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The '''Atlantic mudskipper''' (''Periophthalmus barbarus'') is a [[species]] of [[mudskipper]] native to fresh, marine and brackish waters of the [[tropical]] [[Atlantic]] coasts of [[Africa]], including most offshore islands, through the [[Indian Ocean]] and into the western [[Pacific Ocean]] to [[Guam]].<ref name=Physicochemical>{{cite journal|last=Emuebie|first=Okonji Raphael|year=2011|title=Physicochemical Properties of Mudskipper ( Periophthalmus Barbarus Pallas) Liver Rhodanese.|journal=Australian Journal of Basic and Applied Sciences|volume=5|issue=8|pages=507–514}}</ref> This species occurs on the tidal flats of [[mangrove forest]]s where it readily crosses mud and sand surfaces out of the water, using its pectoral fins to move.
The '''Atlantic mudskipper''' (''Periophthalmus barbarus'') is a [[species]] of [[mudskipper]] native to fresh, marine and brackish waters of the [[tropical]] [[Atlantic]] coasts of [[Africa]], including most offshore islands, through the [[Indian Ocean]] and into the western [[Pacific Ocean]] to [[Guam]].<ref name=Physicochemical>{{cite journal|last=Emuebie|first=Okonji Raphael|year=2011|title=Physicochemical Properties of Mudskipper ( Periophthalmus Barbarus Pallas) Liver Rhodanese.|journal=Australian Journal of Basic and Applied Sciences|volume=5|issue=8|pages=507–514}}</ref> This species occurs on the tidal flats of [[mangrove forest]]s where it readily crosses mud and sand surfaces out of the water, using its pectoral fins to move.

== Anatomy and morphology ==
The Atlantic mudskipper can grow up to 16cm in [[length]].<ref name=":0">{{Cite web|title=Atlantic mudskippers: Meet them at Zoo Leipzig!|url=https://www.zoo-leipzig.de/en/animal/atlantic-mudskipper/|access-date=2020-10-07|website=Zoo Leipzig|language=en}}</ref> The body is covered with [[Scale (anatomy)|scales]], coated with a [[mucus]] layer that helps to retain [[moisture]].<ref name=":0" /> The Atlantic mudskippers have more than 90 scales along the side of its body.<ref name=":12">{{Cite book|last=Murdy|first=Edward O.|url=https://books.google.com.au/books/about/A_Taxonomic_Revision_and_Cladistic_Analy.html?id=t1yLQgAACAAJ&redir_esc=y|title=A Taxonomic Revision and Cladistic Analysis of the Oxudercine Gobies (Gobiidae, Oxudercinae)|date=1989|publisher=Australian Museum|isbn=978-0-7305-6374-7|language=en}}</ref> Atlantic mudskippers also retain [[moisture]] by storing water within [[gill]] chambers, that allows them to breathe when out of water.<ref name=":0" /> In addition to retaining water, the surface of the Atlantic mudskipper which enables them to breathe through their skin, otherwise known as [[cutaneous respiration]].<ref name=":2">{{Cite journal|last=Kutschera|first=Ulrich|last2=Elliott|first2=J. Malcolm|date=2013-03-26|title=Do mudskippers and lungfishes elucidate the early evolution of four-limbed vertebrates?|url=https://doi.org/10.1186/1936-6434-6-8|journal=Evolution: Education and Outreach|volume=6|issue=1|pages=8|doi=10.1186/1936-6434-6-8|issn=1936-6434}}</ref>

Mudskippers have a pair of [[Pectoral muscles|pectoral]] [[Fin|fins]], which allows them to 'skip' on land and maintain stability within water.<ref name=":0" /><ref name=":32">{{Cite web|last=Ansari|first=A.|last2=Trivedi|first2=S.|last3=Saggu|first3=Shalini|last4=Rehman|first4=H.|date=2014|title=Mudskipper: A biological indicator for environmental monitoring and assessment of coastal waters|url=https://www.semanticscholar.org/paper/Mudskipper%3A-A-biological-indicator-for-monitoring-Ansari-Trivedi/4e393c91ef8e3959b7fb557af96c00ed75e07018|access-date=2020-10-07|website=undefined|language=en}}</ref><ref name=":4">{{Cite journal|last=Pace|first=C. M.|last2=Gibb|first2=A. C.|date=2009-07-15|title=Mudskipper pectoral fin kinematics in aquatic and terrestrial environments|url=https://jeb.biologists.org/content/212/14/2279|journal=Journal of Experimental Biology|language=en|volume=212|issue=14|pages=2279–2286|doi=10.1242/jeb.029041|issn=0022-0949|pmid=19561218}}</ref> The Atlantic mudskipper also has a pair of caudal fins that aid in [[aquatic locomotion]], and [[Pelvis|pelvic]] fins that aid the pectoral fins in [[terrestrial locomotion]].<ref name=":4" /> The pelvic fins are adapted to terrestrial-living by acting as a sucker to attach the Atlantic mudskipper to land.<ref name=":2" /> Atlantic mudskippers can also crawl and climb on land using their pelvic and pectoral fins.<ref name=":2" />

