Abstract
Relatively little is known about the biology and ecology of the world’s largest (heaviest) bony fish, the ocean sunfish Mola mola, despite its worldwide occurrence in temperate and tropical seas. Studies are now emerging that require many common perceptions about sunfish behaviour and ecology to be re-examined. Indeed, the long-held view that ocean sunfish are an inactive, passively drifting species seems to be entirely misplaced. Technological advances in marine telemetry are revealing distinct behavioural patterns and protracted seasonal movements. Extensive forays by ocean sunfish into the deep ocean have been documented and broad-scale surveys, together with molecular and laboratory based techniques, are addressing the connectivity and trophic role of these animals. These emerging molecular and movement studies suggest that local distinct populations may be prone to depletion through bycatch in commercial fisheries. Rising interest in ocean sunfish, highlighted by the increase in recent publications, warrants a thorough review of the biology and ecology of this species. Here we review the taxonomy, morphology, geography, diet, locomotion, vision, movements, foraging ecology, reproduction and species interactions of M. mola. We present a summary of current conservation issues and suggest methods for addressing fundamental gaps in our knowledge.
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Ackman RG (1997) Has evolution and long-term coexistence adapted us to cope with trans fatty acids? J Food Lipids 4:295–318. doi:10.1111/j.1745-4522.1997.tb00101.x
Alfaro ME, Santini F, Brock CD (2007) Do reefs drive diversification in marine teleosts? Evidence from the pufferfish and their allies (order Tetraodontiformes). Evolution 61:2104–2126. doi:10.1111/j.1558-5646.2007.00182.x
Arai MN (2005) Predation on pelagic coelenterates: a review. J Mar Biol Assoc UK 85:523–536.
Arai T, Levin AV, Boltunov AN, Miyazaki N (2002) Migratory history of the Russian sturgeon Acipenser guldenstadti in the Caspian Sea, as revealed by pectoral fin spine Sr:Ca ratios. Mar Biol 141:315–319. doi:10.1007/s00227-002-0820-y
Arreola VI, Westneat MW (1996) Mechanics of propulsion by multiple fins: kinematics of aquatic locomotion in the burrfish (Chilomycterus schoepfi). Proc R Soc Lond B 263:1689–1696
Bainbridge R (1958) The speed of swimming of fish as related to size and to the frequency and amplitude of the tail beat. J Exp Biol 35:109–133
Bass AL, Dewar H, Thys T, Streelman JT, Karl SA (2005) Evolutionary divergence among lineages of the ocean sunfish family, Molidae (Tetraodontiformes). Mar Biol 148:405–414. doi:10.1007/s00227-005-0089-z
Baum JK, Myers RA, Kehler DG, Worm B, Harley SJ, Doherty PA (2003) Collapse and conservation of shark populations in the Northwest Atlantic. Science 299:389–392. doi:10.1126/science.1079777
Bjorndal KA (1997) Foraging ecology and nutrition of sea turtles. In: Lutz PL, Musick JA (eds) The biology of sea turtles. CRC Press, Boca Raton, pp 199–231
Bonaparte CL (1841) New systematic arrangement of vertebrated animals. Trans Linn Soc Lond 18:247–304
Britz R, Johnson G (2005) Occipito-vertebral fusion in ocean sunfishes (Teleostei: Tetraodontiformes: Molidae) and its phylogenetic implications. J Morphol 266:74–79. doi:10.1002/jmor.10366
Brongersma LD (1972) European Atlantic turtles. Zool Verh 121:1–318
Brudenall DK, Schwab IR, Fritsches KA (2008) Ocular morphology of the Leatherback sea turtle (Dermochelys coriacea). Vet Ophthalmol 11:99–110
