690:. Their eyes are located high on their head compared to other gobies, and they blink by lowering their eyes as a membrane called the dermal cup rises to cover them. Although other fully aquatic goby species do not have the ability to blink, mudskippers did not evolve different muscles or glands to blink with; their blinking is accomplished with the same muscles in a different configuration. Rather than having specialized glands to produce eye lubricant, the fluid film on their cornea is likely composed of mucus secreted by their skin and water from their environment, possibly stored in the infraorbital space behind the dermal cup membrane. Mudskippers likely evolved blinking in response to conditions of terrestrial life, such as to keep their eyes wet out of water (they blink more frequently in high evaporation conditions and only when colliding with things in water) and to clean and protect the eye from debris, which may adhere to the eye or approach at a faster, more dangerous speed when in air versus when in water. Their eyes are not elevated when they are still juveniles, which are fully aquatic.
575:
457:
70:
392:. These burrows are most often characterised by their smooth and vaulted ceilings. The way the males dig these burrows has been found to be directly linked to their ability to survive submerged in almost anoxic water. It has also been found to play a crucial role in the development of the eggs within the burrow. Mudskippers are quite active when out of water, feeding and interacting with one another, for example, to defend their
438:
851:
42:
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624:. Another important adaptation that aids breathing while out of water is their enlarged gill chambers, where they retain a bubble of air. These chambers close tightly when the fish is above water, due to a ventromedial valve of the gill slit, keeping the gills moist, and allowing them to function while exposed to air. Gill filaments are stiff and do not coalesce when out of water.
591:
404:
Mudskippers are amphibious. When leaving the water and moving into a more dry environment on land, they are still able to breathe using water that is trapped inside their large gill chambers. They are also able to absorb oxygen from the lining of their mouth and throat, allowing them to stay out
396:
and court potential partners. Once the male has completed digging his burrow he will resurface and will begin attempting to attract a female through assorted yet typical displays. These displays consist of body undulations, different postures and energetic movements. Once the female has made her
616:); this is only possible when the mudskippers are wet, limiting them to humid habitats and requiring that they keep themselves moist. The ability to breathe through their skin is associated with increased capillary density in their skin. This mode of breathing, similar to that employed by
1634:
Aiello, Brett R.; Bhamla, M. Saad; Gau, Jeff; Morris, John G. L.; Bomar, Kenji; da Cunha, Shashwati; Fu, Harrison; Laws, Julia; Minoguchi, Hajime; Sripathi, Manognya; Washington, Kendra; Wong, Gabriella; Shubin, Neil H.; Sponberg, Simon; Stewart, Thomas A. (April 24, 2023).
596:
595:
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667:
To reduce toxic ammonia production, mudskippers can suppress amino acid breakdown when exposed to air. Another method they use involves the partial breakdown of amino acids leading to the production of alanine, which also reduces ammonia production.
288:, the males will also develop brightly coloured spots in order to attract females, which can be red, green or blue. Unlike other fish, the mudskipper's eyes protrude from the top of its flat head. Their most noticeable feature however is their side
594:
658:
when the fish and burrow are submerged, and lay their eggs. When the burrow is submerged, several mudskipper species maintain an air pocket inside it, which allows them to breathe in conditions of very low oxygen concentration.
338:
lineage", together with amblyopines. Mudskippers can be defined as oxudercine gobies that are "fully terrestrial for some portion of the daily cycle" (character 24 in Murdy, 1989). This would define the species of the genera
296:, which allow the mudskipper to crawl from place to place. Although having the typical body form of any other gobiid fish, these front fins allow the mudskipper to actively "skip" across muddy surfaces (hence the
518:
are elongated and protrude from the body wall. This unusual morphology creates a pectoral fin with two fin segments (the radials and the rays) and two movable hinge joints: a 'shoulder' joint where the
401:
occurs, the period of cohabitation between the male and female is rather short. Eventually, the female will leave and it is the male that ends up guarding the egg filled burrow from hungry predators.
593:
1850:
Polgar G, Zane L, Babbucci M, Barbisan F, Patarnello T, Rüber L, Papetti C (2014). "Phylogeography and demographic history of two widespread Indo-Pacific mudskippers (Gobiidae:
1986:
487:
Although mudskippers' fins do not have a joint homologous to the elbow, the joint between the radials and the fin rays serves a functionally analogous role.
671:
Mudskippers can reduce the membrane permeability of their skin and acidify the water in their burrows to reduce levels of ammonia from the environment.