The [[Eye|eyes]] of the Atlantic mudskipper are adapted to terrestrial-living by being located closely together, providing the mudskipper with a large [[Visual field|field of vision]].<ref name=":32" /> The eyes can move independent of one another at 360 degrees.<ref name=":5">{{Cite web|title=Atlantic mudskipper|url=https://www.zoo.ch/en/atlantic-mudskipper|access-date=2020-10-11|website=Zoo Zürich|language=en}}</ref> The eyes are also positioned further up on the head, enabling the eyes to remain above the water surface whilst their body is submerged [[Underwater environment|underwater]].<ref name=":32" /> Cup-like structures that hold water are located beneath the eye, which aids in [[Lubrication|lubricating]] the eyes when the Atlantic mudskipper is on land.<ref name=":32" />

Atlantic mudskippers have a [[mouth]],<ref name=":32" /> which can be reorientated. The Atlantic mudskipper has the ability to [[Rotation|rotate]] its mouth opening so that their jaws can be oriented over prey.<ref>{{Cite journal|last=Michel|first=Krijn B.|last2=Adriaens|first2=Dominique|last3=Aerts|first3=Peter|last4=Dierick|first4=Manuel|last5=Wassenbergh|first5=Sam Van|date=2014|title=Functional anatomy and kinematics of the oral jaw system during terrestrial feeding in Periophthalmus barbarus|url=https://onlinelibrary.wiley.com/doi/abs/10.1002/jmor.20318|journal=Journal of Morphology|language=en|volume=275|issue=10|doi=10.1002/jmor.20318|issn=1097-4687}}</ref> Sharp [[Tooth|teeth]], reflective of their [[Carnivore|carnivorous]] diet, are present within the mouth.<ref>{{Cite journal|last=Sponder|first=Debra L.|last2=Lauder|first2=G.|date=2009|title=Terrestrial feeding in the Mudskipper Periophthalmus (Pisces: Teleostei): A cineradiographic analysis|url=https://www.semanticscholar.org/paper/Terrestrial-feeding-in-the-Mudskipper-(Pisces:-A-Sponder-Lauder/06ab53968782cf50a50ec0423715706d24925a58|doi=10.1111/J.1469-7998.1981.TB01501.X}}</ref> Atlantic mudskippers have a short [[Gastrointestinal tract|digestive system]], that is composed of an [[Esophagus|oesophagus]], [[stomach]], [[Gastrointestinal tract|intestine]], and [[rectum]].<ref>{{Cite journal|last=Wołczuk|first=Katarzyna|last2=Ostrowski|first2=Maciej|last3=Ostrowska|first3=Agnieszka|last4=Napiórkowska|first4=Teresa|date=2018-06-01|title=Structure of the alimentary tract in the Atlantic mudskipper Periophthalmus barbarus (Gobiidae: Oxudercinae): anatomical, histological and ultrastructural studies|url=http://www.sciencedirect.com/science/article/pii/S0944200618300060|journal=Zoology|language=en|volume=128|pages=38–45|doi=10.1016/j.zool.2018.04.002|issn=0944-2006}}</ref>

The Atlantic mudskipper has a unique [[Olfaction|olfactory]] [[Organ (anatomy)|organ]] that includes a canal 0.3mm in diameter near its upper [[lip]] that increases in size into a chamber-like sac. The chamber-like sacs only serve a mechanical purpose, which is to circulate water through the canals which are sensory structures.<ref>{{Cite journal|last=Kuciel|first=Michał|date=2013-03-01|title=The mechanism of olfactory organ ventilation in Periophthalmus barbarus (Gobiidae, Oxudercinae)|url=https://doi.org/10.1007/s00435-012-0167-y|journal=Zoomorphology|language=en|volume=132|issue=1|pages=81–85|doi=10.1007/s00435-012-0167-y|issn=1432-234X|pmc=PMC3570760|pmid=23420290}}</ref><ref>{{Cite journal|last=Kuciel|first=Michał|last2=Żuwała|first2=Krystyna|last3=Jakubowski|first3=Michał|date=2011|title=A new type of fish olfactory organ structure in Periophthalmus barbarus (Oxudercinae)|url=https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1463-6395.2010.00459.x|journal=Acta Zoologica|language=en|volume=92|issue=3|pages=276–280|doi=10.1111/j.1463-6395.2010.00459.x|issn=1463-6395}}</ref>

The Atlantic mudskippers have [[Genital papilla|genital papillae]] that are located on the [[abdomen]]. Females can be distinguished from males, who have less rounded papillae.<ref>{{Cite journal|last=Udo|first=M.|last2=Abiaobo|first2=No|last3=Asuquo|first3=I.|date=2016|title=Aspects of the reproductive biology in mudskippers Periophthalmus barbarus ( Gobiidae ) ( Linneaus 1766 ) in mangrove swamps of Iko Estuary , Southeast , Nigeria|url=https://www.semanticscholar.org/paper/Aspects-of-the-reproductive-biology-in-mudskippers-Udo-Abiaobo/6fddc9ca8531e666c3c5a0e474976492db0a49d3}}</ref>