Carey FG (1982) A brain heater in swordfish. Science 216:1327–1329
Carey FG, Robison BH (1981) Daily patterns in the activities of swordfish, Xiphias gladius, observed by acoustic telemetry. Fish Bull 79:277–292
Carey FG, Scharold JV (1990) Movements of blue sharks (Prionace glauca) in depth and course. Mar Biol 106:329–342
Cartamil DP, Lowe CG (2004) Diel movement patterns of ocean sunfish Mola mola off southern California. Mar Ecol Prog Ser 266:245–253
Carwardine M (1995) The Guinness book of animal records. Guinness Publishing, Middlesex, UK
Celona A, Donato N, De Maddalena A (2001) In relation to the captures of a great white shark, Carcharodon carcharias (Linnaeus, 1758), and a shortfin mako, Isurus oxyrinchus Rafinesque, 1809, in the Messina Strait. Ann Ser Hist Nat 11:13–16
Cleland J (1862) On the anatomy of the short sun-fish (Orthragoriscus mola). Nat Hist Rev (Lond) 2:170–185
Clemens WA, Wilby A (1961) Fishes of the Pacific coast of Canada, 2nd edn. Fisheries Research Board of Canada Bulletin 68, Ottawa
Dagorn L, Bach P, Josse E (2000) Movement patterns of large bigeye tuna (Thunnus obesus) in the open ocean, determined using ultrasonic telemetry. Mar Biol 136:361–371
Davenport J, Kjørsvik E (1986) Buoyancy in the lumpsucker Cyclopterus lumpus. J Mar Biol Assoc UK 66:159–174
Doyle TK, Houghton JDR, O’Súilleabháin PF, Hobson VJ, Marnell F, Davenport J, Hays GC (2008) Leatherback turtles satellite tagged in European waters. Endanger Species Res 4:23–31. doi:10.3354/esr00076
Dulčić J, Paklar GB, Grbec B, Morović M, Matić F, Lipej L (2007) On the occurrence of ocean sunfish Mola mola and slender sunfish Ranzanza laevis in the Adriatic Sea. J Mar Biol Assoc UK 87:789–796. doi:10.1017/S0025315407053842
Fergusson IK, Compagno LJ, Marks MA (2000) Predation by white sharks Carcharodon carcharias (Chondrichthyes: Lamnidae) upon chelonians, with new records from the Mediterranean Sea and a first record of the ocean sunfish Mola mola (Osteichthyes: Molidae) as stomach contents. Environ Biol Fish 58:447–453
FishBase World Wide Web Electronic Publication (2008) Froese R, Pauly D (eds) version (12/2008). www.fishbase.org. Accessed 13 February 2009
Fraser-Brunner A (1951) The ocean sunfishes (Family Molidae). Bull Br Mus (Nat Hist) Zool 1:87–121
Fritsches KA, Marshall NJ, Warrant EJ (2003) Retinal specializations in the blue marlin: eyes designed for sensitivity to low light levels. Mar Freshw Res 54:333–341. doi:10.1071/MF02126
Fulling GL, Dagmar F, Knight K, Hoggard W (2007) Distribution of Molidae in the northern Gulf of Mexico. Gulf Caribb Res 19:53–67
Gauldie RW (1990) Vaterite otoliths from the opah, Lampris immaculatus, and two species of sunfish, Mola mola and M. ramsayi. Acta Zool 71:193–199
Giglioli HH (1883) Zoology at the fisheries exhibition. Nature 28:313–316
Gill TN (1897) The distinctive characters of the Molinae and Ranzaniinae. Science 156:966–967
Gladstone W (1988) Killer whale feeding observed underwater. J Mammal 69:629–630
Gleiss AC, Gruber SH, Wilson RP (2009) Multi-channel data-logging: towards determination of behaviour and metabolic rate in free-swimming sharks. In: Nielsen JL, Arrizabalaga H, Fragoso N, Hobday A, Lutcavage M, Sibert J (eds) Tagging and tracking of marine animals with electronic devices, vol 9. Springer, Dordrecht, pp 211–228. doi:10.1007/978-1-4020-9640-213
Gordon MS, Plaut I, Kim D (1996) How pufferfish (Teleostei: Tetraodontidae) swim. J Fish Biol 49:319–328
