Walking fish

A mudskipper, a type of walking fish, perched on land.

A walking fish, or ambulatory fish, is a fish that is able to travel over land for extended periods of time. Some other modes of non-standard fish locomotion include "walking" along the sea floor, for example, in handfish or frogfish.

Types

Pacific leaping blenny (Alticus arnoldorum) hopping

Most commonly, walking fish are amphibious fish. Able to spend longer times out of water, these fish may use a number of means of locomotion, including springing, snake-like lateral undulation, and tripod-like walking. The mudskippers are probably the best land-adapted of contemporary fish and are able to spend days moving about out of water and can even climb mangroves, although to only modest heights.^[1] The climbing gourami is often specifically referred to as a "walking fish", although it does not actually "walk", but rather moves in a jerky way by supporting itself on the extended edges of its gill plates and pushing itself by its fins and tail. Some reports indicate that it can also climb trees.^[2]

The epaulette shark (Hemiscyllium ocellatum) tends to live in shallow waters where swimming is difficult, and can often be seen walking over rocks and sand by using its muscular pectoral fins.^[3] It lives in areas of great variation in water depth, usually where the tide falls below its location. If it finds itself out of water, it can survive for several hours, and is capable of walking over land to get to water. This means that it is easily observed by beachgoers in its natural range.

There are a number of fish that are less adept at actual walking, such as the walking catfish. Despite being known for "walking on land", this fish usually wriggles and may use its pectoral fins to aid in its movement. Walking catfish have a respiratory system that allows them to live out of water for several days. Some are invasive species, for example, the Northern snakehead in the U.S.^[4] Polypterids have rudimentary lungs and can also move about on land, though rather clumsily. The mangrove rivulus can survive for months out of water and can move to places like hollow logs.^[5][6][7][8]

Some species of fish can "walk" along the sea floor but not on land. One such animal is the flying gurnard (it does not actually fly, and should not be confused with flying fish). The batfishes of the Ogcocephalidae family (not to be confused with Batfish of Ephippidae) are also capable of walking along the sea floor. Bathypterois grallator, also known as a "tripodfish", stands on three fins on the bottom of the ocean and hunts for food.^[9] The African lungfish (P. annectens) can use its fins to "walk" along the bottom of its tank in a manner similar to the way amphibians and land vertebrates use their limbs on land. ^[10][11][12]

Evolutionary link

Tiktaalik (reconstruction)

In modern fish, the "walking" ability differs from that of tetrapods (four-limbed animals). The theory of evolution suggests that life originated in the oceans and later moved onto land, and paleontologists have long been looking for a missing evolutionary link between ocean-living and land-living animals. In 2006, a fossil Tiktaalik roseae was found which has many features of wrist, elbow, and neck that are akin to those of tetrapods.^[13] It belongs to a group of lobe-finned fish called Rhipidistia, which according to some theories, were the ancestors of all tetrapods.

Comparison of fish with tetrapod-like features

A number of fish, both extant and prehistoric, have featured some characteristics related to locomotion that are typical of tetrapods.

Species	venturing onto land	tetrapod-like spine			tetrapod-like appendages					digit-like bones
		5 axial regions	interlocking vertebrae	fully ossified vertebrae	shoulder & skull separation	functional 'intra-fin' joints	fins adapted for walking rather than swimming	strong & muscled fins	humerus, radius & ulna bones	differentiated distal radial bones	jointed distal radial bones
Panderichthys rhombolepis †	?	No	No	No	No	?	No	Yes	Yes	Yes[14]	No
Sauripterus taylori †	?	No	No	No	No	?	No	Yes	Yes	Yes	Yes[15]
Tiktaalik roseae †	?	No	No	No	Yes	Yes	No	Yes	Yes	No	No
Tarrasius problematicus †	?	Yes[16]	No	No	No	No	No	No	No	No	No
Leptolepis koonwarriensis †	?	No	No	Yes[17]	No	No	No	No	No	No	No
Eastmanosteus pustulosus †	?	No	No	No	Yes[18]	No	No	No	No	No	No
Atractosteus spatula	No	No	Yes	Yes[19]	No	No	No	No	No	No	No
Periophthalmus papilio	Yes	No	No	No	No	Yes[20][21]	No	No	No	No	No
Brachionichthys hirsutus	No	No	No	No	No	No	Yes	No	No	No	No
Ogcocephalus darwini	No	No	No	No	No	No	Yes	No	No	No	No
Antennarius maculatus	No	No	No	No	No	No	Yes	No	No	No	No
Protopterus annectens	Yes	No	No	No	No	No	?[22]	No	No	No	No
Latimeria chalumnae	No	No	No	No	No	No	No	Yes	No	No	No
Polypterus bichir lapradei	Yes	No	No	No	No	No	No	Yes	No	No	No
Chelidonichthys cuculus	No	No	No	No	No	No	Yes (3 rays)	No	No	? (3 rays)	No
Hemiscyllium ocellatum	Yes	No	No	No	Yes[23]	No	?[24]	No	No	No	No

Popular culture

Another usage of the term walking fish is in reference to the "Darwin fish", a bumper sticker parody of the Ichthys, a symbol of Christianity.

