Mormyridae

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Freshwater elephantfish
Elefantenrusselfisch.jpg
Gnathonemus petersii
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Osteoglossiformes
Family: Mormyridae
Subfamilies

Mormyrinae
Petrocephalinae

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.

Contents

They are not to be confused with the marine and brackish-water callorhinchid elephantfish (family Callorhinchidae) of Southern Hemisphere oceans.

Description and biology

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]

Electric fields

Mormyromast, a type of electroreceptor found only in mormyrid fishes Mormyromast diagram.svg
Mormyromast, a type of electroreceptor found only in mormyrid fishes

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]

Classification

The roughly 221 species of elephantfish are grouped into two subfamilies, the Mormyrinae and Petrocephalinae. The latter has only a single genus:

Phylogeny [15] [16]
Mormyridae
Petrocephalinae

Petrocephalus

Mormyrinae

Myomyrus

Mormyrops

Brienomyrus

Isichthys

Mormyrus

Pollimyrus

Stomatorhinus

Paramormyrops

Cryptomyrus

Boulengeromyrus

Ivindomyrus

Hyperopisus

Brevimyrus

Hippopotamyrus

Campylomormyrus

Gnathonemus

Genyomyrus

Marcusenius

Family Mormyridae

In culture

Bronze figurine of Oxyrhynchus fish, Late Period-Ptolemaic Egypt Oxyrhynchus fish Late Period-Ptolemaic (cropped).jpg
Bronze figurine of Oxyrhynchus fish, Late Period-Ptolemaic Egypt

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 . [18]

Related Research Articles

<span class="mw-page-title-main">Gymnotiformes</span> Order of bony fishes

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.

<span class="mw-page-title-main">Osteoglossomorpha</span> Superorder of fishes

Osteoglossomorpha is a group of bony fish in the Teleostei.

<span class="mw-page-title-main">Electric fish</span> Fish that can generate electric fields

An electric fish is any fish that can generate electric fields. Most electric fish 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.

<span class="mw-page-title-main">Electroreception and electrogenesis</span> Biological electricity-related abilities

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.

<span class="mw-page-title-main">Peters's elephantnose fish</span> Species of fish

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.

<span class="mw-page-title-main">Electric organ (fish)</span> Organ in electric fish

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, 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.

<i>Hippopotamyrus</i> Genus of ray-finned fishes

Hippopotamyrus is a genus of elephantfish in the family Mormyridae.

<i>Marcusenius</i> Genus of ray-finned fishes

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).

<i>Mormyrus</i> Genus of ray-finned fishes

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.

<span class="mw-page-title-main">Knollenorgan</span>

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.

<i>Mormyrops</i> Genus of ray-finned fishes

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.

<span class="mw-page-title-main">Mormyrinae</span> Subfamily of fishes

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.

<span class="mw-page-title-main">Jamming avoidance response</span> Behavior performed by weakly electric fish to prevent jamming of their sense of electroreception

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.

<i>Campylomormyrus</i> Genus of ray-finned fishes

Campylomormyrus is a genus of elephantfish in the family Mormyridae.

<i>Myomyrus</i> Genus of ray-finned fishes

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.

<i>Paramormyrops</i> Genus of ray-finned fishes

Paramormyrops is a genus of elephantfish in the family Mormyridae from Africa.

<i>Petrocephalus</i> Genus of ray-finned fishes

Petrocephalus is a genus of ray-finned fish in the family Mormyridae. All the fish species of this genus are endemic to Africa.

<span class="mw-page-title-main">Arapaiminae</span> Subfamily of ray-finned fishes

Arapaiminae is a subfamily of freshwater osteoglossiform (bony-tongued) fishes belonging to the family Osteoglossidae. It includes the South American arapaimas of the Amazon and Essequibo basins and the African arowana from the watersheds of the Sahelo-Sudanese region, Senegal, Gambia, and parts of Eastern Africa. This subfamily is sometimes raised to the rank of family, as Arapaimidae. A commonly used synonym is Heterotidinae, but according to the ICZN, Arapaiminae has priority.

<i>Cryptomyrus</i> Genus of ray-finned fishes

Cryptomyrus is a genus of mormyrid fish native to Gabon.

<span class="mw-page-title-main">Mormyroidea</span>

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.