308:. Mudskippers have also been found to be able to leap distances of up to 61 centimetres (24 in) by laterally flexing and pushing with their tails.
1278:
1325:
Sasekumar A, Chong VC, Lim KH, Singh HR (1994). "The Fish
Community of Matang Mangrove Waters, Malaysia". In Sudara S, Wilkinson CR, Chou LM (eds.).
913:
Agorreta A.; San Mauro D.; Schliewen U. & Rüber L. (2013). "Molecular phylogenetics of
Gobioidei and phylogenetic placement of European gobies".
786:
is in fact a complex of species, with at least three separate lineages, one in East Africa, and two in the Indo-Malayan region. Another species,
284:
Mudskippers can grow up to 30 cm (12 in) long, and most are a brownish green colour that ranges anywhere from dark to light. During
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superficialis muscle of the pectoral fin is divided into two sections (rather than being a single muscle, as is common with the rest of the
397:
choice she will then proceed to follow the male into the burrow where she will lay hundreds of eggs and allow them to be fertilized. After
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493:
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of water for long periods of time. It has been discovered that they spend up to three quarters of their life on land. They are found in
1248:"One size does not fit all: Variation in anatomical traits associated with emersion behavior in mudskippers (Gobiidae: Oxudercinae)"
2012:
1096:"Propulsive Forces of Mudskipper Fins and Salamander Limbs during Terrestrial Locomotion: Implications for the Invasion of Land"
711:
1579:
Randall, D. J.; Wilson, J. M.; Peng, K. W.; Kok, T. W. K.; Kuah, S. S. L.; Chew, S. F.; Lam, T. J.; Ip, Y. K. (1999-12-01).
1359:
Ishimatsu A, Hishida Y, Takita T, Kanda T, Oikawa S, Takeda T, Khoo KH (1998). "Mudskipper Store Air in Their
Burrows".
69:
1991:
579:
574:
524:
960:"Differentiation and adaptive radiation of amphibious gobies (Gobiidae: Oxudercinae) in semi-terrestrial habitats"
2050:
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that are located more forward and under their elongated body. These fins are jointed and function similarly to
1580:
788:
634:
1402:
Ishimatsu A, Takeda T, Kanda T, Oikawa S, Khoo KH (2000). "Burrow environment of mudskippers in
Malaysia".
1926:
1581:"The mudskipper, Periophthalmodon schlosseri, actively transports NH 4 + against a concentration gradient"
274:
1471:"Five Tropical Air‐Breathing Fishes, Six Different Strategies to Defend against Ammonia Toxicity on Land"
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621:
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2017:
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1911:
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49:
324:). Recent molecular studies do not support this classification, as oxudercine gobies appear to be
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meets the radials and an 'intra-fin' joint where the radials meet the rays. In addition, ... the
393:
225:
64:
1340:
Brillet C (1969). "Etude du comportement constructeur des poissons amphibies
Periophthalmidae".
1999:
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981:
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886:"A Taxonomic Revision and Cladistic Analysis of the Oxudercine Gobies (Gobiidae: Oxudercinae)"
832:
608:
Mudskippers have the ability to breathe through their skin and the lining of their mouth (the
357:
216:
1964:
2004:
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317:
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1526:"Air Breathing and Ammonia Excretion in the Giant Mudskipper, Periophthalmodon schlosseri"
705:
is by far the most diverse and widespread genus of mudskipper. Eighteen species have been
507:
377:
249:
204:
1741:(Teleostei: Gobiidae: Oxudercinae), a valid species of mudskipper, and a re-diagnosis of
1327:
Proceedings, Third ASEAN-Australia
Symposium on Living Coastal Resources. Research papers
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As their name implies, these fish use their fins to move around in a series of skips.
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1189:"Acoustic Communication at the Water's Edge: Evolutionary Insights from a Mudskipper"
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639:
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289:
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186:
1821:
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1637:"The origin of blinking in both mudskippers and tetrapods is linked to life on land"
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1694:(Teleostei: Gobiidae: Oxudercinae) from northern Australia, and a re-diagnosis of
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American
Journal of Physiology. Regulatory, Integrative and Comparative Physiology
41:
1213:
792:, is the only oxudercine goby that inhabits the coastal areas of western Africa.
642:, while others will eat small crabs, insects, snails and even other mudskippers.
475:
adaptations that allow them to move effectively on land as well as in the water.
281:, and the ability to survive prolonged periods of time both in and out of water.