In The Simpsons episode "Treehouse of Horror V" segment "Time and Punishment," Homer accidentally kills a walking fish by sitting on it, resulting in irreparable damage to the timeline.

In another Simpsons episode, "Judge Me Tender", one of Ned's fish walks out of its tank.

Gallery

• Play media

  Alticus arnoldorum climbing up a vertical piece of Plexiglas

• 

  An epaulette shark clambering over the sand on a beach

• 

  Ogcocephalus parvus

See also

• Evolution of fish

References

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2. Climbing Fish
3. http://elasmo-research.org/education/topics/b_dorsal_out.htm
4. "Maryland Suffers Setback in War on Invasive Walking Fish", National Geographic News July 12, 2002
5. Shells, trees and bottoms: Strange places fish live
6. "Tropical fish can live for months out of water". Reuters. 15 November 2007.
7. Fish Lives in Logs, Breathing Air, for Months at a Time
8. Fish Lives in Logs, Breathing Air, for Months at a Time
9. Jones, AT; KJ Sulak (1990). "First Central Pacific Plate and Hawaiian Record of the Deep-sea Tripod Fish Bathypterois grallator (Pisces: Chlorophthalmidae)" (PDF). Pacific Science. 44 (3): 254–7.
10. Fish uses fins to walk and bound
11. Behavioral evidence for the evolution of walking and bounding before terrestriality in sarcopterygian fishes
12. A Small Step for Lungfish, a Big Step for the Evolution of Walking
13. Daeschler, E.B., Shubin, N.H. and Jenkins, Jr. F.A. (6 April 2006). "A Devonian tetrapod-like fish and the evolution of the tetrapod body plan". Nature. 440 (7085): 757–763. doi:10.1038/nature04639. PMID 16598249.
14. C. A. Boisvert, E. Mark-Kurik, P. E. Ahlberg (4 December 2008). "The pectoral fin of Panderichthys and the origin of digits". Nature. 456 (7222): 636–638. Bibcode:2008Natur.456..636B. doi:10.1038/nature07339. PMID 18806778.
15. E. B. Daeschler, N. H. Shubin (8 January 1998). "Fish with Fingers?". Nature. 391 (6663): 133–133. doi:10.1038/34317.
16. Lauren Cole Sallan (23 May 2012). "Tetrapod-like axial regionalization in an early ray-finned fish" (PDF). Proceedings of the Royal Society B. 279 (1741): 3264–3271. doi:10.1098/rspb.2012.0784. PMID 22628471.
17. Shukla, J.P. (2007). Fish & Fisheries. Rastogi Publications. pp. 24–25. ISBN 81-7133-800-3.
18. K. Trinajstic et al. (12 July 2013). "Fossil Musculature of the Most Primitive Jawed Vertebrates". Science. 341 (6142): 160–164. doi:10.1126/science.1237275.
19. Olsen, Stanley J. (1968). Fish, Amphibian and Reptile Remains from Archaeological Sites. Acme Bookbinding. p. 4. ISBN 0-87365-163-4.
20. S. M. Kawano and R. W. Blob (August 2, 2013). "Propulsive Forces of Mudskipper Fins and Salamander Limbs during Terrestrial Locomotion: Implications for the Invasion of Land" (PDF). Integrative and Comparative Biology. 53: 283–294. doi:10.1093/icb/ict051. PMID 23667046.
21. C. M. Pace and A. C. Gibb (July 15, 2009). "Mudskipper pectoral fin kinematics in aquatic and terrestrial environments" (PDF). The Journal of Experimental Biology. 212: 2279–2286. doi:10.1242/jeb.029041. PMID 19561218.
22. H.M. King, N.H. Shubin, M.I. Coates, M.E. Hale (27 December 2011). "Behavioral evidence for the evolution of walking and bounding before terrestriality in sarcopterygian fishes". PNAS. 108 (52): 21146–51. doi:10.1073/pnas.1118669109.
23. Hanken, James; Hall, Brian K. (1993). The Skull, Volume 2: Patterns of Structural and Systematic Diversity. University of Chicago Press. p. 209. ISBN 0-226-31570-3.
24. Goto, T., Nishida, K. and Nakaya, K. (Sep 1999). "Internal morphology and function of paired fins in the epaulette shark, Hemiscyllium ocellatum". Ichthyological Research. 46 (3): 281–287. doi:10.1007/BF02678514.