cichlid ('' Hypsophrys nicaraguensis)'', who drills a tunnel by spinning inside of it. In the case of the mudskippers, the burrows are shaped like a J and can be as much as 2 m deep. Two species, the giant mudskipper (''Periophthalmodon schlosseri)'' and the walking goby ('' Scartelaos histophorus)'', build a special chamber at the bottom of their burrows into which they carry mouthfuls of air. Once released the air accumulates at the top of the chamber and forms a reserve from which the fish can breathe – like all amphibious fishes, mudskippers are good air breathers. If researchers experimentally extract air from the special chambers, the fish diligently replenish it. The significance of this behaviour stems from the facts that at high tide, when water covers the mudflats, the fish stay in their burrow to avoid predators, and water inside the confined burrow is often poorly oxygenated. At such times these air-breathing fishes can tap into the air reserve of their special chambers.
Mounds are easy to build, but can be quite extensive. In North American streams, the male cutlip minnow ''Exoglossum maxillingua'', long, assembles mounds that are high, in diameter, made up of more than 300 pebbles 13–19 mm in diameter (a quarter to half an inch). The fish carry these pebbles one by one in their mouths, sometimes stealing some from the mounds of other males. The females deposit their eggs on the upstream slope of the mounds, and the males cover these eggs with more pebbles. Males of the hornyhead chub ''Nocomis biguttatus'', long, and of the river chub ''Nocomis micropogon'', long, also build mounds during the reproductive season. They start by clearing a slight depression in the substrate, which they overfill with up to 10,000 pebbles until the mounds are long (in the direction of the water current), wide, and high. Females lay their eggs among those pebbles. The stone accumulation is free of sand and it exposes the eggs to a good water current that supplies oxygen.
Males of many mouthbrooding cichlid species in
Lake Malawi
Lake Malawi, also known as Lake Nyasa in Tanzania and Lago Niassa in Mozambique, () is an African Great Lakes, African Great Lake and the southernmost lake in the East African Rift system, located between Malawi, Mozambique and Tanzania.
It is ...
and
Lake Tanganyika
Lake Tanganyika ( ; ) is an African Great Lakes, African Great Lake. It is the world's List of lakes by volume, second-largest freshwater lake by volume and the List of lakes by depth, second deepest, in both cases after Lake Baikal in Siberia. ...
build sand cones that are flattened or crater-shaped at the top. Some of these mounds can be 3 m in diameter and 40 cm high. The mounds serve to impress females or to allow species recognition during courtship.
Male pufferfish, ''Torquigener'' sp., also build sand mounds to attract females. The mounds, up to 2 m in diameter, are intricate with radiating ridges and valleys.
Several species build up mounds of coral pieces either to protect the entrance to their burrows, as in
tilefish
file:Malacanthus latovittatus.jpg, 250px, Blue blanquillo, ''Malacanthus latovittatus''
Tilefishes are mostly small perciform marine fish comprising the family (biology), family Malacanthidae. They are usually found in sandy areas, especially n ...
es and gobies of the genus Valenciennea, or to protect the patch of sand in which they will bury themselves for the night, as in the Jordan's tuskfish ''Choerodon jordani'' and the rockmover wrasse ''Novaculichthys taeniourus''.
Male sticklebacks are well known for their habit of building an enclosed nest made of pieces of vegetation glued together with secretions from their kidneys. Some of them adorn the entrance of the nest with unusually colored algae or even shiny tinfoil experimentally introduced in the environment.
Foam nests, made up of air bubbles glued together with mucus from the mouth, are also well known in gouramis and armoured catfish.
Social intelligence
Fish can remember the attributes of other individuals, such as their competitive ability or past behavior, and modify their own behavior accordingly. For example, they can remember the identity of individuals to whom they have lost in a fight, and avoid these individuals in the future; or they can recognize territorial neighbors and show less aggression towards them as compared to strangers.Reebs, S.G. (2010 Social intelligence in fishes Retrieved 10 July 2014 They can recognize individuals in whose company they obtained less food in the past and preferentially associate with new partners in the future.
Fish can seem mindful of which individuals have watched them in the past. In an experiment with
Siamese fighting fish
The Siamese fighting fish (''Betta splendens''), commonly known as the betta, is a freshwater fish native to Southeast Asia, namely Cambodia, Laos, Myanmar, Malaysia, Indonesia, Thailand, and Vietnam. It is one of 76 species of the genus '' Bet ...
