Kudoa is a genus of Myxozoa and the only genus recognized within the monotypic family Kudoidae. There are approximately 100 species of Kudoa all of which parasitize on marine and estuarine fish. Kudoa are most commonly known and studied for the negative effects the genus has on commercial fishing and aquaculture industries.
Description
The genus Kudoa is identified by the possession of four or more shell valves composed of a fragile membrane and arranged in a or stellate pattern. The maximum number of shell valves in any described Kudoa species is 13. Each of these valves has a polar capsule. The genus Kudoa was originally part of the genus Chloromyxum because of the distribution of their polar capsules, however, it was later determined to be a separate genus. Each Kudoa has two sporoplasm cells, one enclosed by the other. Most Kudoa are histozoic parasites, with a few species being described as coelzoic. Some taxonomists question whether these coelzoic organisms belong to a separate genus. There are approximately 100 described species of Kudoa which can be found in the Atlantic, Pacific, and Indian oceans.
Species
The following species are recognized in the genus Kudoa:
- Kudoa aburakarei <small>Li, Inoue, Tanaka, Zhang & Sato, 2020</small>
- Kudoa acentrogobia <small>Li, Inoue, Tanaka, Zhang & Sato, 2020</small>
- Kudoa aequidens <small>Casal, Matos, Matos & Azevedo, 2008</small>
- Kudoa ajurutellus <small>Araújo-Neto, Cardim, Da Silva, Hamoy, Matos & Abrunhosa, 2020</small>
- Kudoa akihitoi <small>Kasai, Setsuda & Sato, 2017</small>
- Kudoa alliaria <small>Kovaleva, Shulman & Yakolev, 1979</small>
- Kudoa amamiensis <small>Egusa & Nakajima, 1978</small>
- Kudoa amazonica <small>Velasco, Sindeaux Neto, Videira, de Cássia Silva do Nascimento, Gonçalves & Matos, 2019</small>
- Kudoa anatolica <small>ÃÂzer, Okkay, Gürkanlñ, ÃÂiftçi & Yurakhno, 2018</small>
- Kudoa azevedoi <small>Mansour, Thabet, Chourabi, Harrath, Gtari, Omar & Hassine, 2013</small>
- Kudoa azoni <small>Aseeva, 2004</small>
- Kudoa barracudai <small>Abdel-Baki, Al-Quraishy, Omar & Mansour, 2016</small>
- Kudoa bengalensis <small>Sarkar & Mazumder, 1983</small>
- Kudoa boopsi <small>Kpatcha, Diebakate, Faye & Toguebaye, 1999</small>
- Kudoa bora <small>Fujita, 1930</small>
- Kudoa borimiri <small>Yurakhno, Slynko, Chinh, Ha & Whipps, 2022</small>
- Kudoa branchiata <small>Joy, 1972</small>
- Kudoa camarguensis <small>Pampoulie, Marques, Crivelli & Boucherau, 1999</small>
- Kudoa carcharhini <small>Gleeson, Bennett & Adlard, 2010</small>
- Kudoa cascasia <small>Sarkar & Chaudry, 1996</small>
- Kudoa caudata <small>Kovaleva & Gaevskaya, 1983</small>
- Kudoa cerebralis <small>Paperna & Zwerner, 1974</small>
- Kudoa chaetodoni <small>Burger, Cribb & Adlard, 2007</small>
- Kudoa cheilodipteri <small>Heiniger, Cribb & Adlard, 2013</small>
- Kudoa chilkaensis <small>Tripathi, 1951</small>
- Kudoa ciliatae <small>Lom, Rohde & Dyková, 1992</small>
- Kudoa clupeidae <small>Hahn, 1917</small>
- Kudoa cookii <small>Heiniger, Cribb & Adlard, 2013</small>
