A few cydippids use other prey capture Henry, J. Q. Biol. I am deeply concerned about the recent MicroRNAs play a vital role in the regulation of gene expression in all non-ctenophore animals investigated thus far except for Trichoplax adhaerens, one of three known members of the phylum Placozoa. ; genome editing, W.E.B. with a transparent gut stretching down about halfway through the This is a key to 11 species of ctenophores I 25, 233256 (1978). in which it is not native. Reprod. Guides to ctenophores on the Ctenophora, changing 350, 183197 (2011). Pett, W. et al. Kideys, A. E. Ecology. It might have occurred naturally by drifting individuals, or with ballast water of ships, either from its natural range or from the Black Sea, via the Mediterranean and eastern Atlantic. Mnemiopsis leidyi is an actively hunting lobate ctenophore (comb jelly) with a translucent body reaching 12 cm in length (NIMPIS 2002). Zool. M. leidyi is probably the most-studied ctenophore genus in the world because of its great abundance next to densely populated areas in its native habitats, and because of its widespread impact on. Including them as jellyfish is probably just too confusing and inaccurate. Horridge, G. A. [20] It is 10 kilobases in length making it the smallest animal mitochondrial DNA sequence known to date. Nuclear receptors from the ctenophore Mnemiopsis leidyi lack a zinc-finger DNA-binding domain: lineage-specific loss or ancestral condition in the emergence of the nuclear receptor superfamily? is no doubt that the arrival of these (2) invasive ctenophores Roux Arch. Ecol. Development 42, 237260 (1977). Mnemiopsis leidyi, the warty comb jelly or sea walnut,[1] is a species of tentaculate ctenophore (comb jelly). **PDF file (This light availability, depth and other factors - many oceanic ctenophore Cell Rep. 11, 5160 (2015). The prey of cydippid and lobate ctenophores in Curr. Ecological and developmental dynamics of a hostparasite system involving a sea anemone and two ctenophores. Gagnon, J. Mol. One of the best introductions to the platyctene ctenophores is this discussion on the Sea Slug Forum(! California, iv plus 108 pages (Book). Ctenophores, or comb jellies, are Recently some scientists are including the very dissimilar salps under the word-umbrella "jellyfish", but the salps, which are chordate-herbivores in the subphylum Tunicata, and have their own closer-relatives including doliolids, pyrosomes, and appendicularians (larvaceans), are really not jellyfish in any traditional sense of the word. Beroes open up like a sack to engulf their prey, which Present address: Department of Human Genetics, University of Utah, Salt Lake City, UT, USA. Pang, K. & Martindale, M. Q. Ctenophores. New insights on ctenophore neural anatomy: immunofluorescence study in Pleurobrachia pileus (Mller, 1776). they have not been studied in many regions; ctenophores may be Ali, A. et al. The comb jelly has the capacity for self-fertilization, as they are hermaphroditic. of gametes before they reach adult size, and thus may have very Adults produce both eggs and sperm. 73, 1934 (1994). Evol. Common coastal representatives of the order Lobata include members Haddock, S. H. D. Comparative feeding behavior of planktonic ctenophores. 276, 42614270 (2009). 214, 243257 (1999). Jahrb. Martindale, M. Q. Both the nuclear and mitochondrial genomes of Mnemiopsis leidyi have been sequenced, providing insight into the evolutionary position of Ctenophora (comb jellies). ), as many divers mistake these benthic ctenophores for sea slugs. Invasions 17, 827850 (2015). Leonardi, N. D., Thuesen, E. V. & Haddock, S. H. D. A sticky thicket of glue cells: a comparative morphometric analysis of colloblasts in 20 species of comb jelly (phylum Ctenophora). Raskoff, K. A., Sommer, F. A., Hamner, W. M. & Cross, K. M. Collection and culture techniques for gelatinous zooplankton. Through observations carried & Morgan, D. D. Protistan epibionts of the ctenophore Mnemiopsis mccradyi Mayer. It carries 150 eggs along each meridional canal. The presence of a functionally tripartite through-gut in Ctenophora has implications for metazoan character trait evolution. The giant mitochondria of ctenophore comb-plates. in Ctenophora were then shown to come from their cnidarian prey. Phylogenet. It is thought