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[[File:Mola alexandrini specimen.jpg |thumb|[[Taxidermy]] with a total length of {{convert|3|m|ft|abbr=on}} and a height of {{convert|3.5|m|ft|abbr=on}} in [[Aqua World]]]]
[[File:Mola alexandrini specimen.jpg |thumb|[[Taxidermy]] with a total length of {{convert|3|m|ft|abbr=on}} and a height of {{convert|3.5|m|ft|abbr=on}} in [[Aqua World]]]]


''Mola alexandrini'' has a relatively small mouth and its teeth fused into a [[parrot]]-like beak. It can reach up to {{convert|3.3|m|abbr=on}} in length and 2,300&nbsp;kg in mass, making it one of the two heaviest bony fish, only matched by its congener, the [[ocean sunfish]].<ref>{{Cite web | url=https://www.springer.com/gp/about-springer/media/research-news/all-english-research-news/world-s-heaviest-bony-fish-identified-and-correctly-named/15270666 | title=World's heaviest bony fish identified and correctly named}}</ref> Their body is flat and round, with large [[fin]]s that they swish back and forth to propel themselves with as they swim horizontally. Their skin has rough [[Dermal denticles|denticle]]s, [[leather]]y texture, with brown and gray coloring with pale blotches until death when they turn white.<ref name=museum>{{cite web|url=http://australianmuseum.net.au/Southern-Ocean-Sunfish-Mola-ramsayi|title=Southern Ocean Sunfish, Mola ramsayi (Giglioli, 1883) - Australian Museum|website=australianmuseum.net.au}}</ref> The body of the species has a thick white subcutaneous gelatinous layer that is smooth to the touch with a laterally compressed body covered in small rectangular scales <ref name=":0" />'''.''' Both ''mola'' [[species]] have no [[caudal (anatomical term)|caudal]] bones, [[rib]]s, and [[pelvic fin]]s and have fused [[vertebrae]], leaving only their median fins to propel themselves.<ref name=maridiver>{{cite journal|author1=Tierney M. Thys |author2=Jonathan Whitney |author3=Alex Hearn |author4=Kevin C. Weng |author5=Cesar Pen Aherrera |author6=L. Jawad |author7=J. Alfaro-Shigueto |author8=J.C. Mangel |author9=Stephen A. Karl |title=First record of the southern ocean sunfish, Mola ramsayi, in the Galapagos Marine Reserve|journal=Marine Biodiversity Records|volume=6|pages=1–4|doi=10.1017/S1755267213000377|year=2013 }}</ref> It can be recognized from the ''Mola mola'' by their lesser number of [[ossicle]]s and lacking of a vertical band of denticles at its base.<ref name = fishau /> '''I'''n ''Mola'', the lower jaws are intact while the upper jaws are slightly forked <ref>{{Cite book|last=R. Accademia delle scienze dell'Istituto di Bologna.|url=https://www.biodiversitylibrary.org/item/31099|title=Novi Commentarii Academiae Scientiarum Instituti Bononiensis.|last2=Bologna|first2=R. Accademia delle scienze dell'Istituto di|date=1839|volume=3}}</ref>'''.''' Also present on the head are lateral lines with small white rounded otoconia <ref name=":0" />'''.  '''On the sides of the fish are small gill openings that are covered by soft gill membrane and gill rakers which are covered under a subcutaneous gelatinous layer <ref name=":0" />'''.''' In these species, all fins are spineless and triangular with pectoral fins being small and rounded, located midlaterally fitting into shallow grooves on sides of the body <ref name=":0" />'''.''' Additionally, dorsal fins are located on opposite sides of the anal fin <ref name=":0" />'''.'''