1973:
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376:, and exhibit unique adaptations to this environment that are not found in most
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1792:(Teleostei: Gobiidae: Oxudercinae), from Australia, with a key to the genus".
1702:
The Beagle, Records of the
Museums and Art Galleries of the Northern Territory
1187:
Polgar G, Malavasi S, Cipolato G, Georgalas V, Clack JA, Torricelli P (2011).
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Compared with fully aquatic gobies, these specialised fish present a range of
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1949:
1604:
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1247:
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1137:"Mudskipper pectoral fin kinematics in aquatic and terrestrial environments"
1055:"Mudskipper pectoral fin kinematics in aquatic and terrestrial environments"
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1839:(unpubl. Ph.D. dissertation ed.). Brisbane: University of Queensland.
1943:
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Studies on the taxonomy, ecology and physiology of
Queensland mudskippers
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is one of the most widespread and well-known species. It can be found in
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445:
406:
101:
1524:
Randall, D. J.; Ip, Y. K.; Chew, S. F.; Wilson, J. M. (September 2004).
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suggest that this monotypic genus should be included in the definition.
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Richard van der Laan; William N. Eschmeyer & Ronald Fricke (2014).
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The different species have adapted to various diets on the mudflats.
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1329:. Vol. 2. Bangkok: Chulalonghorn University. pp. 457–464.
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Lee HJ, Martinez CA, Hertzberg KJ, Hamilton AL, Graham JB (2005).
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557:
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1641:
1423:"Burrow air phase maintenance and respiration by the mudskipper
1290:
Tytler P, Vaughan T (1983). "Thermal
Ecology of the Mudskippers
655:
381:
321:
269:. They are known for their unusual body shapes, preferences for
260:
1924:
1902:
852:"Classification of fishes from Fishes of the World 5th Edition"
770:
Islands. It grows to a length of about 9.5 cm and is a
1906:
1897:
1788:
Jaafar Z, Larson HL (2008). "A new species of mudskipper,
650:
Digging deep burrows in soft sediments allows the fish to
1006:
Air–breathing Fishes. Evolution, Diversity and Adaptation
774:
opportunist feeder. It feeds on small prey such as small
1022:
Harris VA (1960). "On the locomotion of the mudskipper
1469:
Ip, Y. K.; Chew, S. F.; Randall, D. J. (2004-09-01).
1933:
782:. However, a recent molecular study suggests that
850:Nelson, JS; Grande, TC & Wilson, MVH (2016).
355:as "mudskippers". However, field observations of
1028:Proceedings of the Zoological Society of London
958:Polgar G.; Sacchetti A. & Galli P. (2010).
500:
485:
316:Oxudercinae is sometimes classified within the
1464:
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380:, which typically survive the retreat of the
8:
1017:
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999:
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604:territorial defense call and jumping ability
535:rays and the other section inserting on the
1094:S. M. Kawano; R. W. Blob (August 2, 2013).
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531:gobies) with one section inserting on the
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654:, avoid marine predators during the high
369:Mudskippers typically live in burrows in
328:relative to amblyopine gobies (Gobiidae:
1903:The Mudskipper: a website on mudskippers
1279:Ecology of Kalimantan: Indonesian Borneo
1008:. San Diego California: Academic Press.
801:
332:), thus being included in a distinct "
1856:Molecular Phylogenetics and Evolution
1737:Jaafar Z, Perrig M, Chou LM (2009). "
1530:Physiological and Biochemical Zoology
1475:Physiological and Biochemical Zoology
915:Molecular Phylogenetics and Evolution
811:"Family-group names of Recent fishes"
7:
248:are any of the 23 extant species of
1431:The Journal of Experimental Biology
1144:The Journal of Experimental Biology
1100:Integrative and Comparative Biology
679:Mudskippers evolved the ability to
546:The Journal of Experimental Biology
494:Integrative and Comparative Biology
1312:10.1111/j.1095-8649.1983.tb02912.x
1252:Frontiers in Ecology and Evolution
1135:Pace CM, Gibb AC (July 15, 2009).
1040:10.1111/j.1469-7998.1960.tb05921.x
25:
1597:10.1152/ajpregu.1999.277.6.R1562
977:10.1111/j.1095-8649.2010.02807.x
890:Records of the Australian Museum
68:
1696:Periophthalmus novaeguineaensis
1059:Journal of Experimental Biology
602:Periophthalmodon septemradiatus
899:10.3853/j.0812-7387.11.1989.93
712:Periophthalmus argentilineatus
1:
300:) and even climb low-hanging
1743:Periophthalmus novemradiatus
1688:Larson HK, Takita T (2004).