Graham BS, Grubbs D, Holland K, Popp BN (2007) A rapid ontogenetic shift in the diet of juvenile yellowfin tuna from Hawaii. Mar Biol 150:647–658. doi:10.1007/s00227-006-0360-y
Gudger EW (1936) From atom to colossus. Nat Hist 38:26–30
Hart JL (1973) Pacific fishes of Canada. Fisheries Research Board of Canada Bulletin 180, Ottawa
Hays GC, Doyle TK, Houghton JDR, Lilley MKS, Metcalfe JD, Righton D (2008) Diving behaviour of jellyfish equipped with electronic tags. J Plankton Res 30:325–331. doi:10.1093/plankt/fbn003
Hays GC, Farquhar MR, Luschi P, Teo SLH, Thys TM (2009) Vertical niche overlap by two ocean giants with similar diets: oceanic sunfish and leatherback turtles. J Exp Mar Biol Ecol 370:134–143. doi:10.1016/j.jembe.2008.12.009
Heithaus MR, Frid A, Wirsing AJ, Worm B (2008) Predicting ecological consequences of marine top predator declines. Trends Ecol Evol 23:202–210. doi:10.1016/j.tree.2008.01.003
Hixon MA (1979) The halfmoon, Medialuna californiensis, as a cleaner fish. Calif Fish Game 65:117–118
Hooper S, Ackman R (1970) Trans-6-hexadecenoic acid in the Atlantic leatherback Dermochelys coriacea coriacea L. and other marine turtles. Lipids 5:288–292
Hooper S, Ackman R (1972) Presence of trans-6-hexadecenoic acid in the white jellyfish Aurelia aurita Lamarck and in a Caribbean gorgonian. Lipids 7:624–626
Hooper SN, Paradis M, Ackman RG (1973) Distribution of trans-6-hexadecenoic acid, 7-methyl-7-hexadecenoic acid and common fatty acids in lipids of ocean sunfish Mola mola. Lipids 8:509–516
Houghton JDR, Doyle TK, Davenport J, Hays GC (2006a) The ocean sunfish Mola mola: insights into distribution, abundance and behaviour in the Irish and Celtic Seas. J Mar Biol Assoc UK 86:1237–1243. doi:10.1017/S002531540601424x
Houghton JDR, Doyle TK, Davenport J, Hays GC (2006b) Developing a simple, rapid method for identifying and monitoring jellyfish aggregations from the air. Mar Ecol Prog Ser 314:159–170
Houghton JDR, Doyle TK, Wilson MW, Davenport J, Hays GC (2006c) Jellyfish aggregations and leatherback turtle foraging patterns in a temperate coastal environment. Ecology 87:1967–1972
Houghton JDR, Doyle TK, Davenport J, Wilson RP, Hays GC (2008) The role of infrequent and extraordinary dives in the leatherback turtle (Dermochelys coriacea). J Exp Biol 211:2566–2575. doi:10.1242/jeb.020065
Houghton JDR, Liebsch N, Doyle TK, Gleiss A, Lilley MKS, Wilson RP, Hays GC (2009) Harnessing the sun: testing a novel attachment method to record fine scale movements in Ocean Sunfish (Mola mola). In: Nielsen JL, Arrizabalaga H, Fragoso N, Hobday A, Lutcavage M, Sibert J (eds) Tagging and tracking of marine animals with electronic devices, vol 9. Springer, Dordrecht, pp 229–242. doi:10.1007/978-1-4020-9640-214
Hove JR, O’Bryan LM, Gordon MS, Webb PW, Weihs D (2001) Boxfishes (Teleostei: Ostraciidae) as a model system for fishes swimming with many fins: kinematics. J Exp Biol 204:1459–1471
Iverson SJ, Field C, Bowen WD, Blanchard W (2004) Quantitative fatty acid signature analysis: a new method of estimating predator diets. Ecol Monogr 74:211–235
Johnson G, Britz R (2005) Leis’ conundrum: Homology of the clavus of the ocean sunfishes. 2. Ontogeny of the median fins and axial skeleton of Ranzania laevis (Teleostei, Tetraodontiformes, Molidae). J Morphol 266:11–21. doi:10.1002/jmor.10242
Kino M, Miayzaki T, Iwami T, Kohbara J (2009) Retinal topography of ganglion cells in immature ocean sunfish, Mola mola. Environ Biol Fish 85:33–38. doi:10.1007/s10641-009-9453-z
Kitagawa T, Kimura S, Nakata H, Yamada H (2007) Why do young Pacific bluefin tuna repeatedly dive to depths through the thermocline? Fish Sci 73:98–106