== Etymology ==
== Etymology ==
For the scientific name ''Periophthalmus barbarus,'' in [[Greek language|Greek]] ''<nowiki/>'<nowiki/>''Peri''<nowiki/>'<nowiki/>'' means 'around', and 'ophthalmos''<nowiki/>''' means 'eye',<ref name=":1">{{Cite book|last=Murdy|first=Edward O.|url=https://books.google.com/books?id=t1yLQgAACAAJ|title=A Taxonomic Revision and Cladistic Analysis of the Oxudercine Gobies (Gobiidae, Oxudercinae)|date=1989|publisher=Australian Museum|isbn=978-0-7305-6374-7|language=en}}</ref> in reference to the Atlantic mudskipper's close set eyes that provide them with a larger field of vision.<ref name=":3">{{Cite journal|last1=Ansari|first1=A.|last2=Trivedi|first2=S.|last3=Saggu|first3=Shalini|last4=Rehman|first4=H.|date=2014|title=Mudskipper: A biological indicator for environmental monitoring and assessment of coastal waters|url=https://www.semanticscholar.org/paper/Mudskipper%3A-A-biological-indicator-for-monitoring-Ansari-Trivedi/4e393c91ef8e3959b7fb557af96c00ed75e07018|access-date=2020-10-07|website=undefined|s2cid=54816035|language=en}}</ref> In Greek, 'barbarus' means 'foreign', potentially named after the many differences it displayed when compared to other gobies.<ref name=":1" /> The generic name 'mudskipper', is labelled after their 'skipping' movement on [[Mudflat|mudflats]].<ref>{{Cite web|title=Definition of mudskipper {{!}} Dictionary.com|url=https://www.dictionary.com/browse/mudskipper|access-date=2020-10-07|website=www.dictionary.com|language=en}}</ref>
For the scientific name ''Periophthalmus barbarus,'' in [[Greek language|Greek]] ''<nowiki/>'<nowiki/>''Peri''<nowiki/>'<nowiki/>'' means 'around', and 'ophthalmos''<nowiki/>''' means 'eye',<ref name=":1">{{Cite book|last=Murdy|first=Edward O.|url=https://books.google.com/books?id=t1yLQgAACAAJ|title=A Taxonomic Revision and Cladistic Analysis of the Oxudercine Gobies (Gobiidae, Oxudercinae)|date=1989|publisher=Australian Museum|isbn=978-0-7305-6374-7|language=en}}</ref> in reference to the Atlantic mudskipper's close set eyes that provide them with a larger field of vision.<ref name=":3">{{Cite journal|last1=Ansari|first1=A.|last2=Trivedi|first2=S.|last3=Saggu|first3=Shalini|last4=Rehman|first4=H.|date=2014|title=Mudskipper: A biological indicator for environmental monitoring and assessment of coastal waters|url=https://www.semanticscholar.org/paper/Mudskipper%3A-A-biological-indicator-for-monitoring-Ansari-Trivedi/4e393c91ef8e3959b7fb557af96c00ed75e07018|access-date=2020-10-07|website=undefined|s2cid=54816035|language=en}}</ref> In Greek, 'barbarus' means 'foreign', potentially named after the many differences it displayed when compared to other gobies.<ref name=":1" /> The generic name 'mudskipper', is labelled after their 'skipping' movement on [[Mudflat|mudflats]].<ref>{{Cite web|title=Definition of mudskipper {{!}} Dictionary.com|url=https://www.dictionary.com/browse/mudskipper|access-date=2020-10-07|website=www.dictionary.com|language=en}}</ref>


==Distribution==
== Taxonomy ==
The initial identification and description of the species was outlined by [[Carl Linnaeus|Carl Linneaus]], published in 1766 as ''[[Gobius]] barbarus'' and renamed to ''[[Periophthalmus]] barbarus''.<ref>{{Cite web|title=WoRMS - World Register of Marine Species - Periophthalmus barbarus (Linnaeus, 1766)|url=http://www.marinespecies.org/aphia.php?p=taxdetails&id=277973|access-date=2020-10-07|website=www.marinespecies.org}}</ref> The Atlantic mudskipper has also previously been known or misidentified as ''Gobius koelreuteri'', ''Periophthalmus papilio'', ''Periophthalmus gabonicus'', ''Periophthalmus errythronemus''.<ref name=":13">{{Cite book|last=Murdy|first=Edward O.|url=https://books.google.com.au/books/about/A_Taxonomic_Revision_and_Cladistic_Analy.html?id=t1yLQgAACAAJ&redir_esc=y|title=A Taxonomic Revision and Cladistic Analysis of the Oxudercine Gobies (Gobiidae, Oxudercinae)|date=1989|publisher=Australian Museum|isbn=978-0-7305-6374-7|language=en}}</ref> The Atlantic mudskipper is distinguished from other members of ''Periophthalmus'' by having no spots or having some white spots on its back.<ref name=":13" />