, two males were made to fight each other while being watched by a female, whom the males could also see. The winner and the loser of the fight were then, separately, given a choice between spending time next to the watching female or to a new female. The winner courted both females equally, but the loser spent more time next to the new female, avoiding the watcher female. In this species, females prefer males they have seen win a fight over males they have seen losing, and it therefore makes sense for a male to prefer a female that has never seen him lose as opposed to a female that has seen him lose.
Social interactions also provides the context for a test of transitive inference, that is figuring out that if A > B and B > C, then A > C. In a study with the cichlid '' Astatotilapia burtoni'', observer fish could watch aggressive interactions between pairs of other individuals. They witnessed individual A beat individual B, then individual B beat individual C, then C beat D, and D beat E. The observer fish were then given a choice of associating with either B or D (both of which they had seen win once and lose once). All eight observer fish tested spent more time next to D. Fish in this species prefer to associate with more subordinate individuals, so the preference for D showed that the observers had worked out that given that B was superior to C, and C to D, then D had to be subordinate to B.
Deception
There are several examples of fish being
deceptive
Deception is the act of convincing of one or many recipients of untrue information. The person creating the deception knows it to be false while the receiver of the information does not. It is often done for personal gain or advantage.
Deceit ...
, suggesting to some researchers that they may possess a
theory of mind
In psychology and philosophy, theory of mind (often abbreviated to ToM) refers to the capacity to understand other individuals by ascribing mental states to them. A theory of mind includes the understanding that others' beliefs, desires, intent ...
. However, most of the observations of deception can be understood as instinctive patterns of behavior that are triggered by specific environmental events, and they do not require a fish to understand the point of view of other individuals.
Distraction display
In the threespine stickleback (''Gasterosteus aculeatus''), males sometimes see their nest full of eggs fall prey to groups of marauding females; some males, when they see a group of females approaching, swim away from their nest and start poking their snout into the substrate, as would a female raiding a nest. This
distraction display
Distraction displays, also known as diversionary displays, or paratrepsis are antipredator adaptation, anti-predator behaviors used to attract the attention of an enemy away from something, typically the nest or young, that is being protected by a ...
commonly fools the females into behaving as if a nest has been discovered there and they rush to that site, leaving the male's real nest alone.
Bowfin
The ruddy bowfin (''Amia calva'') is a ray-finned fish native to North America. Common names include mudfish, mud pike, dogfish, grindle, grinnel, swamp trout, and choupique. It is regarded as a relict, being one of only two surviving species ...
(''Amia calva'') males caring for their free-swimming fry exhibit a related distraction display when a potential fry-predator approaches; they move away and thrash about as if injured, drawing the predator's attention toward himself.
False courtship behaviour
In the Malili Lakes of Sulawesi, Indonesia, one species of
sailfin silverside
The Telmatherinidae, the sail-fin silversides are a family of atheriniform fish, inhabiting fresh and brackish water. All but the species '' Kalyptatherina helodes'' are restricted to the Indonesian island of Sulawesi
Sulawesi ( ), also kno ...
(''Telmatherina sarasinorum'') is an egg predator. They often follow courting pairs of the closely related species ''T. antoniae''. When those pairs lay eggs, ''T. sarasinorum'' darts in and eats the eggs. On four different occasions in the field (out of 136 observation bouts in total), a male ''T. sarasinorum'' who was following a pair of courting ''T. antoniae'' eventually chased off the male ''T. antoniae'' and took his place, courting the heterospecific female. That female released eggs, at which point the male darted to the eggs and ate them.
Death feigning
Death feigning as a way to attract prey is another form of deception. In
Lake Malawi
Lake Malawi, also known as Lake Nyasa in Tanzania and Lago Niassa in Mozambique, () is an African Great Lakes, African Great Lake and the southernmost lake in the East African Rift system, located between Malawi, Mozambique and Tanzania.
It is ...
, the predatory cichlid ''
Nimbochromis livingstonii
''Nimbochromis livingstonii'', Livingston's cichlid or, ''kalingono'' (in Chichewa), is a freshwater mouthbrooding cichlid native to Lake Malawi, an African Rift Lake. It is also found in the upper Shire River and Lake Malombe. They are foun ...
'' have been seen first remaining stationary with their abdomen on or near sand and that then dropping onto their sides. In a variant behaviour, some ''N. livingstonii'' fell through the water column and landed onto their side. The fish then remained immobile for several minutes. Their colour pattern was blotchy and suggested a rotting carcass. Small inquisitive cichlids of other species often came near and they were suddenly attacked by the predator. About a third of the death-feigning performances led to an attack, and about one-sixth of the attacks were successful. Another African cichlid, '' Lamprologus lemairii'', from
Lake Tanganyika
Lake Tanganyika ( ; ) is an African Great Lakes, African Great Lake. It is the world's List of lakes by volume, second-largest freshwater lake by volume and the List of lakes by depth, second deepest, in both cases after Lake Baikal in Siberia. ...