Konow N, Fitzpatrick R, Barnett A (2006) Adult emperor angelfish (Pomacanthus imperator) clean giant sunfishes (Mola mola) at Nusa Lembongan, Indonesia. Coral Reefs 25:208. doi:10.1007/s00338-006-0086-9
Liu KM, Lee ML, Joung SJ, Chang YC (2009) Age and growth estimates of the sharptail mola, Masturus lanceolatus, in waters of eastern Taiwan. Fish Res 95:154–160. doi:10.1016/j.fishres.2008.08.013
Love M, Moser M (1983) Technical Report NMFS SSRF-777 A checklist of parasites of California. Oregon and Washington marine and estuarine fishes. U.S. Department of Commerce, Washington, DC
Lynam CP, Gibbons MJ, Axelsen BE, Sparks CAJ, Coetzee J, Heywood BG, Brierley AS (2006) Jellyfish overtake fish in a heavily fished ecosystem. Curr Biol 16:R492–R493. doi:10.1016/j.cub.2006.06.018
MacGinitie GE, MacGinitie N (1968) Natural history of marine animals, 2nd edn. McGraw-Hill Book Company, New York
McMahon CR, Hays GC (2006) Thermal niche, large-scale movements and implications of climate change for a critically endangered marine vertebrate. Glob Change Biol 12:1330–1338. doi:10.1111/j.1365-2486.2006.01174.x
Mills CE (2001) Jellyfish blooms: are populations increasing globally in response to changing ocean conditions? Hydrobiologia 451:55–68
Motani R, Rothschild BM, Wahl W (1999) Large eyeballs in diving ichthyosaurs—the huge eyes of these extinct reptiles may have been useful deep in the ocean. Nature 402:747
Myers RA, Worm B (2003) Rapid worldwide depletion of predatory fish communities. Nature 423:280–283. doi:10.1038/nature01610
Myers RA, Baum JK, Shepherd TD, Powers SP, Peterson CH (2007) Cascading effects of the loss of apex predatory sharks from a coastal environment. Science 315:1846–1850. doi:10.1126/science.1138657
Nakae M, Sasaki K (2006) Peripheral nervous system of the ocean sunfish Mola mola (Tetraodontiformes: Molidae). Ichthyol Res 53:233–246. doi:10.1007/s10228-006-0339-1
Nakatsubo T (2008) A study on the reproductive biology of ocean sunfish Mola mola (in Japanese with English summary). Dissertation, International Marine Biological Institute, Kamagora Sea World, Japan
Nakatsubo T, Hirose H (2007) Growth of captive ocean sunfish, Mola mola (in Japanese with English abstract). Suisan Zoshoku 55:403–407
Nakatsubo T, Kawachi M, Mano N, Hirose H (2007) Spawning period of ocean sunfish Mola mola in waters of the eastern Kanto Region, Japan (in Japanese with English abstract). Suisan Zoshoku 55:613–618
Nelson DR, McKibben JN, Strong WR Jr, Lowe CG, Sisneros JA, Schroeder DM, Lavenberg RJ (1997) An acoustic tracking of a megamouth shark, Megachasma pelagios: a crepuscular vertical migratory. Environ Biol Fish 49:389–399
Norman JR, Fraser FC (1949) Field book of giant fishes. GP Putnam and Sons, New York
Parsons TR, Takahashi M, Hargreave B (1984) Biological oceanographic processes, 3rd edn. Pergamon Press, Oxford
Pascal J, Ackman RG (1975) Occurrence of 7-methyl-7-hexadecenoic acid, the corresponding alcohol, 7-methyl-6-hexadecenoic acid, and 5-methyl-4-hexadecenoic acid in sperm whale oils. Lipids 10:478–482
Pearce R, Stillway L (1976) Trans-6-hexadecenoic acid in the spadefish Chaetodipterus faber. Lipids 11:247–249
Petersen S (2005) Initial bycatch assessment: South Africa’s domestic longline fishery, 2000–2003. Domestic pelagic longline fishery: Bycatch Report 2000–2003, BirdLife South Africa