- Kudoa corniculata <small>Li, Inoue, Zhang & Sato, 2022</small>
- Kudoa crenimugilis <small>Abdel-Baki, Abdel-Haleem, Al-Quraishy, Azevedo & Mansour, 2018</small>
- Kudoa cruciformum <small>Matsumoto, 1954</small>
- Kudoa crumena <small>Iversen & van Meter, 1967</small>
- Kudoa cynoglossi <small>Obiekezie & Lick, 1994</small>
- Kudoa dianae <small>Dyková, Fajer & Fiala, 2002</small>
- Kudoa dicentrarchi <small>SitjÃÂ -Bobadilla & Alvarez-Pellitero, 1992</small>
- Kudoa empressmichikoae <small>Kasai, Setsuda & Sato, 2017</small>
- Kudoa encrasicoli <small>Iglesias, Rangel, Fernández-Vázquez, Santos & GarcÃÂa-Estévez, 2022</small>
- Kudoa eugerres <small>Casal, Soares, Rocha, Silva, Santos, Nascimento, Oliveira & Azevedo, 2019</small>
- Kudoa fujitai <small>Li, Inoue, Zhang & Sato, 2020</small>
- Kudoa funduli <small>Hahn, 1915</small>
- Kudoa grammatorcyni <small>Adlard, Bryant, Whipps & Kent, 2005</small>
- Kudoa guangdongensis <small>Li, Inoue, Tanaka, Zhang & Sato, 2020</small>
- Kudoa gunterae <small>Burger & Adlard, 2009</small>
- Kudoa haridasae <small>Sarkar & Ghosh, 1991</small>
- Kudoa hemiscylli <small>Gleeson, Bennett & Adlard, 2010</small>
- Kudoa hexapunctata <small>Yokoyama, Suzuki & Shirakashi, 2014</small>
- Kudoa hirsuta <small>Li, Inoue, Zhang & Sato, 2022</small>
- Kudoa histolytica <small>Pérard, 1928</small>
- Kudoa hypoepicardialis <small>Blaylock, Bullard & Whipps, 2004</small>
- Kudoa igami <small>Shirakashi, Yamane, Ishitani, Yanagida & Yokoyama, 2014</small>
- Kudoa igori <small>Yurakhno, Slynko, Chinh, Ha & Whipps, 2022</small>
- Kudoa iidae <small>Li, Inoue, Tanaka, Zhang & Sato, 2020</small>
- Kudoa inornata <small>Dykova, de Buron, Fiala, Roumillat, 2009</small>
- Kudoa insolita <small>Shulman & Kovalijova, 1979</small>
- Kudoa intestinalis <small>Maeno, Magasawa & Sorimachi, 1993</small>
- Kudoa islandica <small>Kristmundsson & Freeman, 2014</small>
- Kudoa iwatai <small>Egusa & Shiomitsu, 1983</small>
- Kudoa javaensis <small>Yunus, Yustinasari, Natalia, Ghosh, Sakuma, Inoue & Sato, 2021</small>
- Kudoa kabatai <small>Shulman & Kovalijova, 1979</small>
- Kudoa kenti <small>Burger & Adlard, 2009</small>
- Kudoa konishiae <small>Sakai, Kato, Sakaguchi, Setsuda & Sato, 2018</small>
- Kudoa lateolabracis <small>Yokoyama, Whipps, Kent, Mizuno & Kawakami, 2004</small>
- Kudoa leiostomi <small>Dyková, Lom & Overstreet, 1994</small>
- Kudoa lemniscati <small>Miller & Adlard, 2012</small>
- Kudoa leptacanthae <small>Heiniger & Adlard, 2012</small>
- Kudoa lethrini <small>Burger, Cribb & Adlard, 2007</small>
- Kudoa longichorda <small>Inoue, Kasai, Argamjav & Sato, 2022</small>
- Kudoa lunata <small>Lom, Dyková & Lhotákova, 1983</small>
- Kudoa lutjanus <small>Wang, Huang, Tsai, Cheng, Tsai, Chen, Chen, Chiu, Liaw, Chang & Chen, 2005</small>
- Kudoa megacapsula <small>Yokoyama & Itoh, 2005</small>
- Kudoa miniauriculata <small>Whitaker, Kent & Sakanari, 1996</small>
- Kudoa minithyrsites <small>Whipps, Adlard, Bryant, Lester, Findlay & Kent, 2003</small>
- Kudoa mirabilis <small>Naidenova & Gaevskaya, 1991</small>