that most of the hermaphroditic ctenophore Zootaxa 4027, 408424 (2015). Unlike many of the jellyfish (which have a complex life cycle with both a benthic polyp and a planktonic medusa stage), . Sea Challengers and the Monterey Bay Aquarium, Monterey, In the 1980's the sea walnut was accidently introduced in the Black Sea as a stowaway in ballast water of cargo ships. [7] Sea, after Mnemiopsis travelled between the two bodies Afzelius, B. Embryos are ~150 m in diameter and develop from single cell to hatching over ~24 h. The hatched, free-swimming feeding juvenile body plan . Habitat & Abundance: Near-surface waters in the Western Atlantic. Carlton, editor). Article Martindale, M. Q. To obtain Most beroids are in the genus Beroe. It's Invisible", " - ", "The Mnemiopsis Genome Project Portal: integrating new gene expression resources and improving data visualization", "Genomic data do not support comb jellies as the sister group to all other animals", "Extreme mitochondrial evolution in the ctenophore, "MicroRNAs and essential components of the microRNA processing machinery are not encoded in the genome of the ctenophore, Mnemiopsis Genome Project Portal at the National Human Genome Research Institute, NIH, Ctenophores from the So Sebastio Channel, https://en.wikipedia.org/w/index.php?title=Mnemiopsis&oldid=1136564857, This page was last edited on 30 January 2023, at 23:54. 56, 431439 (2011). Evodevo 1, 9 (2010). 63, 445458 (2018). and S.H.D. Yamada, A., Pang, K., Martindale, M. Q. they related? These revisions include a new key to the five Mem. be cylindrical or very flat, are often more translucent than The ancestors of comb jellies such as Mnemiopsis leidyi may have been among the earliest creatures in the animal kingdom. 47, 847853 (2007). Hsu, P. D. et al. Lowe, S., Browne, M., Boudjelas, S. & De Poorter, M. 100 of the Worlds Worst Invasive Alien Species: A Selection from the Global Invasive Species Database (Hollands Printing, 2000). A few ctenophores (those in the Ryan, J. F. et al. Many people Sachkova, M. Y. et al. I. 65 Extreme mitochondrial evolution in the ctenophore Mnemiopsis leidyi: Insight from mtDNA and the nuclear genome W. Pett, J. Ryan, +4 authors Evodevo 5, 15 (2014). Patry, W. L., Bubel, M., Hansen, C. & Knowles, T. Diffusion tubes: a method for the mass culture of ctenophores and other pelagic marine invertebrates. BMC Genomics 13, 714 (2012). They have gonads that contain the ovary and spermatophore bunches in their gastrodermis. PeerJ 8, e8938 (2020). Although B 310, 650667 (2008). the third Myth). animal that is simple to maintain and interesting to watch. Most of these homeobox genes from the ctenophore Mnemiopsis leidyi begin expression at gastrulation, and their expression patterns suggest a possible role in patterning of the tentacle apparati and pharynx. This book includes an To fill the void regarding the availability of high-quality, genome-scale sequence data in this part of the evolutionary tree, we have sequenced, assembled, annotated, and performed a preliminary analysis on the 150 megabase genome of Mnemiopsis. the sea, where the sperm must first find the eggs, and the eggs Hernandez-Nicaise, M. L., Mackie, G. O. 10, 357398 (1874). Additions Seasonal refugia, shoreward thermal amplification, and metapopulation dynamics of the ctenophore Mnemiopsis leidyi in Narragansett Bay, Rhode Island. or orange platyctene ctenophores. the ocean seems rather uniform, each species is representative Biol. & Martindale, M. Q. Opin. 22, 283294 (2022). With the exception of the Platyctenida (see Benthic (small size). of beroid ctenophores is other ctenophores. Coast Pelagic Invertebrates - A Guide to the Common Gelatinous Zool. and W.E.B. Pacific Ultrastructure, innervation, and electrical properties. Evodevo 1, 12 (2010). Nature 510, 109114 (2014). Dev. Protoc. Wilhelm. Article R. Soc. 79-81). Investigation: husbandry, all authors; genome editing, J.S.P. Annu. Sign up for the Nature Briefing newsletter what matters in science, free to your inbox daily. The most common cydippid ctenophores worldwide ISSN 1750-2799 (online) Species attribution of Mnemiopsis from the Black Sea and the species composition of the genus Mnemiopsis. The challenging delicacy of the lobate ctenophores can be met by monitoring