''Mola alexandrini'' has a relatively small mouth and its teeth fused into a [[parrot]]-like beak. It can reach up to {{convert|3.3|m|abbr=on}} in length and 2,300&nbsp;kg in mass, making it one of the two heaviest bony fish, only matched by its congener, the [[ocean sunfish]].<ref>{{Cite web | url=https://www.springer.com/gp/about-springer/media/research-news/all-english-research-news/world-s-heaviest-bony-fish-identified-and-correctly-named/15270666 | title=World's heaviest bony fish identified and correctly named}}</ref> Their bodies are flat and round, with large [[fin]]s that they swish back and forth to propel themselves with as they swim horizontally. Their skin has rough [[Dermal denticles|denticle]]s, [[leather]]y texture, with brown and gray coloring with pale blotches until death when they turn white.<ref name=museum>{{cite web|url=http://australianmuseum.net.au/Southern-Ocean-Sunfish-Mola-ramsayi|title=Southern Ocean Sunfish, Mola ramsayi (Giglioli, 1883) - Australian Museum|website=australianmuseum.net.au}}</ref> The body of the species has a thick white subcutaneous gelatinous layer that is smooth to the touch with a laterally compressed body covered in small rectangular scales <ref name=":0" />'''.''' Both ''mola'' [[species]] have no [[caudal (anatomical term)|caudal]] bones, [[rib]]s, and [[pelvic fin]]s and have fused [[vertebrae]], leaving only their median fins to propel themselves.<ref name=maridiver>{{cite journal|author1=Tierney M. Thys |author2=Jonathan Whitney |author3=Alex Hearn |author4=Kevin C. Weng |author5=Cesar Pen Aherrera |author6=L. Jawad |author7=J. Alfaro-Shigueto |author8=J.C. Mangel |author9=Stephen A. Karl |title=First record of the southern ocean sunfish, Mola ramsayi, in the Galapagos Marine Reserve|journal=Marine Biodiversity Records|volume=6|pages=1–4|doi=10.1017/S1755267213000377|year=2013 }}</ref> It can be recognized from the ''Mola mola'' by their lesser number of [[ossicle]]s and lacking of a vertical band of denticles at its base.<ref name = fishau /> '''I'''n ''Mola'', the lower jaws are intact while the upper jaws are slightly forked <ref>{{Cite book|last=R. Accademia delle scienze dell'Istituto di Bologna.|url=https://www.biodiversitylibrary.org/item/31099|title=Novi Commentarii Academiae Scientiarum Instituti Bononiensis.|last2=Bologna|first2=R. Accademia delle scienze dell'Istituto di|date=1839|volume=3}}</ref>'''.''' Also present on the head are lateral lines with small white rounded otoconia <ref name=":0" />'''.  '''On the sides of the fish are small gill openings that are covered by soft gill membrane and gill rakers which are covered under a subcutaneous gelatinous layer <ref name=":0" />'''.''' In these species, all fins are spineless and triangular with pectoral fins being small and rounded, located midlaterally fitting into shallow grooves on sides of the body <ref name=":0" />'''.''' Additionally, dorsal fins are located on opposite sides of the anal fin <ref name=":0" />'''.'''