== Distribution ==
''Periophthalmus barbarus'' is found throughout West Africa, in mangrove swamps and primarily brackish bodies of water near the coast.<ref name=iucn/> Countries where the mudskipper are found include [[Angola]], the [[Democratic Republic of the Congo]], [[Cameroon]] and Ghana <ref name=iucn/>
''Periophthalmus barbarus'' is found throughout West Africa, in mangrove swamps and primarily brackish bodies of water near the coast.<ref name=iucn/> Countries where the mudskipper are found include [[Angola]], the [[Democratic Republic of the Congo]], [[Cameroon]] and Ghana <ref name=iucn/>

== Ecology and behaviour ==

=== Habitat ===
Atlantic mudskippers are [[Semiaquatic|semi-aquatic]] animals that live in areas with water that is [[Brackish water|slightly salty]], such as [[Estuary|river estuaries]] and [[Mudflat|mudflats]].<ref name=":52">{{Cite web|title=Atlantic mudskipper|url=https://www.zoo.ch/en/atlantic-mudskipper|access-date=2020-10-11|website=Zoo Zürich|language=en}}</ref><ref name=":6">{{Cite web|title=Periophthalmus barbarus (Atlantic Mudskipper) — Seriously Fish|url=https://www.seriouslyfish.com/species/periophthalmus-barbarus/|access-date=2020-10-07}}</ref> Generally, Atlantic mudskippers spend the majority of the day on land.<ref name=":52" /> In tidal regions, Atlantic mudskippers may appear only during low [[tide]] to feed; whereas, they hide in their [[burrow]] at high tide.<ref name=":6" />

=== Predator-prey behaviour ===
The Atlantic mudskipper is [[Carnivore|carnivorous]],<ref>{{Cite journal|last=Clayton|first=David|date=1993|title=Mudskippers|url=https://www.academia.edu/7148041/Mudskippers_Literature_Review_|journal=Oceanography and Marine Biology|language=en|volume=31|pages=507-577|via=}}</ref> and has adopted an [[Ambush predator|ambushing strategy]] to capture terrestrial prey.<ref name=":7">{{Cite journal|last=Kutschera|first=U.|last2=Burghagen|first2=H.|last3=Ewert|first3=J.|date=2008|title=Prey-Catching Behaviour in Mudskippers and Toads: A Comparative Analysis|url=https://www.semanticscholar.org/paper/Prey-Catching-Behaviour-in-Mudskippers-and-Toads:-A-Kutschera-Burghagen/bd24664dc0a451662a126f37cc09b1f861eecf2f|doi=10.3844/OJBSCI.2008.41.43}}</ref> To detect prey, the Atlantic mudskipper submerges itself underwater whilst leaving its eyes out, using only [[Visual perception|sight]] to identify and locate prey.<ref name=":7" /> To ambush prey, Atlantic mudskippers launch onto land using predominantly their pectoral fins, and catch the prey using their mouth.<ref name=":7" />

=== Feeding behaviour ===
On land, the Atlantic mudskipper feeds by covering their prey with water, then sucking back the water and prey into their mouth, named as the 'hydrodynamic tongue'.<ref>{{Cite journal|last=Michel|first=Krijn B.|last2=Heiss|first2=Egon|last3=Aerts|first3=Peter|last4=Van Wassenbergh|first4=Sam|date=2015-04-22|title=A fish that uses its hydrodynamic tongue to feed on land|url=https://royalsocietypublishing.org/doi/10.1098/rspb.2015.0057|journal=Proceedings of the Royal Society B: Biological Sciences|volume=282|issue=1805|pages=20150057|doi=10.1098/rspb.2015.0057}}</ref> In water, the Atlantic mudskipper feeds through [[Aquatic feeding mechanisms|suction feeding]], similar to other aquatic species.<ref name=":8">{{Cite journal|last=Michel|first=K. B.|last2=Aerts|first2=P.|last3=Wassenbergh|first3=S. Van|date=2016-11-15|title=Environment-dependent prey capture in the Atlantic mudskipper (Periophthalmus barbarus)|url=https://bio.biologists.org/content/5/11/1735|journal=Biology Open|language=en|volume=5|issue=11|pages=1735–1742|doi=10.1242/bio.019794|issn=2046-6390|pmid=27765755}}</ref> Although the feeding technique is similar in both environments, the Atlantic mudskipper alters the force of suction, such that the flow underwater is stronger than on land.<ref name=":8" /> The Atlantic mudskippers are [[Diurnality|diurnal]], which means they are active and feed during the day.<ref name=":52" />

Atlantic mudskippers are flexible in regard to their diet choices. Larger Atlantic mudskippers ingest larger sized prey, potentially due to the correlation between their mouth gape and prey size. The feeding choices also may vary between [[Habitat|habitats]] and [[Season|seasonally]], depending on what [[Resource|resources]] are most abundant. The Atlantic mudskippers feed more during [[dry season]] than [[Wet season|wet]], reflective of [[optimal foraging theory]], which proposes that diet flexibility increases with lower food availability. Atlantic mudskippers feed frequently, where larger Atlantic mudskippers feed at higher intensities as they are less prone to predation.<ref>{{Cite journal|last=Udo|first=Mfon T.|title=Tropic attributes of the mudskipper, Periophthalmus barbarus (Gobiidae: Oxudercinae) in the mangrove swamps of Imo River estuary, Nigeria|url=http://www.jesc.ac.cn/jesc_en/ch/reader/view_abstract.aspx?file_no=20020412|language=en|volume=14|issue=4|pages=508–517}}</ref>