, has been reported to do the same thing. A South American cichlid, the yellowjacket cichlid '' Parachromis friedrichsthalii'', also uses death feigning. They turn over onto their sides at the bottom of the sinkholes they inhabit and remain immobile for as long as 15 minutes, during which they attack the small mollies that come too close to them. The comb grouper ''Mycteroperca acutirostris'' may also be an actor, though in this case the behaviour should be called dying or illness feigning, rather than death feigning, because while lying on its side the fish occasionally undulates its body. In 1999, off the coast of southeastern Brazil, one juvenile comb grouper was observed using this tactic to catch five small prey in 15 minutes.
Cooperation
Cooperative foraging reflects some mental flexibility and planning, and could therefore be interpreted as intelligence. There are a few examples in fishes.
Yellowtail amberjack can form packs of 7-15 individuals that maneuver in U-shaped formations to cut away the tail end of prey shoals ( jack mackerels or Cortez grunts) and herd the downsized shoal next to seawalls where they proceed to capture the prey.
In the coral reefs of the Red Sea, roving coralgrouper that have spotted a small prey fish hiding in a crevice sometimes visit the sleeping hole of a giant moray and shake their head at the moray, and this seems to be an invitation to group hunting as the moray often swims away with the grouper, is led to the crevice where the prey hides, and proceeds to probe that crevice (which is too small to let the grouper in), either catching the prey by itself or flushing it into the open where the grouper grabs it. The closely related coral trout also enrolls the help of moray eels in this way, and they only do so when the prey they seek is hidden in crevices, where only the eel can flush them. They also quickly learn to invite preferentially those individual eels that collaborate most often.
Similarly, zebra lionfish that detect the presence of small prey fishes flare up their fins as an invitation to other zebra lionfish, or even to another species of lionfish (''
Pterois antennata
''Pterois'' is a genus of venomous fish, venomous marine fish, commonly known as the lionfish, native to the Indo-Pacific. It is characterized by conspicuous aposematism, warning coloration with red or black bands and ostentatious dorsal fins tip ...
''), to join them in better cornering the prey and taking turns at striking the prey so that every individual hunter ends up with similar capture rates.
Numeracy
Mosquitofish (''Gambusia holbrooki'') can distinguish between doors marked with either two or three geometric symbols, only one of which allows the fish to rejoin its shoalmates. This can be achieved when the two symbols have the same total surface area, density and brightness as the three symbols. Further studies have shown that this discrimination extends to 4 vs 8, 15 vs 30, 100 vs 200, 7 vs 14, and 8 vs 12 symbols, again controlling for non-numerical factors. Other species similarly tested with some measure of success include
goldfish
The goldfish (''Carassius auratus'') is a freshwater fish in the family Cyprinidae of the order Cypriniformes. It is commonly kept as a pet in indoor aquariums, and is one of the most popular aquarium fish. Goldfish released into the w ...
(discriminating between 2 vs. 3, and 10 vs. 15),
guppy
The Greater Underwater Propulsion Power Program (GUPPY) was initiated by the United States Navy after World War II to improve the submerged speed, maneuverability, and endurance of its submarines. (The "Y" in the acronym was added for pronouncea ...
(3 vs. 4, and 4 vs. 5), and
zebrafish
The zebrafish (''Danio rerio'') is a species of freshwater ray-finned fish belonging to the family Danionidae of the order Cypriniformes. Native to South Asia, it is a popular aquarium fish, frequently sold under the trade name zebra danio (an ...
(2 vs. 3, 3 vs. 4, and 4 vs. 5, but not 5 vs. 6, nor 6 vs. 7).
Many studies have shown that when given a choice, shoaling fish prefer to join the larger of two shoals. It has been argued that several aspects of such choice reflect an ability by fish to distinguish between numerical quantities.
A laboratory study with zebra mbuna and ocellate river stingray has demonstrated that these fish can add and subtract 'one' from 2, 3, or 4. The fish had to learn that if presented with, say, 3 yellow symbols, then in a subsequent choice test between 2 and 4 symbols they had to choose 2 to get a food reward (thus, yellow meant "choose one less"); but if presented with 3 blue symbols, then they had to choose 4 rather than 2 (thus, blue meant "choose one more"). The fish easily learned this task (the success criterion was 70% correct choice). Crucially, if after 3 yellow symbols they were given a choice between 1 and 2, they chose 2; conversely, if after 3 blue symbols the fish were given a choice between 4 or 5 symbols, they chose 4. This latter test showed that the fish had learned the concept of "one less" or "one more", and not simply "less" or "more".