[[File:Bump head sunfish.jpg|thumb|right|Bump head sunfish, dwarfs diver]]
[[File:Bump head sunfish.jpg|thumb|right|Bump head sunfish, dwarfs diver]]

Revision as of 12:58, 2 June 2022

Giant sunfish
CITES Appendix II (CITES)[1]
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Tetraodontiformes
Family: Molidae
Genus: Mola
Species:
M. alexandrini
Binomial name
Mola alexandrini
(Ranzani, 1839)[2]
Synonyms
  • Orthragoriscus eurypterus Philippi 1892
  • Orthragoriscus ramsayi Giglioli 1839[3]
  • Mola ramsayi Giglioli 1883[4]

The southern sunfish (Mola alexandrini[4]), also known as the Ramsay's sunfish, southern ocean sunfish, short sunfish or bump-head sunfish in many parts of the world,[5] is a fish belonging to the family Molidae. It is closely related to its congener, the more widely known Mola mola, and is found in the Southern Hemisphere.[6] It can be found basking on its side occasionally near the surface, which is thought to be used to re-heat themselves after diving in cold water for prey, recharge their oxygen stores, and attract gulls to free them of parasites.[3]

Taxonomy

In December 2017, it was demonstrated that Mola alexandrini may be a senior synonym of Mola ramsayi (Ranzani 1839) through both historically and newly published morphological data.[7] In July 2020, building upon this scientific learning, the larval forms of these species were discovered for the first time and confirmed with DNA analyses by Australian and New Zealander scientific teams.[8][9][10][11]

A huge bump head mola caught at Darling harbour

The ocean sunfish are in the genus Mola, currently composed of three species: Mola mola, Mola alexandrini, Mola tecta (Yasemi and Narari Bejgan, 2013). Also known as the southern ocean sunfish or southern sunfish, Mola alexandrini are commonly found in the epipelagic zone of the ocean which is the part of the ocean where enough light penetrates for photosynthesis to occur (Matsuura, 2002) although recent studies also suggest that the sunfish are more common in deep waters than previously thought (Phillips et al. 2015).

Description

Taxidermy with a total length of 3 m (9.8 ft) and a height of 3.5 m (11 ft) in Aqua World

Mola alexandrini has a relatively small mouth and its teeth fused into a parrot-like beak. It can reach up to 3.3 m (11 ft) in length and 2,300 kg in mass, making it one of the two heaviest bony fish, only matched by its congener, the ocean sunfish.[12] Their bodies are flat and round, with large fins that they swish back and forth to propel themselves with as they swim horizontally. Their skin has rough denticles, leathery texture, with brown and gray coloring with pale blotches until death when they turn white.[4] The body of the species has a thick white subcutaneous gelatinous layer that is smooth to the touch with a laterally compressed body covered in small rectangular scales [7]. Both mola species have no caudal bones, ribs, and pelvic fins and have fused vertebrae, leaving only their median fins to propel themselves.[13] It can be recognized from the Mola mola by their lesser number of ossicles and lacking of a vertical band of denticles at its base.[5] In Mola, the lower jaws are intact while the upper jaws are slightly forked [14]. Also present on the head are lateral lines with small white rounded otoconia [7].  On the sides of the fish are small gill openings that are covered by soft gill membrane and gill rakers which are covered under a subcutaneous gelatinous layer [7]. In these species, all fins are spineless and triangular with pectoral fins being small and rounded, located midlaterally fitting into shallow grooves on sides of the body [7]. Additionally, dorsal fins are located on opposite sides of the anal fin [7].

File:Bump head sunfish.jpg
Bump head sunfish, dwarfs diver

M. alexandrini was found to be synonymous with M. ramsayi in July 2017 and can be distinguished by their unique characteristics of head bump, a chin bump, rectangular body scales, and rounded clavus. Although adult sunfish look generally similar, they are distinguishable using the seven characteristics: number of clavus fin rays (Yoshita et al 2009), number of clavus ossicles (Yoshita et al 2009), shape of clavus margin (Yoshita et al 2009), presence of head bump (Yoshita et al 2009), proportion of body length compared to body height (Yoshita et al 2009), shape of body scales (Fraser-Brunner 1951), and the presence of a chin bump (Sawai et al. 2017).

Alongside these species, there are no external differences between sexes however, the shape of gonads differs in males and females with females having a single ball-shaped ovary and males having a pair of elongated, rod-like testis [7].

Overall, the maximum recorded weight of M. alexandrini is 2,300 kg for a 272-cm total length female caught off the coast of Kamogawa, Japan in 1996.[7] With these records, this specimen is currently regarded as the world’s heaviest bony fish specimen [7].

Distribution

Mola alexandrini has been found all over the globe and is widely distributed throughout the world’s oceans except for two polar regions; the Arctic and Antarctic [7]. These species have been collected from waters off Japan, Taiwan, Galapagos Islands, New Zealand, Australia, Turkey, Oman, and Spain [7]. It can be found in the southwest Pacific, especially around Australia and New Zealand, and the southeast Pacific around Chile. Its range also extends to the southeast Atlantic near South Africa. During different seasons on the Pacific side of Japan, M. alexandrini will move northward in the summer and southward in the winter [15]. Seasonal migration is driven by temperatures and productive frontal areas [16].