Petersen S, McDonell Z (2007) A bycatch assessment of the Cape horse mackerel Trachurus trachurus capensis mid-water trawl fishery off South Africa. Birdlife/WWF Responsible Fisheries Programme Report 2002–2005
Powell DC (2001) A fascination for fish: adventures of an underwater pioneer. University of California Press, Berkeley
Purcell JE, Arai MN (2001) Interactions of pelagic cnidarians and ctenophores with fish: a review. Hydrobiologia 451:27–44
Richardson AJ, Bakun A, Hays GC, Gibbons MJ (2009) The jellyfish joyride: causes, consequences and management actions. Trends Ecol Evol. doi:10.1016/j.tree.2009.01.010
Roach J (2003) World’s heaviest bony fish discovered? National Geographic. Available via http://news.nationalgeographic.com/news/2003/05/0513_030513_sunfish.html
Rooker JR, Secor DH, DeMetrio G, Schloesser R, Block BA, Neilson JD (2008) Natal homing and connectivity in Atlantic bluefin tuna populations. Science 322:742–744. doi:10.1126/science.1161473
Sagara K, Ozawa T (2002) Landing statistics of molids in four prefectures of Japan (in Japanese with English abstract). Mem Fac Fish Kagoshima Univ 51:27–33
Santini F, Tyler JC (2002) Phylogeny of the ocean sunfishes (Molidae, Tetraodontiformes), a highly derived group of teleost fishes. Ital J Zool 69:37–43
Santini F, Tyler JC (2003) A phylogeny of the families of fossil and extant tetraodontiform fishes (Acanthomorpha, Tetraodontiformes), Upper Cretaceous to recent. Zool J Linn Soc 139:565–617
Santini F, Tyler JC (2004) The importance of even highly incomplete fossil taxa in reconstructing the phylogenetic relationships of the Tetraodontiformes (Acanthomorpha: Pisces). Integr Comp Biol 44:349–357
Schmidt J (1921) New studies of sun-fishes made during the ‘‘Dana’’ Expedition, 1920. Nature 107:76–79
Shepard ELC, Ahmed MZ, Southall EJ, Witt MJ, Metcalfe JD, Sims DW (2006) Diel and tidal rhythms in diving behaviour of pelagic sharks identified by signal processing of archival tagging data. Mar Ecol Prog Ser 328:205–213
Shepard ELC, Wilson RP, Quintana F, Albareda DA, Gómez Laich A, Halsey LG, Liebsch N, Gleiss A, Morgan DT, Myers AE, Newman C, Macdonald DW (2008) Identification of animal movement patterns using tri-axial accelerometry. Endanger Species Res. doi:10.3354/esr00084
Silvani L, Gazo M, Aguilar A (1999) Spanish driftnet fishing and incidental catches in the western Mediterranean. Biol Conserv 90:79–85
Sims DW, Southall EJ (2002) Occurrence of ocean sunfish, Mola mola near fronts in the western English Channel. J Mar Biol Assoc UK 82:927–928
Sims DW, Southall EJ, Richardson AJ, Reid PC, Metcalfe JD (2003) Seasonal movements and behaviour of basking sharks from archival tagging: no evidence of winter hibernation. Mar Ecol Prog Ser 248:187–196
Sims DW, Southall EJ, Humphries NE, Hays GC, Bradshaw CJA, Pitchford JW, James A, Ahmed MZ, Brierley AS, Hindell MA, Morritt D, Musyl MK, Righton D, Shepard ELC, Wearmouth VJ, Wilson RP, Witt MJ, Metcalfe JD (2008) Scaling laws of marine predator search behaviour. Nature 451:1098–1102. doi:10.1038/nature06518
Sims DW, Queiroz N, Doyle TK, Houghton JDR, Hays GC (2009a) Satellite tracking of the world’s largest bony fish, the ocean sunfish (Mola mola) in the North East Atlantic. J Exp Mar Biol Ecol 370:127–133. doi:10.1016/j.jembe.2008.12.011
Sims DW, Queiroz N, Humphries NE, Lima FP, Hays GC (2009b) Long-term GPS tracking of ocean sunfish Mola mola offers a new direction in fish monitoring. PLoS ONE 4:e7351. doi:10.1371/journal.pone.0007351
Thys T (1994) Swimming heads. Nat Hist 103:36–39