Social learning
Fish can learn how to perform a behavior simply by watching other individuals in action. This is variously called
observational learning
Observational learning is learning that occurs through observing the behavior of others. It is a form of Social learning theory, social learning which takes various forms, based on various processes. In humans, this form of learning seems to not n ...
, cultural transmission, or social learning. For example, fish can learn a particular route after following an experienced leader a few times. One study trained guppies to swim through a hole marked in red while ignoring another one marked in green in order to get food on the other side of a partition; when these experienced fish ("demonstrators") were joined by a naive one (an "observer"), the observer followed the demonstrators through the red hole, and kept the habit once the demonstrators were removed, even when the green hole now allowed food access. In the wild, juvenile French grunt follow traditional migration routes, up to 1 km long, between their daytime resting sites and their nighttime foraging areas on coral reefs; if groups of 10-20 individuals are marked and then transplanted to new populations, they follow the residents along what is for them – the transplants – a new migration route, and if the residents are then removed two days later, the transplanted grunts continue to use the new route, as well as the resting and foraging sites at both ends.
Through cultural transmission, fishes could also learn where good food spots are. Ninespine stickleback, when given a choice between two food patches they have watched for a while, prefer the patch over which more fish have been seen foraging, or over which fish were seen feeding more intensively. Similarly, in a field experiment where Trinidadian guppies were given a choice between two distinctly marked feeders in their home rivers, the subjects chose the feeder where other guppies were already present, and in subsequent tests when both feeders were deserted, the subjects remembered the previously popular feeder and chose it.
Through social learning, fishes might learn not only where to get food, but also what to get and how to get it. Hatchery-raised salmon can be taught to quickly accept novel, live prey items similar to those they will encounter once they will be released in the wild, simply by watching an experienced salmon take such prey. The same is true of young
perch
Perch is a common name for freshwater fish from the genus ''Perca'', which belongs to the family Percidae of the large order Perciformes. The name comes from , meaning the type species of this genus, the European perch (''P. fluviatilis'') ...
. In the laboratory, juvenile European seabass can learn to push a lever in order to obtain food just by watching experienced individuals use the lever.
Fishes can also learn from others the identity of predatory species.
Fathead minnow
Fathead minnow (''Pimephales promelas''), also known as fathead or tuffy, is a species of freshwater Actinopterygii, ray-finned fish belonging to the Family (biology), family Leuciscidae, the shiners, daces and minnows. The natural geographic ra ...
s, for example, can learn the smell of a predatory pike just by being simultaneously exposed to that smell and the sight of experienced minnows reacting with fear, and brook stickleback can learn the visual identity of a predator by watching the fright reaction of experienced fathead minnows. Fish can also learn to recognize the odor of dangerous sites when they are simultaneously exposed to it and to other fish that suddenly show a fright reaction. Hatchery-raised salmon can learn the smell of a predator by being simultaneously exposed to it and to the alarm substance released by injured salmon.
Latent learning
Latent learning is a form of learning that is not immediately expressed in an overt response; it occurs without any obvious reinforcement of the behaviour or associations that are learned. One example in fish comes from research with male three spot gouramis (''Trichopodus trichopterus''). This species quickly form dominance hierarchies. To appease dominants, subordinates adopt a typical body posture angled at 15-60º to the horizontal, all fins folded and pale body colors. Individuals trained to associate a light-stimulus with the imminent arrival of food exhibit this associative learning by approaching the surface where the food is normally dropped immediately the light-stimulus is presented. However, if a subordinate is placed in a tank with a dominant individual and the light-stimulus is presented, the subordinate immediately assumes the submissive posture rather than approaching the surface. The subordinate has predicted that going to the surface to get food would place it in competition with the dominant, and to avoid potential aggression, it immediately attempts to appease the dominant.
Cleaner fish
The bluestreak cleaner wrasse (''Labroides dimidiatus'') performs a service for "client" fishes (belonging to other species) by removing and eating their
ectoparasites
Parasitism is a close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The en ...