Habitat

Although members of the genus Mola are found in many oceans throughout the world, this species thrives best in the open ocean of tropical and temperate seas preferring warmer temperatures with temperatures being between 16.8 C -25.6 C and averaging 19.9 C [7]. Many occurrences of these fish are linked to influences of ocean currents.[17] With the environment, sunfish have different movement patterns. During the nighttime, these species stay in the same areas but, during the daytime, they stay below thermocline [16]. Vertical movement patterns correlated with thermocline depth and were distinct from December to May [16].

Development

Overtime, as this species develops, there are physical changes to the body. This includes a head bump forming from above the eyes to the front of the dorsal-fin base and a chin bump developing from beneath the lower jaw to beneath the pectoral fins [7]. Additionally, developing with age are lateral ridges from above the head and below the eyes to beyond the pectoral fins [7]. Characteristics that distinguish Mola alexandrini from other species in genus Mola are clavus ossicles, a snout ossicle and a chin ossicle that develop overtime with age [7]. Transitioning from eggs into larvae, specimens can reach between 1.42 and 1.84 mm.[18] At 1.42 mm, a globular shape is portrayed [18]. As they move into the pre-juvenile stage, specimens reach between 5 mm to 59 mm.[18] As they continue to grow, their body transitions into adult body proportions, including an elongated body.[18] Once these species reach the juvenile stage, specimens are described to be as big as 305–750 mm in total length.[18] Moving into adulthood, specimens are claimed to reach 4000 mm with well-defined features along with pigmentations of a gray, olive gray or blackish with a brown cast [18].

Reproduction - general behavior/parental investment

Spawning of sunfish happen in the outer circulation of temperate Atlantic, Pacific Oceans, and in the Mediterranean Sea.[18] There is no absolute determination of when the best time is for spawning season, but research has shown that spawning in the Fall season and Winter seasons during the month of September result in bigger fish.[18] Fertilization occurs when sperm and eggs are shed in the water.[19]  Being that sunfish are so large, a single adult female can produce 300 million eggs.[19] Unfertilized eggs were measured at 0.42-0.45 mm in diameter.[18]

Lifespan/Longevity

Just like many other fish, high mortality rates are common for eggs, larvae, pre-juveniles, and small juveniles due to predators.[20] There have been few reports on predation of Mola species however, predation by fish are from families Scombridae, Carangidae, Coryphaenidae, Xiphiidae, and Alepisauridae.[20] Although there is not much research on the lifespan of ocean sunfish, it is said for ocean sunfish to take about 20 years for the species to reach a length of 3 m.[21]

Behavior

Sunfish swim by moving its dorsal and anal fins back and forth while moving both fins in the same direction at the same time.[19] Adults are reported to travel mainly in pairs and sometimes in groups.[18] Migrating from one place to another requires high tolerance and it is found that sunfish have high thermal tolerance undergoing quick and large temperature changes diving down the ocean several hundred meters.[17] Sometimes, sunfish come up to the shallow water to recover from hypoxia from feeding below the thermocline.[17] Like many other fish, sunfish adapt in response to the environment.[17] During the daytime, sunfish showed diel movement patterns with depths deeper than the nighttime.[16] During the nighttime, sunfish remained in temperatures between 18-24 C while the fish moved to different areas during the day time.[16] Vertical shifts were correlated with temperature shifts.[16]  Sunfish may dive deeper into cooler waters to cool the muscles or repay oxygen debts.[16]

Diet

They mainly consume jellyfish, which are of low nutritional content but abundant, and they will also eat brittle stars, small fish, plankton, algae, salps, and mollusks.[3] Sunfish also feed on ctenophores, hydrozonans, and small crustaceans.[19] It is also known for sunfish to swim up to the shallow water to eat cleaner fish and seabirds at the surface.[17] Juvenile sunfish feed in coastal areas in the coastal food web while larger sunfish dive deeper.[20] These species are active predators hunting in dynamic frontal systems.[17]

