Freshwater elephantfish | |
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Gnathonemus petersii | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Actinopterygii |
Order: | Osteoglossiformes |
Family: | Mormyridae |
Genera | |
see text |
The Mormyridae, sometimes called "elephantfish" (more properly freshwater elephantfish), are a superfamily of weakly electric fish in the order Osteoglossiformes native to Africa. [1] It is by far the largest family in the order, with around 200 species. Members of the family can be popular, if challenging, aquarium species. These fish have a large brain size and unusually high intelligence.
They are not to be confused with the marine and brackish-water callorhinchid elephantfish (family Callorhinchidae) of Southern Hemisphere oceans.
The elephantfish are a diverse family, with a wide range of different sizes and shapes. The smallest are just 5 cm (2.0 in) in adult length, while the largest reach up to 1.5 m (4.9 ft). They do, however, have a number of unique features in common. Firstly, their cerebellum is greatly enlarged, enabling them to interpret complex bio-electrical signals, [2] and to the large size of the valve. [3] [4] Secondly, an auditory vesicle (a small bladder) is present inside the labyrinth of the left and right inner ears. This vesicle, together with a bag with an otolith (sacculum containing the otolith sagitta), itself communicating to the lagena (containing the otolith asteriscus), is in fact unique among vertebrates, completely independent of the other organs; it is neither connected to the labyrinth to which only one otolith bag (the utriculus containing the otolith lapillus) is attached, nor is it connected to the swim bladder (except in embryos) of which it has the same histological structure, nor is it therefore related to the pharynx. [5]
Some species possess modifications of their mouthparts to facilitate electrolocating and feeding on small invertebrates buried in muddy substrates. The shape and structure of these leads to the popular name "elephant-nosed fish" for those species with particularly prominent mouth extensions. The extensions to the mouthparts usually consist of a fleshy elongation attached to the lower jaw. They are flexible, and equipped with touch, and possibly taste, sensors. The mouth is not protrusible, and the head (including the eyes), the dorsum, and belly are covered by a thin layer of skin that is perforated with small pores leading to electroreceptors.[ citation needed ]
The retina is called a "grouped retina", an eye structure seen in mormyrids and a few other fishes. [6] Instead of being smooth, their retina is composed of tiny cups, acting like parabolic mirrors. Because of the murky waters they inhabit, the cones in their eyes have adapted to see only red light. The cups are made of four layers of light-reflecting proteins, funneling red light to areas of cones, intensifying its brightness 10-fold, while the rods are hit by light from other wavelengths. [7] Only a single gonad is present, located on the left side of their body. [8] The Mormyridae and the closely related genus Gymnarchus are also unique in being the only vertebrates where the male sperm cell does not have a flagellum. [9]
Elephantfish possess electric organs that generate weak electric fields, and electroreceptors (ampullae of Lorenzini, knollenorgans, and Mormyromasts) that detect small variations in these electric fields caused by the presence of prey or other objects of different conductivities. This allows them to sense their environment in turbid waters where vision is impaired by suspended matter. [10] [11]
Electric fish can be classified into two types: pulse fish or wave fish. Pulse-type discharges are characterized by long intervals between electric discharges, whereas wave-type discharges occur when the interval between consecutive pulses is so brief that the discharges fuse together to form a wave. [12] The electric discharge is produced from an electric organ that evolved from muscle, as can also be seen in gymnotiform electric fish, electric rays, and skates. The convergent evolution between the South American gymnotiforms and the African Mormyridae is remarkable, with the electric organ being produced by the substitution of the same amino acid in the same voltage-gated sodium channel despite the two groups of fish being on different continents and the evolution of the electric sense organ being separated in time by around 60 million years. [13] Convergent changes to other key transcription factors and regulatory pathways in both Gymnotiforms and Mormyridae also contributed to the evolution of the electric sense organ. [14]
The roughly 221 species of elephantfish which are sometimes grouped into two subfamilies, the Mormyrinae and Petrocephalinae. The latter has only a single genus:
Phylogeny [15] [16] | |||
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Eschmeyer's Catalog of Fishes classifies the family as follows [17] Family Mormyridae
The Medjed was a sacred fish in Ancient Egypt. At the city of Per-Medjed, better known as Oxyrhynchus, whose name means "sharp-nosed" after the fish, archaeologists have found fishes depicted as bronze figurines, mural paintings, or wooden coffins in the shape of fishes with downturned snouts, with horned sun-disc crowns like those of the goddess Hathor. The depictions have been described as resembling members of the genus Mormyrus . [19]
The Gymnotiformes are an order of teleost bony fishes commonly known as Neotropical knifefish or South American knifefish. They have long bodies and swim using undulations of their elongated anal fin. Found almost exclusively in fresh water, these mostly nocturnal fish are capable of producing electric fields to detect prey, for navigation, communication, and, in the case of the electric eel, attack and defense. A few species are familiar to the aquarium trade, such as the black ghost knifefish, the glass knifefish, and the banded knifefish.