. Clients can invite a cleaning session by adopting a typical posture or simply by remaining immobile near a wrasse's cleaning station. They can even form queues while doing so. But cleaning sessions do not always end up well, because wrasses (or wrasse-mimicking parasitic sabre-toothed blennies) may cheat and eat the nutritious body mucus of their clients, rather than just the ectoparasites, something that makes the client jolt and sometimes flee. This system has been the subject of extensive observations which have suggested cognitive abilities on the part of the cleaner wrasses and their clients. For example, clients refrain from soliciting a cleaning session if they have witnessed the cleaning session of the previous client ending badly. Cleaners give the impression of trying to maintain a good reputation, because they cheat less when they see a big audience (a long queue of clients) watching. Cleaners sometimes work as male-female teams, and when the smaller female cheats and bites the client, the larger male chases her off, as if to punish her for having tarnished their reputation.
mirror test
The mirror test—sometimes called the mark test, mirror self-recognition (MSR) test, red spot technique, or rouge test—is a behavioral technique developed in 1970 by American psychologist Gordon Gallup Jr. to determine whether an animal posse ...
. According to the authors, their findings mean that either prior conceptions of animal self-awareness or the validity of the mark test need to be reexamined. Gordon Gallup, the originator of the test, believes that the results are best explained by the fish having instincts to scrape off parasites. In 2022, a larger study was performed with positive results, but the fish only passed when the mark resembled a parasite. The authors then reiterated their conclusion that either self-awareness in animals or the validity of the mark test as a method need to be reconsidered.
Play
Play behaviour is often considered a correlate of intelligence. One possible example in fish is provided by the electrolocating Peters' elephantnose fish (mentioned above as having one of the largest brain-to-body weight ratios of all known vertebrates). One captive individual was observed carrying a small ball of aluminum foil to the outflow tube of the aquarium filter, letting the current push the ball away before chasing after it and repeating the behaviour. The same species has been reported trying to balance tubes vertically at the water surface until they fell, and then try again. Captive white-spotted cichlids have also been seen hitting a floating thermometer hundreds of times to make it wobble and bob.
Food stocking
Food stocking can be viewed potentially as an animal planning for the future. One example of short-term stocking involves climbing perch (''Anabas testudineus''). Individuals were kept singly in aquaria and fed with pellets dropped at the surface. When the pellets were dropped one after the other at 1-s intervals, the fish took them as they reached the surface and stocked them inside the mouth. On average, the fish placed 7 pellets in their mouth before moving away to consume them. When starved for 24-h before the feeding test, they doubled the number of pellets stocked (14 on average); the underside of their heads bulged under the load. The behaviour may be an indication that competition for food is normally severe in this species and that any adaptation to secure food would be beneficial.Binoy, V.V., and Thomas, K.J. 2008. The influence of hunger on food-stocking behaviour of climbing perch ''Anabas testudineus''. Journal of Fish Biology 73: 1053-1057.
See also
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Animal cognition
Animal cognition encompasses the mental capacities of non-human animals, including insect cognition. The study of animal conditioning and learning used in this field was developed from comparative psychology. It has also been strongly influ ...
*
Bird intelligence
The difficulty of defining or measuring intelligence in non-human animals makes the subject difficult to study scientifically in birds. In general, birds have relatively large brains compared to their head size. Furthermore, bird brains have two ...
*
Cephalopod intelligence
Cephalopod intelligence is a measure of the cognitive ability of the cephalopod class of molluscs.
Intelligence is generally defined as the process of acquiring, storing, retrieving, combining, and comparing information and skills. Though these ...
Shoaling and schooling
In biology, any group of fish that stay together for social reasons are shoaling, and if 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. Ab ...
The Sydney Morning Herald
''The Sydney Morning Herald'' (''SMH'') is a daily Tabloid (newspaper format), tabloid newspaper published in Sydney, Australia, and owned by Nine Entertainment. Founded in 1831 as the ''Sydney Herald'', the ''Herald'' is the oldest continuous ...
TimesOnline
''The Times'' is a British daily national newspaper based in London. It began in 1785 under the title ''The Daily Universal Register'', adopting its modern name on 1 January 1788. ''The Times'' and its sister paper ''The Sunday Times'' (fo ...
TimesOnline
''The Times'' is a British daily national newspaper based in London. It began in 1785 under the title ''The Daily Universal Register'', adopting its modern name on 1 January 1788. ''The Times'' and its sister paper ''The Sunday Times'' (fo ...
'', 1 September 2003.
{{animal cognition
Intelligence
Intelligence has been defined in many ways: the capacity for abstraction, logic, understanding, self-awareness, learning, emotional knowledge, reasoning, planning, creativity, critical thinking, and problem-solving. It can be described as t ...