Predation

Predators are those who can cripple portions of a species; including sharks and orcas.[20] Sharks are the most common predators of ocean sunfish.[20] Specifically, tiger sharks can predate on ocean sunfish, however, it is rare.[20] The lack of cases on shark predation on the family Molidae is sporadic, leading to the idea that ocean sunfish are of low quality or unpleasant for tiger sharks.[20] Sharks tend to prey on larger prey as they become adults because there is an increase in size of the mouth and jaw.[20] With smart tactics, tiger sharks can stalk and ambush their prey and are able to bite through thick dermis.[20]  The thick dermis is a subcutaneous gelatinous layer consisting mostly of water making it undesirable.[20] In a tiger sharks’ diet, there is a low importance of sunfish but although cases are rare, it is never zero.[20]

Ecosystem Roles

With little to no information, the importance of ocean sunfish in marine food webs are unknown.[20]  However, since ocean sunfish feed on gelatinous prey with a generalist diet, this suggests that these species play an important role in coastal food webs.[22] If sunfish were to be removed as bycatch, it can drive localized trophic cascades with top-down control being reduced.[23]

Economic Importance for Humans

Sunfish have economic value in tourism industries.[24]  Seeing these fish are unpredictable, however, their unpredictable occurrences make exciting tours and recreational scuba diving.[24] In locations like the Galapagos Islands and the Alboran Coast in the Mediterranean, sunfish are spotted frequent enough to have sunfish tours.[17] Also, scuba diving tourism in Bali and the Nusa Penida islands have rapidly increased over the last few decades.[24] Only a small number of fisheries target sunfish, including those in Taiwan and Japan.[24] In Indonesia, sunfish are released, eaten by locals, used as bait, or end up at a fish market on rare occasions.[24]

Conservation Status

The conservation status of M. alexandrini has not been evaluated aside from M. mola [24]. Fisheries around the world catch ocean sunfish as bycatch.[24] M.mola was listed by the international Union for the Conservation of Nature (IUCN) as “vulnerable” due to the high level of estimated bycatch in South African longline fishery with an annual estimated 340,000 annual catches.[24] Also, both M. mola and M. alexandrini were listed as “high risk” bycatch species in the longline fishery off eastern Australia.[24]  Threat levels are lower than what is stated on the IUCN listing in Australian, New Zealand, and South African fisheries.[24] Currently, the Indonesian government, Ministry of Marine Affairs and Fisheries has placed sunfish on a plan for protection.[24]

