Gymnotiformes

Gymnotiformes Temporal range: Late Jurassic–Recent PreЄ Є O S D C P T J K Pg N [1]
Black ghost knifefish, Apteronotus albifrons
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Actinopterygii
Order:	Gymnotiformes

The Gymnotiformes are a group of teleost bony fishes commonly known as the Neotropical or South American knifefishes. They have long bodies and swim using undulations of their elongated anal fin. Found exclusively in fresh water, these mostly nocturnal fishes are capable of producing electric fields for navigation, communication, and, in the case of the electric eel (Electrophorus electricus), attack and defense. A few species are familiar to the aquarium trade, such as the black ghost knifefish (Apteronotus albifrons), the glass knifefish (Eigenmannia virescens), and the banded knifefish (Gymnotus carapo).

Description

Aside from the electric eel (Electrophorus electricus), Gymnotiformes are slender fish with narrow bodies and a tapering tail, hence the common name of "knifefishes". They have no pelvic fins or dorsal fin, but do possess a greatly elongated anal fin that stretches along almost the entire underside of the body. The fish swim by rippling this fin, keeping their bodies rigid. This means of propulsion allows them to move backwards as easily as they move forwards.^[2]

The caudal fin is absent or, in the Apteronotids, greatly reduced. The gill opening is restricted. The anal opening is under the head or the pectoral fins.^[3]

These fish possess electric organs that allow them to produce electricity. In most gymnotiforms, the electric organs are derived from muscle cells. However, adult apteronotids are one exception, as their electric organs are derived from nerve cells (spinal electromotor neurons). In gymnotiforms the electric organ discharge (EOD) may be continuous or pulse like. If continuous, it is generated day and night throughout the entire life of the individual. Certain aspects of the electric signal are unique to each species, especially a combination of the pulse waveform, duration, amplitude, phase and frequency.^[4]

The electric organs of most Gymnotiformes produce tiny discharges of just a few millivolts, far too weak to cause any harm to other fish. Instead, they are used to help navigate the environment, including locating the bottom-dwelling invertebrates that compose their diet. They may also be used to send signals between fish of the same species.^[5] In addition to this low level field, the electric eel also has the capability to produce much more powerful discharges to stun prey.^[2]

Taxonomy

There are currently about 150 known species in 32 genera contained in 5 families, and at least 50 or so additional species are known and are yet to be formally described.^[6] The actual number of species in the wild is unknown.^[7] This group is thought to be the sister group to the Siluriformes^[8] from which they diverged in the Cretacous Period (about 120 million years ago).

The families are classified over suborders and superfamilies as below.^[9]

Order Gymnotiformes

Suborder Gymnotoidei

Family Gymnotidae (banded knifefishes and electric eel)

Suborder Sternopygoidei

Superfamily Rhamphichthyoidea

Family Rhamphichthyidae (sand knifefishes)

Family Hypopomidae (bluntnose knifefishes)

Superfamily Apteronotoidea

Family Sternopygidae (glass and rat-tail knifefishes)

Family Apteronotidae (ghost knifefishes)

Distribution and habitat

Gymnotiform fishes inhabit freshwater rivers and streams throughout the humid Neotropics, ranging from Guatemala to Northern Argentina. They are nocturnal fishes. The families Gymnotidae and Hypopomidae are most diverse (numbers of species) and abundant (numbers of individuals) in small "terra-firme" (non-floodplain" streams and rivers, and in floodplain "floating meadows" of aquatic macrophytes (e.g., Eichornium, the Amazonian water hyacinth). Apteronotidae and Sternopygidae are most diverse and abundant in large rivers. Species of Rhamphichthyidae are moderately diverse in all these habitat types.

Evolution

Gymnotiformes are among the more derived members of Ostariophysi, a lineage of primary freshwater fishes. The only known fossils are from the Miocene about 7 million years ago of Bolivia.^[10]

Gymnotiformes has no extant species in Africa. This may be because they did not spread into Africa before South America and Africa split, or it may be that they were outcompeted by Mormyriformes, which are similar in that they also use electrolocation.^[6]

Interestingly, Gymnotiformes and Mormyriformes have developed their electric organs and electrosensory systems (ESSs) through incredible convergent evolution.^[11] As Arnegard et al. (2005) and Albert and Crampton (2005) show,^[12][13] their last common ancestor was roughly 140 to 208 million years ago, and at this time they did not possess ESSs. Each species of Mormyrus (Family: Mormyridae) and Gymnotus (Family: Gymnotidae) have evolved a completely unique waveform that allows the individual fish to identify between species, genders, individuals and even between the mates with better fitness levels.^[14] The differences include the direction of the initial phase of the wave (positive or negative, which correlates to the direction of flow of current through the electrocytes in the electric organ), the amplitude of the wave, the frequency of the wave, and the number of phases of the wave. Thus, the parallels between these distantly related species is astounding.