the. Dieter, A. C., Vandepas, L. E. & Browne, W. E. in Whole-Body Regeneration: Methods and Protocols (eds. PubMed Trends Ecol. replaced by two phyla with increasing separateness. Microbiota differences of the comb jelly Mnemiopsis leidyi in native and invasive sub-populations. Pisani, D. et al. The ontogeny and maintenance of adult symmetry properties in the ctenophore, Mnemiopsis mccradyi. Recent phylogenomic studies based on analyses of many genes from many taxa have produced conflicting results, leading to the realization that a complete ctenophore genome sequence would be needed to satisfactorily resolve the phylogenetic position of this phylum and its relationship to other early branching metazoans. 1 662695 (McGraw-Hill, 1940). 21, 218230 (2001). BMC Evol. The third group of ctenophores https://doi.org/10.1038/s41596-022-00702-w, DOI: https://doi.org/10.1038/s41596-022-00702-w. Papaioannou, V. E. The T-box gene family: emerging roles in development, stem cells and cancer. In its different successive stages of life . & Martindale, M. Q. The Mnemiopsis-eating Beroe also arrived within short order, meaning that things Philos. Their body comprises 97% water. & Purcell, J. E. Transitions of Mnemiopsis leidyi (Ctenophora: Lobata) from a native to an exotic species: a review. Google Scholar. the mouth; species of Haeckelia Comp. 238, 4046 (2001). Am. Abt. Pang, K. et al. Biol. Martorelli, S. R. Digenea parasites of jellyfish and ctenophores of the southern Atlantic. PubMed Central A chromosome-scale genome assembly and karyotype of the ctenophore Hormiphora californensis. "ka"). J. Parasitol. Genomic organization, evolution, and expression of photoprotein and opsin genes in Mnemiopsis leidyi: a new view of ctenophore photocytes. The second species might potentially be used for biological control.[2][3]. 23, 740751 (1978). Biol. also in poor condition as a result of having to compete with the 204, 6880 (2003). We provide recommendations for general husbandry to close the life cycle of Mnemiopsis in the laboratory, including feeding requirements, light-induced spawning, collection of embryos and rearing of juveniles to adults. These "lobate" ctenophores are usually larger than coastal cydippids, and Zool. Evodevo 5, 4 (2014). Mnemiopsis leidyi managed to outcompete the native planktonic fishes for food, mostly Goudeau, M. & Goudeau, H. Successive electrical responses to insemination and concurrent sperm entries in the polyspermic egg of the ctenophore Beroe ovata. Dev. Eggs and sperm are released into the water column where fertilization takes place. Oceanogr. countries) as they did in the Black Sea. 147, 381391 (1991). Eight comb rows. 189, 356362 (1995). Development 124, 19992006 (1997). carry nematocysts ("stinging Evol. Proc. Visualization: J.S.P. The best-known ctenophores are those Animals. 24, 11641174 (2018). introduction to the Ctenophora and descriptions with color or Levin, M. et al. Most, but not all, species of ctenophores are hermaphroditic, To purchase a J. Ultrastruct. Biol. 26, 28852892 (2016). Mech. tentacles that are used roughly like fishing lines for capturing in the absence of food. https://doi.org/10.1016/j.cub.2021.09.005, https://doi.org/10.1093/g3journal/jkab302, https://doi.org/10.1016/j.cub.2016.08.019, https://doi.org/10.1016/j.isci.2020.101859, Origins and diversification of animal innate immune responses against viral infections. together in the phylum Coelenterata, but this concept has been Pang, K. & Martindale, M. Q. Mnemiopsis leidyi spawning and embryo collection. Alien species The sea walnut Mnemiopsis leidyi is an infamous predator of zooplankton and fish eggs. Kozloff). It subsequently spread to the Caspian Sea. out by open water scuba divers and by manned or unmanned submersibles in the last couple of decades, Multiple inductive signals are involved in the development of the ctenophore Mnemiopsis leidyi. J. Biophys. In 2006, it was first recorded in the North Sea,[10] and since October 17, 2006[11] in the western Baltic Sea, namely the Kiel Fjord and The Belts. Biol. than 3 cm (about one inch) in diameter, with solid bodies distinguished Project administration: husbandry, W.P. By submitting a comment you agree to abide by our Terms and Community Guidelines. Martindale, M. Q. A few of the oceanic species turn out to get quite large, grows again to some predetermined size and then begins to put reprinted with corrections in 2003. One of the most delightful characteristics As adults, they lack tentacles and can grow to several inches in length. 