1214:10.1371/journal.pone.0021434
1868:10.1016/j.ympev.2014.01.014
927:10.1016/j.ympev.2013.07.017
2077:
1898:Gobioid Research Institute
1893:FishBase entry on Gobiidae
1298:(Pallas), of Kuwait Bay".
1292:Periophthalmus koelreuteri
1246:Corush J, Zhang J (2022).
1024:Periophthalmus koelreuteri
580:Coringa Mangrove Sanctuary
1739:Periophthalmus variabilis
1053:Pace CM, Gibb AC (2009).
828:10.11646/zootaxa.3882.1.1
231:
224:
159:
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65:Scientific classification
63:
48:
39:
34:
1427:(Gobiidae: Oxudercinae)"
1296:Boleophthalmus boddaerti
1265:10.3389/fevo.2022.967067
1656:10.1073/pnas.2220404120
1300:Journal of Fish Biology
1004:Graham JB, ed. (1997).
964:Journal of Fish Biology
789:Periophthalmus barbarus
635:Boleophthalmus boddarti
461:Periophthalmus barbarus
442:Periophthalmus gracilis
417:regions, including the
54:Periophthalmus barbarus
1425:Scartelaos histophorus
1404:Journal of Biosciences
605:
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571:
551:
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275:terrestrial locomotion
1790:Periophthalmus takita
1026:(Pallas): Gobiidae".
622:cutaneous respiration
600:
577:
561:
463:(from western Africa)
459:
440:
1835:Milward, NE (1974).
1690:"Two new species of
479:Terrestrial movement
384:by hiding under wet
1649:(18): e2220404120.
1373:1998Natur.391..237I
1205:2011PLoSO...621434P
510:fishes in that the
50:Atlantic mudskipper
27:Subfamily of fishes
1806:10.2108/zsj.25.946
1794:Zoological Science
1759:10.2108/zsj.26.309
1747:Zoological Science
1591:(6): R1562–R1567.
1157:10.1242/jeb.029041
1113:10.1093/icb/ict051
1072:10.1242/jeb.029041
892:. Suppl 11: 1–93.
784:P. argentilineatus
748:Northern Australia
663:Ammonia regulation
612:) and throat (the
606:
588:
572:
514:of the mudskipper
506:differs from most
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273:habitats, limited
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2000:Open Tree of Life
1927:Taxon identifiers
1912:Mudskipper photos
1444:10.1242/jeb.01361
1437:(Pt 1): 169–177.
1367:(6664): 237–238.
1258:: 967067–967075.
1150:(14): 2279–2286.
1065:(14): 2279–2286.
884:Murdy EO (1989).
734:east through the
686:from terrestrial
598:
429:coast of Africa.
378:intertidal fishes
358:Zappa confluentus
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502:The mudskipper
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1887:External links
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1852:Periophthalmus
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1827:
1800:(9): 946–952.
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1753:(4): 309–314.
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1692:Periophthalmus
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1626:
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1542:10.1086/423745
1536:(5): 783–788.
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1487:10.1086/422057
1481:(5): 768–782.
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744:Southeast Asia
703:Periophthalmus
695:
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673:
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652:thermoregulate
647:
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629:
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620:, is known as
584:Andhra Pradesh
578:Mudskipper at
564:Point Calimere
562:Mudskipper at
555:
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541:
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434:
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349:Periophthalmus
341:Boleophthalmus
335:Periophthalmus
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286:mating seasons
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199:Periophthalmus
175:Boleophthalmus
157:
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139:
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119:
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112:Actinopterygii
109:
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1410:(1–2): 17–28.
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1381:10.1038/34560
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1348:(4): 496–520.
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684:independently
682:
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640:detritivorous
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399:fertilization
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302:tree branches
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187:Parapocryptes
183:
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148:
143:
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77:
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59:
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47:
43:
38:
33:
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2061:Walking fish
1934:
1859:
1855:
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1836:
1830:
1797:
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1789:
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1750:
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1103:
1099:
1089:
1062:
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1031:
1027:
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963:
953:
936:10261/123985
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787:
783:
750:, southeast
710:
702:
697:
678:
670:
666:
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633:
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607:
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544:
516:pectoral fin
504:pectoral fin
501:
492:
486:
482:
466:
460:
448:to northern
441:
403:
368:
356:
352:
348:
344:
340:
333:
326:paraphyletic
315:
283:
256:
245:
244:
215:
209:
203:
197:
191:
185:
179:
173:
169:Apocryptodon
167:
161:
160:
141:
53:
29:
2056:Oxudercinae
2046:Gobiiformes
2041:Mudskippers
1974:iNaturalist
1965:Oxudercinae
1959:Wikispecies
1935:Oxudercinae
1862:: 161–176.