Trudel M, Welch DW, Morris JFT, Candy JR, Beacham TD (2004) Using genetic markers to understand the coastal migration of juvenile coho (Oncorhynchus kisutch) and chinook salmon (O. tshawytscha). N Pac Anadr Fish Comm Tech Rep 5:52–54
Tudela S, Kai Kai A, Maynou F, El Andalossi M, Guglielmi P (2005) Driftnet fishing and biodiversity conservation: the case study of the large-scale Moroccan driftnet fleet operating in the Alboran Sea (SW Mediterranean). Biol Conserv 121:65–78. doi:10.1016/j.biocon.2004.04.010
Vogel S (1994) Life in moving fluids: the physical biology of flow, 2nd edn. Princeton Univ Press, Princeton
Walker JA, Westneat MW (2000) Mechanical performance of aquatic rowing and flying. Proc R Soc Lond B 267:1875–1881
Walls GL (1942) The vertebrate eye and its adaptive radiation. Hafner, New York
Watanabe Y, Sato K (2008) Functional dorsoventral symmetry in relation to lift-based swimming in the ocean sunfish Mola mola. PLoS ONE 3:e3446. doi:10.1371/journal.pone.0003446
Wilson RP, Shepard ELC, Liebsch N (2008) Prying into the intimate details of animal lives: use of a daily diary on animals. Endanger Species Res 4:123–137. doi:10.3354/esr00064
Yamanoue Y, Miya M, Matsuura K, Katoh M, Sakai H, Nishida M (2004) Mitochondrial genomes and phylogeny of the ocean sunfishes (Tetraodontiformes:Molidae). Ichthyol Res 51:269–273. doi:10.1007/s10228-004-0218-6
Yamanoue Y, Miya M, Matsuura K, Katoh M, Sakai H, Nishida M (2008) A new perspective on phylogeny and evolution of tetraodontiform fishes (Pisces:Acanthopterygii) based on whole mitochondrial genome sequences: basal ecological diversification? BMC Evol Biol 8:212. doi:10.1186/1471-2148-8-212
Yancey PH, Lawrence-Berrey R, Douglas MD (1989) Adaptations in mesopelagic fishes. Mar Biol 103:453–459
Yoshita Y, Yamanoue Y, Sagara K, Nishibori M, Kuniyoshi H, Umino T, Sakai Y, Hashimoto H, Gushima K (2009) Phylogenetic relationship of two Mola sunfishes (Tetraodontiformes: Molidae) occurring around the coast of Japan, with notes on their geographical distribution and morphological characteristics. Ichthyol Res 56:232–244. doi:10.1007/s10228-008-0089-3
Acknowledgments
We are indebted to Francesco Santini for his comments on molid phylogeny and to Toshiyuki Nakatsubo and Rand Rasmussen for their kind contribution of fisheries and bycatch data. We also gratefully acknowledge Mike Johnson, Wolfgang Sterrer, Tao-Jen Kuo, Kamogawa Seaworld and Birdlife International for the photographs used in this review. We would like to thank an anonymous reviewer for their constructive advice and suggestions. ECP was supported by the Welsh Institute for Sustainable Environments (WISE) Network. GCH, TKD and VJH were supported by the EcoJel project funded by INTERREG IVA, a component of the European Regional Development Fund. TMT was supported by the Adopt a Sunfish Project and a grant from the National Geographic Society’s Committee for Research and Exploration #7369-02. NQ was funded by a Fundação para a Ciência e a Tecnologia (FCT) Ph.D. grant SFRH/BD/21354/2005. DWS was supported by the UK Natural Environment Research Council Oceans 2025 Strategic Research Programme and an MBA Senior Research Fellowship.
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Pope, E.C., Hays, G.C., Thys, T.M. et al. The biology and ecology of the ocean sunfish Mola mola: a review of current knowledge and future research perspectives. Rev Fish Biol Fisheries 20, 471–487 (2010). https://doi.org/10.1007/s11160-009-9155-9
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DOI: https://doi.org/10.1007/s11160-009-9155-9