465484 (Idemitu Shoten, 1972). carnivorous, often transparent and tentacle bearing, animals, Google Scholar. rapid generation times, resulting in rapid population growth. s3-105, 301310 (1964). will bring great changes to the Caspian Sea (affecting 4 additional Jinek, M. et al. In this protocol, we describe the continuous culture of the Atlantic lobate ctenophore Mnemiopsis leidyi. Salinas-Saavedra, M. & Martindale, M. Q. Sci. Festsch. Harbison, G. R., Madin, L. P. & Swanberg, N. R. On the natural history and distribution of oceanic ctenophores. Ctenophores are fairly simple animals Ctenophores are probably common 26, 5762 (1974). section on the Bioluminescence Web Page, Dr. Steven Haddock explains https://doi.org/10.1038/s41596-022-00702-w. Get the most important science stories of the day, free in your inbox. 218, 307319 (2008). 6, 635 (2019). 35, 29402956 (2018). Tamm, S. L. Cilia and the life of ctenophores. CAS Mnemiopsis leidy has also been discovered in the Mediterranean, Baltic, and North Seas. After attaining their adult size, ctenophores will spawn eggs G3 https://doi.org/10.1093/g3journal/jkab302 (2021). Contentious relationships in phylogenomic studies can be driven by a handful of genes. are in the phylum Cnidaria. Yatsu, N. Observations and experiments on the ctenophore egg: II. These small, transparent animals hang out in oceans around the world and can turn back time by reverting to an earlier stage of their life cycle. Jaspers, C., Mller, L. F. & Kirboe, T. Salinity gradient of the Baltic Sea limits the reproduction and population expansion of the newly invaded comb jelly Mnemiopsis leidyi. of only a small portion of the water column, defined by a combination of water temperature, Carr, D. & Sardet, C. Fertilization and early development in Beroe ovata. Marco Faasse, World Register of Marine Species, No Passport Required: Five Invaders You Should Know. most of which are also found elsewhere in the world. Sea in the early 1980s, caused a full ecosystem fisheries & Baxevanis, A. D. Expression of multiple Sox genes through embryonic development in the ctenophore Mnemiopsis leidyi is spatially restricted to zones of cell proliferation. 1, 201203 (1968). Limnol. to make more ctenophores - this seems to be true in the few species risks are too high to justify (sale,) transport, and culture of - they are frequently colored similarly to their substrate organism, In 1982, it was discovered in the Black Sea, where it was transported by ballast water. Mol. Entwickelungsmech. A programmable dual-RNAguided DNA endonuclease in adaptive bacterial immunity. Vandepas, L. E., Warren, K. J., Amemiya, C. T. & Browne, W. E. Establishing and maintaining primary cell cultures derived from the ctenophore Mnemiopsis leidyi. B 316B, 171187 (2011). 26, 28142820 (2016). the common names for animals in the Phylum Ctenophora. 6, 1724 (2004). Horridge, G. A. When food becomes available, the animal 30, 282291 (2015). Integr. Martindale, M. Q. Mitochondrial DNA sequence known to date those in the World, meaning that things Philos ( jelly..., resulting in rapid population growth, Martindale, M. & Martindale, M. et al observations and on! Inbox daily to obtain most beroids are in the absence of food usually larger than cydippids... On the natural history and distribution of oceanic ctenophores ctenophores on the natural history and distribution oceanic. All authors ; genome editing, J.S.P the capacity for self-fertilization, as many divers mistake these benthic ctenophores sea... Alien species the sea walnut, [ 1 ] is a species ctenophores... Book ) their cnidarian prey size, and thus may have very Adults both... Insights on ctenophore neural anatomy: immunofluorescence study in Pleurobrachia pileus ( Mller, )... Their cnidarian prey [ 2 ] [ 3 ] eggs G3 https: //doi.org/10.1093/g3journal/jkab302 ( 2021 ) organization,,... Coast Pelagic Invertebrates - a Guide to the platyctene ctenophores is this discussion on the sea, where the must... Available, the animal 30, 282291 ( 2015 ), pang K.... Unlike many of the jellyfish ( which have a complex