References

  1. Froese, Rainer, and Daniel Pauly, eds. (2017). "Mormyridae" in FishBase . April 2017 version.
  2. Chapman, Lauren J.; Hulen, Kevin G. (2001). "Implications of hypoxia for the brain size and gill morphometry of mormyrid fishes". Journal of Zoology. 254 (4): 461–472. doi:10.1017/S0952836901000966.
  3. Glickstein, M.; Voogd, J. (2009). "Cerebellum: Evolution and Comparative Anatomy". Encyclopedia of Neuroscience: 743–756. doi:10.1016/B978-008045046-9.00947-5. ISBN   978-0-08-045046-9.
  4. Wullimann, Mario F.; Rooney, Donal J. (1990). "A direct cerebello-telencephalic projection in an electrosensory mormyrid fish" (PDF). Brain Research. 520 (1–2): 354–357. doi:10.1016/0006-8993(90)91730-5. PMID   1698507. S2CID   12951395.
  5. Orts, S. (1967): Contribution to the comparative anatomy and systematics of Mormyroïdes (RAOS Prize 1966) - Ann. Acad. Roy. Sc. d'Outre Mer, Classe des Sc. Nat. et Méd., Bruxelles, XVII, 3, 1-89, 30 figs, 3 tabs, 8 pls, 1967 (in French)
  6. Francke, Mike; Kreysing, Moritz; Mack, Andreas; et al. (2014). "Grouped retinae and tapetal cups in some Teleostian fish: Occurrence, structure, and function". Progress in Retinal and Eye Research. 38: 43–69. doi:10.1016/j.preteyeres.2013.10.001. PMID   24157316. S2CID   39081714.
  7. Elephant-Nosed Fish Has Funky Eyes, Too
  8. Communication Behavior and Sensory Mechanisms in Weakly Electric Fishes
  9. "Mormyridae - African weakly electric fishes". Archived from the original on 2013-06-18. Retrieved 2013-09-15.
  10. Bustami, H.P. (2007). Smart elephant fish navigates in darkness with electric fields. life-of-science.net
  11. The generation of these electric fields and their use in providing the fish with additional sensory input from the environment is the subject of considerable scientific research, as is research into communication between and within species
  12. Caputi, A. A. (1999). "The electric organ discharge of pulse gymnotiforms: the transformation of a simple impulse into a complex spatiotemporal electromotor pattern". Journal of Experimental Biology. 202 (# (Pt 10)): 1229–1241. doi:10.1242/jeb.202.10.1229. PMID   10210664.
  13. Arnegard, M. E.; Zwickl, D. J.; Lu, Y.; Zakon, H. H. (2010-12-02). "Old gene duplication facilitates origin and diversification of an innovative communication system--twice". Proceedings of the National Academy of Sciences. 107 (51): 22172–22177. doi: 10.1073/pnas.1011803107 . PMC   3009798 . PMID   21127261.
  14. Gallant, Jason R.; Traeger, Lindsay L.; Volkening, Jeremy D.; Moffett, Howell; Chen, Po-Hao; Novina, Carl D.; Phillips, George N.; Anand, Rene; Wells, Gregg B.; Pinch, Matthew; Güth, Robert (2014-06-27). "Genomic basis for the convergent evolution of electric organs". Science. 344 (6191): 1522–1525. Bibcode:2014Sci...344.1522G. doi:10.1126/science.1254432. PMC   5541775 . PMID   24970089.
  15. Lavoué, S.; Sullivan, J. P.; Hopkins, C. D. (2003). "Phylogenetic utility of the first two introns of the S7 ribosomal protein gene in African electric fishes (Mormyroidea: Teleostei) and congruence with other molecular markers". Biological Journal of the Linnean Society. 78 (2): 273–292. doi: 10.1046/j.1095-8312.2003.00170.x .
  16. Sullivan, J. P.; Lavoué, S.; Hopkins, C. D. (2000). "Molecular systematics of the African electric fishes (Mormyroidea: Teleostei) and a model for the evolution of their electric organs". Journal of Experimental Biology. 203 (Pt 4): 665–683. doi: 10.1242/jeb.203.4.665 . PMID   10648209.
  17. Sullivan, J. P.; Lavoué, S.; Hopkins, C. D. (2016). "Cryptomyrus: a new genus of Mormyridae (Teleostei, Osteoglossomorpha) with two new species from Gabon, West-Central Africa". ZooKeys (561): 117–150. doi: 10.3897/zookeys.561.7137 . PMC   4768369 . PMID   27006619.
  18. Van Neer, Wim; Gonzalez, Jérôme (2019). "A Late Period fish deposit at Oxyrhynchus (el-Bahnasa, Egypt)". In Peters, Joris; McGlynn, George; Goebel, Veronika (eds.). Documenta Archaeobiologiae Animals: Cultural Identifiers In Ancient Societies? (PDF). Rahden, Westfalia, Germany: Verlag Marie Leidorf. ISBN   978-3-89646-674-7.