- Kudoa monodactyli <small>Gunter, Cribb, Whipps & Adlard, 2006</small>
- Kudoa muscularis <small>Cheung, Nigrellu & Ruggieri, 1983</small>
- Kudoa musculoliquefaciens <small>Matsumoto & Arai, 1954</small>
- Kudoa neothunni <small>Arai & Matsumoto, 1953</small>
- Kudoa neurophila <small>Grossel, Dyková, Handlinger & Munday, 2003</small>
- Kudoa niluferi <small>ÃÂzer, Okkay, Gürkanlñ, ÃÂiftçi & Yurakhno, 2018</small>
- Kudoa nova <small>Naidenova, 1975</small>
- Kudoa obicularis <small>Azevedo, Rocha, Matos, Oliveira, Matos, Al-Quraishy & Casal, 2015</small>
- Kudoa ocellatus <small>Da Silva, Da Silva, Lima, Matos, De Carvalho Sanches, Matos & Hamoy, 2022</small>
- Kudoa ogawai <small>Yokoyama, Yanagida & Shirakashi, 2012</small>
- Kudoa ovivora <small>Swearer & Robertson, 1999</small>
- Kudoa pagrusi <small>Al Quraishy, Koura, Abdel-Baki, Bashtar, El Deed, Al Rasheid & Abdel Ghaffar, 2008</small>
- Kudoa paniformis <small>Kabata & Whitaker, 1981</small>
- Kudoa paralichtys <small>Cho & Kim, 2003</small>
- Kudoa paraquadricornis <small>Burger & Adlard, 2009</small>
- Kudoa parathyrsites <small>Kasai, Li, Mafie & Sato, 2016</small>
- Kudoa parvibulvosa <small>Li, Inoue, Zhang & Sato, 2022</small>
- Kudoa pericardialis <small>Nakajima & Egusa, 1978</small>
- Kudoa permulticapsula <small>Whipps, Adlard, Bryant & Kent, 2003</small>
- Kudoa peruvianus <small>Mateo Salas, 1972</small>
- Kudoa pleurogrammi <small>Kasai, Li, Mafie & Sato, 2016</small>
- Kudoa prunusi <small>Meng, Yokoyama, Shirakashi, Grabner, Ogawa, Ishimaru, Sawada & Murata, 2011</small>
- Kudoa quadratum <small>Thélohan, 1895</small>
- Kudoa quadricornis <small>Whipps, Adlard, Bryant & Kent, 2003</small>
- Kudoa quraishii <small>Mansour, Harrath, Abd-Elkader, Alwasel, Abdel-Baki & Al Omar, 2014</small>
- Kudoa ramsayi <small>Kalavati, Brickle & MacKenzie, 2000</small>
- Kudoa rayformis <small>Shin, Shirakashi, Hamano, Kato, Lasso & Yokoyama, 2016</small>
- Kudoa rosenbuschi <small>Gelormini, 1966</small>
- Kudoa rousseauxii <small>Velasco, Eduard, Neto, Dias, Matos & Gonçalves, 2022</small>
- Kudoa saudiensis <small>Mansour, Harrath, Abdel-Baki, Alwasel, Al-Quraishy & Al Omar, 2015</small>
- Kudoa schulmani <small>Naidenova & Zaika, 1970</small>
- Kudoa sciaenae <small>Teran, Llicán & Lugue, 1990</small>
- Kudoa scomberi <small>Li, Sato, Tanaka, Ohnishi, Kamata & Sugita-Konishi, 2013</small>
- Kudoa scomberomori <small>Adlard, Bryant, Whipps & Kent, 2005</small>
- Kudoa sebastea <small>Aseeva, 2004</small>
- Kudoa septempunctata <small>Matsukane, Sato, Tanaka, Kamata & Sugita-Konishi, 2010</small>
- Kudoa shiomitsui <small>Egusa & Shiomitsu, 1983</small>
- Kudoa shkae <small>Dyková, Lom & Overstreet, 1994</small>
- Kudoa sphyraeni <small>Narasimhamurti & Kalavati, 1979</small>
- Kudoa stellula <small>Yurakhno, 1991</small>
- Kudoa surabayaensis <small>Yunus, Yustinasari, Natalia, Ghosh, Sakuma, Inoue & Sato, 2021</small>
- Kudoa tachysurae <small>Sarkar & Mazumder, 1983</small>
- Kudoa tetraspora <small>Narasimhamurti & Kalavati, 1979</small>
- Kudoa thalassomi <small>Adlard, Bryant, Whipps & Kent, 2005</small>