== Reproduction ==
Females reach [[sexual maturity]] at a body length of around 10.2cm, and males approximately 10.8cm. The Atlantic mudskipper can [[Spawn (biology)|spawn]] throughout the year.<ref name=":9">{{Cite journal|last=Chukwu|first=K. O.|last2=Deekae|first2=S. N.|last3=Gabriel|first3=U. U.|date=2010|title=Reproductive biology of Periopthalmus barbarus (Linneaus 1766) in New Calabar River, Nigeria.|url=https://www.semanticscholar.org/paper/Reproductive-biology-of-Periopthalmus-barbarus-in-Chukwu-Deekae/7c9ad0ca7569f95d8276fa9fc34099f168e37292|doi=10.5251/ABJNA.2010.1.6.1158.1161}}</ref> However, spawning mainly occurs for male Atlantic mudskippers between February and May, and females between March and May.<ref name=":10">{{Cite journal|last=Etim|first=Lawrence|last2=King|first2=Richard P.|last3=Udo|first3=Mfon T.|date=2002-06-01|title=Breeding, growth, mortality and yield of the mudskipper Periophthalmus barbarus (Linneaus 1766) (Teleostei: Gobiidae) in the Imo River estuary, Nigeria|url=http://www.sciencedirect.com/science/article/pii/S0165783601003277|journal=Fisheries Research|language=en|volume=56|issue=3|pages=227–238|doi=10.1016/S0165-7836(01)00327-7|issn=0165-7836}}</ref> The peak spawning times are associated with high food availability for their young.<ref name=":10" /> The [[fertility]] of Atlantic mudskippers increases with the length of their body, as females may be able to carry more eggs at larger sizes.<ref name=":9" /> Females lay thousands of eggs at one time; however, the eggs are highly prone to predation.<ref name=":33">{{Cite web|last=Ansari|first=A.|last2=Trivedi|first2=S.|last3=Saggu|first3=Shalini|last4=Rehman|first4=H.|date=2014|title=Mudskipper: A biological indicator for environmental monitoring and assessment of coastal waters|url=https://www.semanticscholar.org/paper/Mudskipper%3A-A-biological-indicator-for-monitoring-Ansari-Trivedi/4e393c91ef8e3959b7fb557af96c00ed75e07018|access-date=2020-10-07|website=undefined|language=en}}</ref>

During [[courtship]], after the male pairs with a female mudskipper, the male and female go into the male's burrow to [[Copulation (zoology)|mate]].<ref name=":9" /> After the female releases her eggs onto the burrow wall,<ref name=":9" /> the male displays a large amount of [[paternal care]].<ref name=":33" /> The male Atlantic mudskipper will guard and take care of the eggs within the burrow.<ref name=":33" />

=== Lifecycle ===
The average [[Life expectancy|lifespan]] of an Atlantic mudskipper is approximately five years. The egg laid by the mother in the burrow, will hatch into [[Larva|larvae]] that swim out of the burrow and drift for 30-50 days. As Atlantic mudskippers are territorial, [[Juvenile fish|juvenile]] mudskippers will hide in mud until they mature enough to protect their established territory.<ref name=":33" />


==Usage by humans==
==Usage by humans==

Revision as of 02:54, 3 November 2020

Atlantic mudskipper
Periophthalmus barbatus from Gambia
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Gobiiformes
Family: Oxudercidae
Genus: Periophthalmus
Species:
P. barbarus
Binomial name
Periophthalmus barbarus
(Linnaeus, 1766)
Synonyms
  • Gobius barbarus Linnaeus, 1766
  • Gobius koelreuteri Pallas, 1770
  • Periophtalmus koelreuteri (Pallas, 1770)
  • Periophthalmus koelreuteri (Pallas, 1770)
  • Periophthalmus papilio Bloch & Schneider, 1801
  • Periophthalmus koelreuteri papilio Bloch & Schneider, 1801
  • Periophthalmus gabonicus Duméril, 1861
  • Periophthalmus erythronemus Guichenot, 1858

The Atlantic mudskipper (Periophthalmus barbarus) is a species of mudskipper native to fresh, marine and brackish waters of the tropical Atlantic coasts of Africa, including most offshore islands, through the Indian Ocean and into the western Pacific Ocean to Guam.[2] This species occurs on the tidal flats of mangrove forests where it readily crosses mud and sand surfaces out of the water, using its pectoral fins to move.