Osteoglossiformes is a relatively primitive order of ray-finned fish that contains two sub-orders, the Osteoglossoidei and the Notopteroidei. All of at least 245 living species inhabit freshwater. They are found in South America, Africa, Australia and southern Asia, having first evolved in Gondwana before that continent broke up. In 2008, several new species of marine osteoglossiforms were described from the Danish Eocene Fur Formation, dramatically increasing the diversity of this group. This implies that the Osteoglossomorpha is not a primary freshwater fish group with the osteoglossiforms having a typical Gondwana distribution.
Osteoglossomorpha is a group of bony fish in the Teleostei.
An electric fish is any fish that can generate electric fields, whether to sense things around them, for defence, or to stun prey. Most fish able to produce shocks are also electroreceptive, meaning that they can sense electric fields. The only exception is the stargazer family (Uranoscopidae). Electric fish, although a small minority of all fishes, include both oceanic and freshwater species, and both cartilaginous and bony fishes.
Electroreception and electrogenesis are the closely related biological abilities to perceive electrical stimuli and to generate electric fields. Both are used to locate prey; stronger electric discharges are used in a few groups of fishes to stun prey. The capabilities are found almost exclusively in aquatic or amphibious animals, since water is a much better conductor of electricity than air. In passive electrolocation, objects such as prey are detected by sensing the electric fields they create. In active electrolocation, fish generate a weak electric field and sense the different distortions of that field created by objects that conduct or resist electricity. Active electrolocation is practised by two groups of weakly electric fish, the Gymnotiformes (knifefishes) and the Mormyridae (elephantfishes), and by Gymnarchus niloticus, the African knifefish. An electric fish generates an electric field using an electric organ, modified from muscles in its tail. The field is called weak if it is only enough to detect prey, and strong if it is powerful enough to stun or kill. The field may be in brief pulses, as in the elephantfishes, or a continuous wave, as in the knifefishes. Some strongly electric fish, such as the electric eel, locate prey by generating a weak electric field, and then discharge their electric organs strongly to stun the prey; other strongly electric fish, such as the electric ray, electrolocate passively. The stargazers are unique in being strongly electric but not using electrolocation.
Peters's elephant-nose fish is an African freshwater elephantfish in the genus Gnathonemus. Other names in English include elephantnose fish, long-nosed elephant fish, and Ubangi mormyrid, after the Ubangi River. The Latin name petersii is probably for the German naturalist Wilhelm Peters. The fish uses electrolocation to find prey, and has the largest brain-to-body oxygen use ratio of all known vertebrates.
In biology, the electric organ is an organ that an electric fish uses to create an electric field. Electric organs are derived from modified muscle or in some cases nerve tissue, called electrocytes, and have evolved at least six times among the elasmobranchs and teleosts. These fish use their electric discharges for navigation, communication, mating, defence, and in strongly electric fish also for the incapacitation of prey.
Marcusenius is a genus of the elephantfish group native to Africa. Its members are highly diverse in size, with the smallest species reaching less than 15 cm (6 in) and the largest more than 1 m (3.3 ft).
Mormyrus is a genus of ray-finned fish in the family Mormyridae. They are weakly electric, enabling them to navigate, to find their prey, and to communicate with other electric fish.
A Knollenorgan is an electroreceptor in the skin of weakly electric fish of the family Mormyridae (Elephantfish) from Africa. The structure was first described by Viktor Franz (1921), a German anatomist unaware of its function. They are named after "Knolle", German for "tuberous root" which describes their structure.
Mormyrops is a genus of weakly electric fish in the family Mormyridae from freshwater in Africa. They are characterized by an elongate head measuring twice as long as high, and no teeth on the palate or the tongue. The genus includes the largest member of the mormyrid family, the cornish jack at up to 1.5 m (4.9 ft) in length.
The subfamily Mormyrinae contains all but one of the genera of the African freshwater fish family Mormyridae in the order Osteoglossiformes. They are often called elephantfish due to a long protrusion below their mouths used to detect buried invertebrates that is suggestive of a tusk or trunk. They can also be called tapirfish.
The jamming avoidance response is a behavior of some species of weakly electric fish. It occurs when two electric fish with wave discharges meet – if their discharge frequencies are very similar, each fish shifts its discharge frequency to increase the difference between the two. By doing this, both fish prevent jamming of their sense of electroreception.
Campylomormyrus is a genus of elephantfish in the family Mormyridae.
Myomyrus is a genus of elephantfish in the family Mormyridae. Its members reach about 25–30 cm (10–12 in) in length and are restricted to the Congo River Basin in Africa.
Paramormyrops is a genus of elephantfish in the family Mormyridae from Africa.
Petrocephalus is a genus of ray-finned fish in the family Mormyridae. All the fish species of this genus are endemic to Africa.
Stomatorhinus is a genus of small elephantfish in the family Mormyridae.
Cryptomyrus is a genus of mormyrid fish native to Gabon.
The Mormyroidea are a superfamily of fresh water fishes endemic to Africa that, together with the families Hiodontidae, Osteoglossidae, Pantodontidae and Notopteridae, represents one of the main groups of living Osteoglossiformes. They stand out for their use of weak electric fields, which they use to orient themselves, reproduce, feed, and communicate.