- Kudoa thunni <small>Matsukane, Sato, Tanaka, Kamata & Sugita-Konishi, 2011</small>
- Kudoa thyrsites <small>Gilchrist, 1924</small>
- Kudoa trachuri <small>Matsukane, Sato, Tanaka, Kamata & Sugita-Konishi, 2011</small>
- Kudoa trifolia <small>Holzer, Blasco-Costa, Sarabeev, Ovcharenko & Balbuena, 2006</small>
- Kudoa unicapsula <small>Yurakhno, Ovcharenko, Holzer, Sarabeev & Balbuena, 2007</small>
- Kudoa viseuensis <small>Monteiro, Da Silva, Hamoy, Sanches & Matos, 2019</small>
- Kudoa whippsi <small>Burger & Adlard, 2009</small>
- Kudoa yasai <small>Cardim, Araújo-Neto, da Silva, Hamoy, Matos & Abrunhosa, 2020</small>
- Kudoa yasunagai <small>Hsieh & Chen, 1984</small>
Development
In Myxozoan development, the Myxosporean life-stage develops inside a fish host, while the Actinosporean life-stage develops in an annelid host. Fully-developed Myxospores are consumed by annelids and reproduce asexually via schzogony in the gut epithelium of worms. Gametes are formed in the gut of the worm and these gametes fuse together to create eight zygotes. The zygotes become spores with three valves, 3 polar capsules, and a sporoplasm that are released in the worm's feces and attach to the surface of a fish host. Once attached to the fish, the gamete injects the sporoplasm cell into the fish. The sporoplast divides forming a fully-developed Myxosporean.
Diet
Kudoa parasitize on marine and estuarine fish. Kudoa typically feed on the skeletal muscle tissue, although some may feed on other parts of the body such as the central nervous system, heart, intestines, ovaries, or gills. Most Kudoa are histozoic parasites, however, a few species are coelozoic. The genus Kudoa attack a wide range of hosts, however, individual species only feed on specific hosts. Kudoa feed via pinocytosis across the host-parasite membrane. One species of Kudoa, K. thyrsites, are distributed worldwide and are believed to have been reported in over 20 different species of fish but it is possible that the species described as K. thyrsites is actually multiple different species.
Implications for the fishing industry
Kudoa are most well known for the economic loss they cause for the commercial fishing and aquaculture industries. When Kudoa attach to hosts, they leave unsightly cysts that lower the price fish can be sold for at market. Kudoa also release proteolytic enzymes that degenerate muscle in fish to aid in their own growth and development, further decreasing the fish's value. Some species in the genus Kudoa with the most notable effects of the commercial fishing and aquaculture industries are: K. musculoliquefaciens in Broadbill Swordfish (Xiphias gladius), K. thyrsites in Atlantic Salmon (Salmo salar), K. clupeidae in Atlantic Herring (Clupea harengus), K. septempunctata in Olive Flounder (Paralichthys olivaceus), K. thunni in Yellowfin Tuna (Thunnus albacares) and K. paniformis in Pacific Hake (Merluccius productus).
Some studies have found evidence to suggest that some species of Kudoa are linked to food-borne illness in humans.
References
Encyclopedia of Life