References

  1. ^ "Appendices | CITES". cites.org. Retrieved 2022-01-14.
  2. ^ "Mola alexandrini summary page".
  3. ^ a b c Appeltans, W., Bouchet, P., Boxshall, G.A., Fauchald, K., Gordon, D.P., Hoeksema, B.W., Poore, G.C.B., van Soest, R.W.M., Stöhr, S., Walter, T.C., Costello, M.J. (eds.) (2010) World Register of Marine Species (WoRMS)
  4. ^ a b c "Southern Ocean Sunfish, Mola ramsayi (Giglioli, 1883) - Australian Museum". australianmuseum.net.au.
  5. ^ a b Diane J. Bray, 2011, Short Sunfish, or even. Mola ramsayi, in Fishes of Australia, accessed 02 Feb 2014, http://www.fishesofaustralia.net.au/home/species/784 Archived 2015-02-20 at the Wayback Machine
  6. ^ "Molas - Mola - Overview - Encyclopedia of Life". Encyclopedia of Life.[permanent dead link‍]
  7. ^ a b c d e f g h i j k l m n o Sawai, Etsuro; Yamanoue, Yusuke; Nyegaard, Marianne; Sakai, Yoichi (2017-12-05). "Redescription of the bump-head sunfish Mola alexandrini (Ranzani 1839), senior synonym of Mola ramsayi (Giglioli 1883), with designation of a neotype for Mola mola (Linnaeus 1758) (Tetraodontiformes: Molidae)". Ichthyological Research. 65: 142–160. doi:10.1007/s10228-017-0603-6. ISSN 1341-8998. S2CID 10364811.
  8. ^ http://www.medianet.com.au/releases/189537; https://www.9news.com.au/national/giant-bumphead-sunfish-australian-museum-discovery-mola-larve/a1899741-912e-47bf-b5f8-eebe578e2c20
  9. ^ "Giant Sunfish larva identified for the first time - and it is tiny".
  10. ^ "Scientists crack the enigmatic beginnings of the sunfish". 21 July 2020.
  11. ^ "Sunfish no more a quandary for scientists". 22 July 2020.
  12. ^ "World's heaviest bony fish identified and correctly named".
  13. ^ Tierney M. Thys; Jonathan Whitney; Alex Hearn; Kevin C. Weng; Cesar Pen Aherrera; L. Jawad; J. Alfaro-Shigueto; J.C. Mangel; Stephen A. Karl (2013). "First record of the southern ocean sunfish, Mola ramsayi, in the Galapagos Marine Reserve". Marine Biodiversity Records. 6: 1–4. doi:10.1017/S1755267213000377.
  14. ^ R. Accademia delle scienze dell'Istituto di Bologna.; Bologna, R. Accademia delle scienze dell'Istituto di (1839). Novi Commentarii Academiae Scientiarum Instituti Bononiensis. Vol. 3.
  15. ^ Sawai, Etsuro; Yamada, Morihiko (2020). "Bump-head sunfish Mola alexandrini photographed in the north-west Pacific Ocean mesopelagic zone". Journal of Fish Biology. 96 (1): 278–280. doi:10.1111/jfb.14214. ISSN 1095-8649.
  16. ^ a b c d e f g Chang, Ching-Tsun; Lin, Shian-Jhong; Chiang, Wei-Chuan; Musyl, Michael K.; Lam, Chi-Hin; Hsu, Hung-Hung; Chang, Yung-Chou; Ho, Yuan-Shing; Tseng, Chen-Te (2020-05-01). "Horizontal and vertical movement patterns of sunfish off eastern Taiwan". Deep Sea Research Part II: Topical Studies in Oceanography. Oceanic biodiversity under climate change: shifts in natural and human systems. 175: 104683. doi:10.1016/j.dsr2.2019.104683. ISSN 0967-0645.
  17. ^ a b c d e f g Nyegaard, Marianne; Loneragan, Neil; Hall, Steve; Andrew, James; Sawai, Etsuro; Nyegaard, Mette (2018-07-31). "Giant jelly eaters on the line: Species distribution and bycatch of three dominant sunfishes in the Southwest Pacific". Estuarine, Coastal and Shelf Science. 207: 1–15. doi:10.1016/j.ecss.2018.03.017. ISSN 0272-7714.
  18. ^ a b c d e f g h i j Faber, Daniel J.; McAllister, Don E. (1979-06-01). "Development of Fishes of the Mid-Atlantic Bight. An Atlas of egg, larval and juvenile stages". Journal of the Fisheries Research Board of Canada. 36 (6): 706–707. doi:10.1139/f79-107. ISSN 0015-296X.
  19. ^ a b c d Parkinson, Kerryn (4 July 2021). "Ocean Sunfish, Mola Mola (Linnaeus 1758)". The Australian Museum.{{cite web}}: CS1 maint: url-status (link)
  20. ^ a b c d e f g h i j k l Nyegaard, Marianne; Andrzejaczek, Samantha; Jenner, Curt S.; Jenner, Micheline-Nicole M. (2019-12-01). "Tiger shark predation on large ocean sunfishes (Family Molidae) – two Australian observations". Environmental Biology of Fishes. 102 (12): 1559–1567. doi:10.1007/s10641-019-00926-y. ISSN 1573-5133.
  21. ^ 俊之, 中坪; 一美, 廣瀬 (2007). "飼育下におけるマンボウの成長". 水産増殖. 55 (3): 403–407. doi:10.11233/aquaculturesci1953.55.403.
  22. ^ Mangel, Jeffrey C.; Pajuelo, Mariela; Pasara‐Polack, Andrea; Vela, German; Segura‐Cobeña, Eduardo; Alfaro‐Shigueto, Joanna (2018-12-11). "The effect of Peruvian small‐scale fisheries on sunfishes (Molidae)". Journal of Fish Biology. doi:10.1111/jfb.13862. ISSN 0022-1112.
  23. ^ Phillips, Natasha D.; Kubicek, Lukas; Payne, Nicholas L.; Harrod, Chris; Eagling, Lawrence E.; Carson, Carol D.; Cappanera, Valentina; Houghton, Jonathan D.R. (September 2018). "Isometric growth in the world's largest bony fishes (genus Mola )? Morphological insights from fisheries bycatch data". Journal of Morphology. 279 (9): 1312–1320. doi:10.1002/jmor.20872. ISSN 0362-2525.
  24. ^ a b c d e f g h i j k Nyegaard, Marianne (2018). There be giants! The importance of taxonomic clarity of the large ocean sunfishes (genus Mola, Family Molidae) for assessing sunfish vulnerability to anthropogenic pressures (phd thesis). Murdoch University.