Anatomy and morphology

The Atlantic mudskipper can grow up to 16cm in length.[3] The body is covered with scales, coated with a mucus layer that helps to retain moisture.[3] The Atlantic mudskippers have more than 90 scales along the side of its body.[4] Atlantic mudskippers also retain moisture by storing water within gill chambers, that allows them to breathe when out of water.[3] In addition to retaining water, the surface of the Atlantic mudskipper which enables them to breathe through their skin, otherwise known as cutaneous respiration.[5]

Mudskippers have a pair of pectoral fins, which allows them to 'skip' on land and maintain stability within water.[3][6][7] The Atlantic mudskipper also has a pair of caudal fins that aid in aquatic locomotion, and pelvic fins that aid the pectoral fins in terrestrial locomotion.[7] The pelvic fins are adapted to terrestrial-living by acting as a sucker to attach the Atlantic mudskipper to land.[5] Atlantic mudskippers can also crawl and climb on land using their pelvic and pectoral fins.[5]

The eyes of the Atlantic mudskipper are adapted to terrestrial-living by being located closely together, providing the mudskipper with a large field of vision.[6] The eyes can move independent of one another at 360 degrees.[8] The eyes are also positioned further up on the head, enabling the eyes to remain above the water surface whilst their body is submerged underwater.[6] Cup-like structures that hold water are located beneath the eye, which aids in lubricating the eyes when the Atlantic mudskipper is on land.[6]

Atlantic mudskippers have a mouth,[6] which can be reorientated. The Atlantic mudskipper has the ability to rotate its mouth opening so that their jaws can be oriented over prey.[9] Sharp teeth, reflective of their carnivorous diet, are present within the mouth.[10] Atlantic mudskippers have a short digestive system, that is composed of an oesophagus, stomach, intestine, and rectum.[11]

The Atlantic mudskipper has a unique olfactory organ that includes a canal 0.3mm in diameter near its upper lip that increases in size into a chamber-like sac. The chamber-like sacs only serve a mechanical purpose, which is to circulate water through the canals which are sensory structures.[12][13]

The Atlantic mudskippers have genital papillae that are located on the abdomen. Females can be distinguished from males, who have less rounded papillae.[14]

Etymology

For the scientific name Periophthalmus barbarus, in Greek 'Peri' means 'around', and 'ophthalmos' means 'eye',[15] in reference to the Atlantic mudskipper's close set eyes that provide them with a larger field of vision.[16] In Greek, 'barbarus' means 'foreign', potentially named after the many differences it displayed when compared to other gobies.[15] The generic name 'mudskipper', is labelled after their 'skipping' movement on mudflats.[17]

Taxonomy

The initial identification and description of the species was outlined by Carl Linneaus, published in 1766 as Gobius barbarus and renamed to Periophthalmus barbarus.[18] The Atlantic mudskipper has also previously been known or misidentified as Gobius koelreuteri, Periophthalmus papilio, Periophthalmus gabonicus, Periophthalmus errythronemus.[19] The Atlantic mudskipper is distinguished from other members of Periophthalmus by having no spots or having some white spots on its back.[19]

Distribution

Periophthalmus barbarus is found throughout West Africa, in mangrove swamps and primarily brackish bodies of water near the coast.[1] Countries where the mudskipper are found include Angola, the Democratic Republic of the Congo, Cameroon and Ghana [1]

Ecology and behaviour

Habitat

Atlantic mudskippers are semi-aquatic animals that live in areas with water that is slightly salty, such as river estuaries and mudflats.[20][21] Generally, Atlantic mudskippers spend the majority of the day on land.[20] In tidal regions, Atlantic mudskippers may appear only during low tide to feed; whereas, they hide in their burrow at high tide.[21]

Predator-prey behaviour

The Atlantic mudskipper is carnivorous,[22] and has adopted an ambushing strategy to capture terrestrial prey.[23] To detect prey, the Atlantic mudskipper submerges itself underwater whilst leaving its eyes out, using only sight to identify and locate prey.[23] To ambush prey, Atlantic mudskippers launch onto land using predominantly their pectoral fins, and catch the prey using their mouth.[23]

Feeding behaviour

On land, the Atlantic mudskipper feeds by covering their prey with water, then sucking back the water and prey into their mouth, named as the 'hydrodynamic tongue'.[24] In water, the Atlantic mudskipper feeds through suction feeding, similar to other aquatic species.[25] Although the feeding technique is similar in both environments, the Atlantic mudskipper alters the force of suction, such that the flow underwater is stronger than on land.[25] The Atlantic mudskippers are diurnal, which means they are active and feed during the day.[20]

Atlantic mudskippers are flexible in regard to their diet choices. Larger Atlantic mudskippers ingest larger sized prey, potentially due to the correlation between their mouth gape and prey size. The feeding choices also may vary between habitats and seasonally, depending on what resources are most abundant. The Atlantic mudskippers feed more during dry season than wet, reflective of optimal foraging theory, which proposes that diet flexibility increases with lower food availability. Atlantic mudskippers feed frequently, where larger Atlantic mudskippers feed at higher intensities as they are less prone to predation.[26]

Reproduction

Females reach sexual maturity at a body length of around 10.2cm, and males approximately 10.8cm. The Atlantic mudskipper can spawn throughout the year.[27] However, spawning mainly occurs for male Atlantic mudskippers between February and May, and females between March and May.[28] The peak spawning times are associated with high food availability for their young.[28] The fertility of Atlantic mudskippers increases with the length of their body, as females may be able to carry more eggs at larger sizes.[27] Females lay thousands of eggs at one time; however, the eggs are highly prone to predation.[29]