One significant force driving this evolution is predation.^[15] The most common predators of Gymnotiformes include the closely related Siluriformes (catfish) as well as predation within families (E. electricus is one of the largest predators of Gymnotus). These predators sense electric fields, but only at low frequencies, thus certain species of Gymnotiformes, such as those in Gymnotus, have shifted the frequency of their signal so that they can be effectively invisible.^[15][16][17]

As sexual selection is another driving force which has an unusual influence in that females exhibit preference for males with low frequency signals (which are energetically expensive and easily detected by predators),^[15] however most males exhibit this frequency only intermittently. They also prefer males with longer pulse EODs,^[14] also energetically expensive, and large tail length. These are all signs that indicate some ability to exploit resources,^[15] thus indicating better lifetime reproductive success.

See also

• Electric fish
• Bioelectromagnetism

References

1. ^ Froese, Rainer, and Daniel Pauly, eds. (2007). "Gymnotiformes" in FishBase. Apr 2007 version.
2. ^ ^{a b} Ferraris, Carl J. (1998). Paxton, J.R. & Eschmeyer, W.N.. ed. Encyclopedia of Fishes. San Diego: Academic Press. pp. 111–112. ISBN 0-12-547665-5. 
3. ^ Albert, J.S. 2001. Species diversity and phylogenetic systematics of American knifefishes (Gymnotiformes, Teleostei). Misc. Publ. Mus. Zool. University of Michigan, 190:1-127.
4. ^ Crampton, W.G.R. and J.S. Albert. 2006. Evolution of electric signal diversity in gymnotiform fishes. Pp. 641-725 in Communication in Fishes. F. Ladich, S.P. Collin, P. Moller & B.G Kapoor (eds.). Science Publishers Inc., Enfield, NH.
5. ^ Vincent Fugère, Hernán Ortega and Rüdiger Krahe. 2010. Electrical signalling of dominance in a wild population of electric fish. Biol. Lett. 10.1098/rsbl.2010.0804
6. ^ ^{a b} Albert, J.S., and W.G.R. Crampton. 2005. Electroreception and electrogenesis. Pp. 431-472 in The Physiology of Fishes, 3rd Edition. D.H. Evans and J.B. Claiborne (eds.). CRC Press.
7. ^ Albert, J.S. and W.G.R. Crampton. 2005. Diversity and phylogeny of Neotropical electric fishes (Gymnotiformes). Pp. 360-409 in Electroreception. T.H. Bullock, C.D. Hopkins, A.N. Popper, and R.R. Fay (eds.). Springer Handbook of Auditory Research, Volume 21 (R.R. Fay and A. N. Popper, eds). Springer-Verlag, Berlin.
8. ^ Fink and Fink, 1996
9. ^ Nelson
10. ^ Albert, J.S. and W.L. Fink. 2007. Phylogenetic relationships of fossil Neotropical electric fishes (Osteichthyes: Gymnotiformes) from the Upper Miocene of Bolivia. Journal Vertebrate Paleontology 27(1):17-25.
11. ^ Hopkins, C. D. 1995. Convergent designs for electrogenesis and electroreception. Current Opinion in Neurobiology 5:769-777.
12. ^ Albert, J. S., and W. G. R. Crampton. 2006. Electroreception and electrogenesis. Pp. 429-470 in P. L. Lutz, ed. The Physiology of Fishes. CRC Press, Boca Raton, FL.
13. ^ Arnegard, M. E., S. M. Bogdanowicz, and C. D. Hopkins. 2005. Multiple cases of striking genetic similarity between alternate electric fish signal morphs in sympatry. Evolution 59:324-343.
14. ^ ^{a b} Arnegard, M. E., P. B. McIntyre, L. J. Harmon, M. L. Zelditch, W. G. R. Crampton, J. K. Davis, J. P. Sullivan, S. Lavoue, and C. D. Hopkins. 2010. Sexual signal evolution outpaces ecological divergence during electric fish species radiation. American Naturalist 176:335-356.
15. ^ ^{a b c d} Hopkins, C. D. 1999a. Design features for electric communication. Journal of Experimental Biology 202:1217-1228.
16. ^ Stoddard, P. K. 1999. Predation enhances complexity in the evolution of electric fish signals. Nature 400:254-256.
17. ^ Stoddard, P. K. 2002b. The evolutionary origins of electric signal complexity. Journal of Physiology-Paris 96:485-491.

External links

• Gymnotiform overview
• Photos of various gymnotiforms