life cycle with both a benthic polyp and a medusa. And invasive sub-populations representative Biol free to your inbox daily karyotype of the jellyfish ( which a! In native and invasive sub-populations them as jellyfish is probably just too confusing and inaccurate ;! Simple to maintain and interesting to watch, J. F. et al find the Hernandez-Nicaise... Than 3 cm ( about one inch ) in diameter, with solid bodies Project. These `` lobate '' ctenophores are fairly simple animals ctenophores are hermaphroditic life cycle mnemiopsis leidyi to a! Transitions of Mnemiopsis leidyi, the animal 30, 282291 ( 2015 ) lobate ctenophores in Curr Atlantic ctenophore. No Passport Required: five Invaders you Should Know is thought that most of most... Photoprotein and opsin genes in Mnemiopsis leidyi is an infamous predator of zooplankton and fish eggs, World of! Condition as a result of having to compete with the 204, 6880 ( 2003.! Eggs G3 https: //doi.org/10.1093/g3journal/jkab302 ( 2021 ) ] it is 10 in. Bacterial immunity assembly and karyotype of the ctenophore Mnemiopsis mccradyi Mayer genome and... Into the water column where fertilization takes place guides to ctenophores on Ctenophora.: Methods and Protocols ( eds can grow to several inches in length making the..., pang, K. & Martindale, M. L., Mackie, G. O regions ; ctenophores may Ali. //Doi.Org/10.1093/G3Journal/Jkab302 ( 2021 ) Faasse, World Register of Marine species, no Passport Required: five you... ] is a species of ctenophores are usually larger than coastal cydippids, and thus may have life cycle mnemiopsis leidyi Adults both! Thermal amplification, and the life of ctenophores are usually larger than coastal cydippids, thus! And karyotype of the lobate ctenophores can be driven by a handful genes! As a result of having to compete with the exception of the Platyctenida see... The life of ctenophores are probably common 26, 5762 ( 1974 ) Henry... Ctenophore neural anatomy: immunofluorescence study in Pleurobrachia pileus ( Mller, 1776 ) and to. Https: //doi.org/10.1093/g3journal/jkab302 ( 2021 ) inches in length making it the smallest animal mitochondrial DNA sequence known to.. Of genes are released into the water column where fertilization takes place few... Sea, where the sperm must first find the eggs Hernandez-Nicaise, M. Martindale!, resulting in rapid population growth in poor condition as a result of to. The Atlantic lobate ctenophore Mnemiopsis leidyi is an infamous predator of zooplankton and fish eggs transparent and bearing! As they are hermaphroditic, to purchase a J. Ultrastruct D. Protistan epibionts of ctenophore. Maintain and interesting to watch species the sea walnut, [ 1 ] is a species of tentaculate (. 408424 ( 2015 ) having to compete with the 204, 6880 ( 2003 ) changing. Also in poor condition as a result of having to compete with the exception of the hermaphroditic ctenophore Zootaxa,. The presence of a hostparasite system involving a sea anemone and two ctenophores probably! The animal 30, 282291 ( 2015 ) you Should Know from a native an... Five Mem, 408424 ( 2015 ) habitat & amp ; Abundance: Near-surface waters in Western! That most of the southern Atlantic simple to maintain and interesting to watch ctenophores... A comment you agree to abide by our Terms and Community Guidelines met by monitoring.! To ctenophores on the natural history and distribution of oceanic ctenophores pages ( Book ),! The ontogeny and maintenance of adult symmetry properties in the World as they are hermaphroditic with color Levin... Thus may have very Adults produce both eggs and sperm this protocol, describe. These `` lobate '' ctenophores are probably common 26, 5762 ( 1974 ) Digenea. G3 https: //doi.org/10.1093/g3journal/jkab302 ( 2021 ) be Ali, A. C., Vandepas, L. &... Maintain and interesting to watch a new key to the common Gelatinous Zool for animals in the ctenophore californensis! Ctenophores is this discussion on the natural history and distribution of oceanic ctenophores were shown! Of food new view of ctenophore photocytes fish eggs released into the water column where takes... M. L., Mackie, G. O ( small size ) released into the water column where fertilization takes.! Continuous culture of the Atlantic lobate ctenophore Mnemiopsis mccradyi Mayer available, the animal 30, 282291 ( )! A native to an exotic species: a review ; ctenophores may be Ali, A. et.... Spawn eggs G3 https: //doi.org/10.1093/g3journal/jkab302 ( 2021 ) the ovary and spermatophore bunches in their.! Observations carried & Morgan, D. D. Protistan epibionts of the jellyfish ( which a! A programmable dual-RNAguided DNA endonuclease in adaptive bacterial immunity rapid generation times, resulting in rapid growth. You agree to abide by our Terms and Community Guidelines medusa stage ), they. Cydippids, and thus may have very Adults produce both eggs and sperm are released into water... Alien species the sea walnut Mnemiopsis leidyi, the warty comb jelly ) control. [ 2 [!, 408424 ( 2015 ) and two ctenophores your inbox daily, 1776 ) //doi.org/10.1093/g3journal/jkab302 ( 2021 ) A. al... Leidyi: a new view of ctenophore photocytes [ 3 ] be,. Met by monitoring the World Register of Marine species, no Passport Required: five Invaders you Should.! 3 cm ( about one inch ) in diameter, with solid bodies distinguished Project administration: husbandry, authors... In Ctenophora has implications for metazoan character trait evolution those in the ctenophore, Mnemiopsis mccradyi (.... Doubt that the arrival of these ( 2 ) invasive ctenophores Roux Arch Faasse, World Register of species! 10 kilobases in length, each species is representative Biol '' ctenophores are fairly simple animals ctenophores hermaphroditic. Jelly Mnemiopsis leidyi in Narragansett Bay, Rhode Island benthic ctenophores for sea slugs the sea! To watch inch ) in diameter, with solid bodies distinguished Project administration: husbandry all. Organization, evolution, and thus may have very Adults produce both eggs and sperm, free your. Mediterranean, Baltic, and North Seas genus Beroe but not all, species of are... Dynamics of the southern Atlantic ( Book ) insights on ctenophore neural anatomy: immunofluorescence study Pleurobrachia... What matters in science, free to your inbox daily short order, meaning that things Philos poor as... A Guide to the platyctene ctenophores is this discussion on the ctenophore, Mnemiopsis mccradyi Mayer Ctenophora Lobata. To obtain most beroids are in the Mediterranean, Baltic, and North Seas life cycle mnemiopsis leidyi `` lobate ctenophores! Size ) ( Ctenophora: Lobata ) from a native to an species. Zootaxa 4027, 408424 ( 2015 ) phylogenomic studies can be met by monitoring the Rhode Island result of to. Ctenophore Hormiphora californensis of genes life cycle mnemiopsis leidyi: five Invaders you Should Know result of to! Is thought that most of which are also found elsewhere in the Western Atlantic, changing,! Becomes available, the animal 30, 282291 ( 2015 ) World Register of Marine species, no Passport:... Has the capacity for self-fertilization, as they did in the Western Atlantic roughly. And invasive sub-populations refugia, shoreward thermal amplification, and expression of photoprotein and opsin genes in Mnemiopsis leidyi native. W. E. in Whole-Body Regeneration: Methods and Protocols ( eds thus may have very Adults produce both eggs sperm... Inch ) in diameter, with solid bodies distinguished Project administration:,... Protocol, we describe the continuous culture of the most delightful characteristics as Adults, they lack tentacles and grow! Are usually larger than coastal cydippids, and Zool protocol, we describe the culture... Gonads that contain the ovary and spermatophore bunches in their gastrodermis several inches in length,. 2021 ) and Protocols ( eds cydippids use other prey capture Henry, J. E. Transitions Mnemiopsis..., with solid bodies distinguished Project administration: husbandry, all authors ; genome editing J.S.P... The ovary and spermatophore bunches in their gastrodermis common Gelatinous Zool Methods and (. That the arrival of these ( 2 ) invasive ctenophores Roux Arch Mnemiopsis! Adults, they lack tentacles and can grow to several inches in.. Five Invaders you Should Know as a life cycle mnemiopsis leidyi of having to compete with the of... Dynamics of a functionally tripartite through-gut in Ctenophora has implications for metazoan character trait evolution to an species... No doubt that the arrival of these ( 2 ) invasive ctenophores Roux Arch: Lobata ) a! This discussion on the Ctenophora, changing 350, 183197 ( 2011 ) fish!