During courtship, after the male pairs with a female mudskipper, the male and female go into the male's burrow to mate.[27] After the female releases her eggs onto the burrow wall,[27] the male displays a large amount of paternal care.[29] The male Atlantic mudskipper will guard and take care of the eggs within the burrow.[29]

Lifecycle

The average lifespan of an Atlantic mudskipper is approximately five years. The egg laid by the mother in the burrow, will hatch into larvae that swim out of the burrow and drift for 30-50 days. As Atlantic mudskippers are territorial, juvenile mudskippers will hide in mud until they mature enough to protect their established territory.[29]

Usage by humans

It is important to local indigenous peoples as a food fish and can also be found in the aquarium trade.[1] Fishing has caused population declines in parts of the species' range.[30]

Diet

Periophthalmus barbarus is an omnivore.[31] In the wild, mudskippers prefer to eat worms, crickets, flies, meal worms, beetles, small fish, and small crustaceans (sesarmid crabs).[31] Mudskippers kept as pets can eat frozen fare such as bloodworm or artemia and flake. It cannot eat dried food, however, because its stomach would swell up. It is recommended to feed it frozen food for a healthy diet.[32]

Characteristics

Periophthalmus barbarus and mudskippers generally are able to tolerate high concentrations of toxic substances produced by industrial waste, including cyanide, in their environments.[2] They are also able to survive in a variety of environments, including waters with different temperatures and salinity levels.[2]