1708:: 175–185.
1034:: 107–135.
772:carnivorous
728:East Africa
566:Sanctuary,
529:Oxudercinae
473:ethological
433:Adaptations
411:subtropical
394:territories
330:Amblyopinae
298:common name
271:semiaquatic
267:Oxudercidae
257:Oxudercinae
246:Mudskippers
142:Oxudercinae
138:Subfamily:
132:Oxudercidae
122:Gobiiformes
35:Mudskipper
18:Oxudercinae
2035:Categories
796:References
780:arthropods
778:and other
758:, and the
736:Sundarbans
732:Madagascar
720:ecosystems
618:amphibians
568:Tamil Nadu
469:anatomical
390:tide pools
371:intertidal
353:Scartelaos
320:Gobiidae (
211:Scartelaos
163:Apocryptes
58:The Gambia
1724:131712323
1605:0363-6119
1550:1522-2152
1495:1522-2152
707:described
688:tetrapods
646:Burrowing
554:Breathing
521:cleithrum
450:Australia
415:temperate
365:Behaviour
254:subfamily
252:from the
181:Oxuderces
88:Kingdom:
82:Eukaryota
1944:Wikidata
1876:24486991
1822:10659541
1814:19267605
1775:31647216
1767:19798926
1675:37094121
1666:10160996
1621:29598011
1613:10600900
1566:36864021
1558:15547796
1511:20545085
1503:15547795
1453:15601887
1233:21738663
1193:PLOS ONE
1174:18176905
1166:19561218
1122:23667046
1081:19561218
986:21078024
945:23911892
837:25543675
724:mudflats
717:mangrove
675:Blinking
542:—
525:abductor
490:—
446:Malaysia
427:Atlantic
425:and the
407:tropical
374:habitats
312:Taxonomy
226:Synonyms
128:Family:
102:Chordata
98:Phylum:
92:Animalia
78:Domain:
1950:Q828079
1389:4429394
1369:Bibcode
1224:3125184
1201:Bibcode
815:Zootaxa
760:Ryukyus
694:Species
614:pharynx
586:, India
570:, India
537:ventral
512:radials
423:Pacific
386:seaweed
279:jumping
259:of the
155:Genera
147:Gunther
118:Order:
108:Class:
2018:267096
2005:897333
1992:497678
1979:125852
1916:Flickr
1874:
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1361:Nature
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860:10 May
835:
756:Taiwan
740:Bengal
610:mucosa
533:dorsal
444:(from
413:, and
388:or in
351:, and
322:gobies
318:family
306:scrubs
264:family
2013:WoRMS
1818:S2CID
1771:S2CID
1720:S2CID
1617:S2CID
1562:S2CID
1507:S2CID
1385:S2CID
1170:S2CID
1140:(PDF)
855:(PDF)
776:crabs
768:Tonga
764:Samoa
762:, to
752:China
700:genus
681:blink
539:rays.
294:limbs
217:Zappa
56:) in
1987:NCBI
1872:PMID
1854:)".
1810:PMID
1763:PMID
1671:PMID
1642:PNAS
1609:PMID
1601:ISSN
1554:PMID
1546:ISSN
1499:PMID
1491:ISSN
1449:PMID
1229:PMID
1162:PMID
1118:PMID
1077:PMID
982:PMID
941:PMID
862:2018
833:PMID
819:3882
766:and
730:and
722:and
698:The
656:tide
628:Diet
471:and
419:Indo
382:tide
304:and
277:and
261:goby
149:1861
1914:on
1905:or
1864:doi
1802:doi
1755:doi
1745:".
1710:doi
1661:PMC
1651:doi
1647:120
1593:doi
1589:277
1538:doi
1483:doi
1439:doi
1435:208
1377:doi
1365:391
1308:doi
1260:doi
1219:PMC
1209:doi
1152:doi
1148:212
1108:doi
1067:doi
1063:212
1036:doi
1032:134
972:doi
931:hdl
923:doi
894:doi
823:doi
746:to
738:of
726:of
638:is
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