Further reading

  • Glover, C.J.M. in Gomon, M.F., Glover, C.J.M. & R.H. Kuiter (Eds). (1994). The Fishes of Australia's South Coast. State Print, Adelaide. Pp. 992.
  • Hutchins, B. & R. Swainston. (1986). Sea Fishes of Southern Australia. Complete Field Guide for Anglers and Divers. Swainston Publishing. Pp. 180.
  • Hutchins, B. & M. Thompson. 1983. The Marine and Estuarine Fishes of South-western Australia. Western Australian Museum. Pp. 103.
  • Last, P.R., E.O.G. Scott & F.H. Talbot. (1983). Fishes of Tasmania. Tasmanian Fisheries Development Authority. Pp. 563.
  • Matsuura, K., 2002. Molidae. Molas (ocean sunfishes, headfishes). In Carpenter K.E. (ed.)
  • Phillips ND, Harrod C, Gates AR, Thys TM, Houghton JDR (2015) Seeking the sun in deep, dark places: mesopelagic sightings of ocean sunfishes (Molidae). Journal of Fish Biology, 4, 1118-1126.
  • Phillips ND, Reid N, Thys T et al. (2017) Applying species distribution modelling to a data poor, pelagic fish complex: the ocean sunfishes. Journal of Biogeography, 44, 2176-2187.
  • Sawai E, Yamanoue Y, Nyegaard M, Sakai Y (2018) Redescription of the bump-head sunfish Mola alexandrini (Ranzani 1839), senior synonym of Mola ramsayi (Giglioli 1883), with designation of a neotype for Mola mola (Linnaeus 1758) (Tetraodontiformes: Molidae). Ichthyological Research, 65, 142-160.
  • Sawai E, Yamanoue Y, Jawad L, Al-Mamry J, Sakai Y (2017) Molecular and morphological identification of Mola sunfsh specimens (Actinopterygii: Tetraodontiformes: Molidae) from the Indian Ocean. Species Divers, 2, 99–104.
  • Yasemi M, Nazari Bejgan AR (2013) The first record of southern ocean sunfish, Mola ramsayi from Northern Oman Sea, Iran. Iranian Journal of Fisheries Sciences, 13, 242-246.
  • Yoshita Y, Yamanoue Y, Sagara K, Nishibori M, Kuniyoshi H, Umino T, Sakai Y, Hashimoto H, Gushima K (2009) Phylogenetic relationships of two Mola sunfshes (Tetraodontiformes: Molidae) occurring around the coasts of Japan, with notes on their geographical distribution and morphological characteristics. Ichthyol Res, 56, 232–244.
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