References

  1. ^ a b c d Bousso, T.; Lalèyè, P. & Moelants, T. (2010). "Periophthalmus barbarus". The IUCN Red List of Threatened Species. 2010: e.T182285A7850201. doi:10.2305/IUCN.UK.2010-3.RLTS.T182285A7850201.en.
  2. ^ a b c Emuebie, Okonji Raphael (2011). "Physicochemical Properties of Mudskipper ( Periophthalmus Barbarus Pallas) Liver Rhodanese". Australian Journal of Basic and Applied Sciences. 5 (8): 507–514.
  3. ^ a b c d "Atlantic mudskippers: Meet them at Zoo Leipzig!". Zoo Leipzig. Retrieved 2020-10-07.
  4. ^ Murdy, Edward O. (1989). A Taxonomic Revision and Cladistic Analysis of the Oxudercine Gobies (Gobiidae, Oxudercinae). Australian Museum. ISBN 978-0-7305-6374-7.
  5. ^ a b c Kutschera, Ulrich; Elliott, J. Malcolm (2013-03-26). "Do mudskippers and lungfishes elucidate the early evolution of four-limbed vertebrates?". Evolution: Education and Outreach. 6 (1): 8. doi:10.1186/1936-6434-6-8. ISSN 1936-6434.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  6. ^ a b c d e Ansari, A.; Trivedi, S.; Saggu, Shalini; Rehman, H. (2014). "Mudskipper: A biological indicator for environmental monitoring and assessment of coastal waters". undefined. Retrieved 2020-10-07.
  7. ^ a b Pace, C. M.; Gibb, A. C. (2009-07-15). "Mudskipper pectoral fin kinematics in aquatic and terrestrial environments". Journal of Experimental Biology. 212 (14): 2279–2286. doi:10.1242/jeb.029041. ISSN 0022-0949. PMID 19561218.
  8. ^ "Atlantic mudskipper". Zoo Zürich. Retrieved 2020-10-11.
  9. ^ Michel, Krijn B.; Adriaens, Dominique; Aerts, Peter; Dierick, Manuel; Wassenbergh, Sam Van (2014). "Functional anatomy and kinematics of the oral jaw system during terrestrial feeding in Periophthalmus barbarus". Journal of Morphology. 275 (10). doi:10.1002/jmor.20318. ISSN 1097-4687.
  10. ^ Sponder, Debra L.; Lauder, G. (2009). "Terrestrial feeding in the Mudskipper Periophthalmus (Pisces: Teleostei): A cineradiographic analysis". doi:10.1111/J.1469-7998.1981.TB01501.X. {{cite journal}}: Cite journal requires |journal= (help)
  11. ^ Wołczuk, Katarzyna; Ostrowski, Maciej; Ostrowska, Agnieszka; Napiórkowska, Teresa (2018-06-01). "Structure of the alimentary tract in the Atlantic mudskipper Periophthalmus barbarus (Gobiidae: Oxudercinae): anatomical, histological and ultrastructural studies". Zoology. 128: 38–45. doi:10.1016/j.zool.2018.04.002. ISSN 0944-2006.
  12. ^ Kuciel, Michał (2013-03-01). "The mechanism of olfactory organ ventilation in Periophthalmus barbarus (Gobiidae, Oxudercinae)". Zoomorphology. 132 (1): 81–85. doi:10.1007/s00435-012-0167-y. ISSN 1432-234X. PMC 3570760. PMID 23420290.{{cite journal}}: CS1 maint: PMC format (link)
  13. ^ Kuciel, Michał; Żuwała, Krystyna; Jakubowski, Michał (2011). "A new type of fish olfactory organ structure in Periophthalmus barbarus (Oxudercinae)". Acta Zoologica. 92 (3): 276–280. doi:10.1111/j.1463-6395.2010.00459.x. ISSN 1463-6395.
  14. ^ Udo, M.; Abiaobo, No; Asuquo, I. (2016). "Aspects of the reproductive biology in mudskippers Periophthalmus barbarus ( Gobiidae ) ( Linneaus 1766 ) in mangrove swamps of Iko Estuary , Southeast , Nigeria". {{cite journal}}: Cite journal requires |journal= (help)
  15. ^ a b Murdy, Edward O. (1989). A Taxonomic Revision and Cladistic Analysis of the Oxudercine Gobies (Gobiidae, Oxudercinae). Australian Museum. ISBN 978-0-7305-6374-7.
  16. ^ Ansari, A.; Trivedi, S.; Saggu, Shalini; Rehman, H. (2014). "Mudskipper: A biological indicator for environmental monitoring and assessment of coastal waters". undefined. S2CID 54816035. Retrieved 2020-10-07.
  17. ^ "Definition of mudskipper | Dictionary.com". www.dictionary.com. Retrieved 2020-10-07.
  18. ^ "WoRMS - World Register of Marine Species - Periophthalmus barbarus (Linnaeus, 1766)". www.marinespecies.org. Retrieved 2020-10-07.
  19. ^ a b Murdy, Edward O. (1989). A Taxonomic Revision and Cladistic Analysis of the Oxudercine Gobies (Gobiidae, Oxudercinae). Australian Museum. ISBN 978-0-7305-6374-7.
  20. ^ a b c "Atlantic mudskipper". Zoo Zürich. Retrieved 2020-10-11.
  21. ^ a b "Periophthalmus barbarus (Atlantic Mudskipper) — Seriously Fish". Retrieved 2020-10-07.
  22. ^ Clayton, David (1993). "Mudskippers". Oceanography and Marine Biology. 31: 507–577.
  23. ^ a b c Kutschera, U.; Burghagen, H.; Ewert, J. (2008). "Prey-Catching Behaviour in Mudskippers and Toads: A Comparative Analysis". doi:10.3844/OJBSCI.2008.41.43. {{cite journal}}: Cite journal requires |journal= (help)
  24. ^ Michel, Krijn B.; Heiss, Egon; Aerts, Peter; Van Wassenbergh, Sam (2015-04-22). "A fish that uses its hydrodynamic tongue to feed on land". Proceedings of the Royal Society B: Biological Sciences. 282 (1805): 20150057. doi:10.1098/rspb.2015.0057.
  25. ^ a b Michel, K. B.; Aerts, P.; Wassenbergh, S. Van (2016-11-15). "Environment-dependent prey capture in the Atlantic mudskipper (Periophthalmus barbarus)". Biology Open. 5 (11): 1735–1742. doi:10.1242/bio.019794. ISSN 2046-6390. PMID 27765755.
  26. ^ Udo, Mfon T. "Tropic attributes of the mudskipper, Periophthalmus barbarus (Gobiidae: Oxudercinae) in the mangrove swamps of Imo River estuary, Nigeria". 14 (4): 508–517. {{cite journal}}: Cite journal requires |journal= (help)
  27. ^ a b c d Chukwu, K. O.; Deekae, S. N.; Gabriel, U. U. (2010). "Reproductive biology of Periopthalmus barbarus (Linneaus 1766) in New Calabar River, Nigeria". doi:10.5251/ABJNA.2010.1.6.1158.1161. {{cite journal}}: Cite journal requires |journal= (help)
  28. ^ a b Etim, Lawrence; King, Richard P.; Udo, Mfon T. (2002-06-01). "Breeding, growth, mortality and yield of the mudskipper Periophthalmus barbarus (Linneaus 1766) (Teleostei: Gobiidae) in the Imo River estuary, Nigeria". Fisheries Research. 56 (3): 227–238. doi:10.1016/S0165-7836(01)00327-7. ISSN 0165-7836.
  29. ^ a b c d Ansari, A.; Trivedi, S.; Saggu, Shalini; Rehman, H. (2014). "Mudskipper: A biological indicator for environmental monitoring and assessment of coastal waters". undefined. Retrieved 2020-10-07.
  30. ^ Etim, Lawrence (June 2002). "Breeding, growth, mortality and yield of the mudskipper Periophthalmus barbarus (Linnaeus 1766) (Teleostei: Gobiidae) in the Imo River estuary, Nigeria". Fisheries Research. 56 (3): 227–238. doi:10.1016/s0165-7836(01)00327-7.
  31. ^ a b "Periophthalmus barbarus". Mudskipper.it. Retrieved 28 October 2016.
  32. ^ Froese, Rainer; Pauly, Daniel (eds.) (2013). "Periophthalmus barbarus" in FishBase. June 2013 version.
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