Invasive Species Profile

Oriental Weatherfish, Misgurnus anguillicaudatus (Candor, 1824)

Ben Frable

FISH 423

December 3rd, 2008

Diagnostic Information are imbedded and irregularly distributed along the body; Berg (1949) gave scale counts of 105- Common names: Oriental weatherfish, oriental 180 laterally. Another characteristic to weather loach, weather loach, dojo and dojo distinguish it from other species is that it has loach elongated peduncular keels with pre-current caudal rays creating conspicuous adipose crests Classification: on the dorsal and ventral sides of the caudal Kingdom: Animalia peduncle, see Fig. 1b (Berg 1949; Kottelat Phylum: Chordata 1998). The caudal peduncle is also fairly short Class: Actinopterygii and less than the length of the head (Nichols Order: Cypriniformes 1943). On many specimens there is a large Family: Cobitidae pigmented spot located dorsally above the base Genus: Misgurnus of the caudal fin (Berg 1949). Also, it has been Species: anguillicaudatus noted that the eyes of M. anguillicaudatus are visibly larger than those of the European Basic identification key weather loach Misgurnus fossilis (Berg 1949). This species is also somewhat sexually The identification of the oriental dimorphic with males having pectoral fins that weatherfish is fairly distinct; however, no are longer than their head. Also, the females modern meristic analysis or comparisons have tend to have a fuller abdomen than males that is been done on the genus. In general, the species mainly associated with the large size of their in long with an anguillid body shape having a ovaries (Yamamoto and Tagawa 2000). A less ventrally located mouth with ten rostral barbells, noticeable character of the males is that the 4 of which are receded. The dorsal fin placed ventral base of the second pectoral ray broadens two-thirds the length of the body posteriorally into a rounded plate and covers the bases of the and the pectoral fins are readily placed ventrally other pectoral rays (Berg 1949). behind the opercular opening. The body color is In Oregon, Logan et al. (1996) collected one yellow-brown to brown with irregularly specimen of the Chinese fine-scaled loach distributed areas of pigmentation without Misgurnus mizolepis, another non-native species distinct lateral banding (Berg 1949). Nichols that is closely related to M. anguillicaudatus. (1943) provides a dichotomous key for This species can be readily distinguished from identifying M. anguillicaudatus from other the oriental weatherfish because it has a great Misgurnus species. Some of the characteristics number of fine surface scales covering its body he lists that are used in the key are that the scales rather than irregularly embedded scales (Rendahl 1934). Also, the caudal fin of M. These fish, like many other cypriniform fishes, mizolepis is greatly reduced in relation to its are generally considered r-selected organisms in body length in comparison with that of M. that they mature quickly, reproduce multiple anguillicaudatus. Although one specimen was times and produce a great number of offspring. found in Oregon, no other specimens have since Members of this species are sexually distinct and been collected there or anywhere else in the reproduce sexually. Males of the species mature United States, so it can be assumed at this point rapidly within a year and females mature within this related species has not established non- one or two years (Lei and Wang 1990). In their native populations (Tabor pers. comm.). native range, these fish spawn multiple times per year during a spawning season that lasts from Life history and basic ecology mid-April until mid-October. However, in established non-native populations in Indiana, Life Cycle and Reproductive Habits females with ripe ova were collected in mid- M. anguillicaudatus inhabits still water, June and based on other physiological characters shallow pools and near shore shallow water it appears that they had just started the breeding mudflat areas of lakes, rivers, streams, swamp, season (Simon et al 2006). This could indicate rice patties and ponds. In these environments it that these fish are altering their breeding season is commonly found burrowed into mud, silt, based on different environmental conditions in macrophyte beds or leaf detritus (Simon et al. their non-native ranges. 2006). In other areas where it has invaded, such Reproduction occurs at night in which the male as Hawaii, it has been found under mats of locates a female based on scent and wraps his honohono (Commelina diffusa) and California body around the female stimulating her to grass (Brachiara nuatica) (Simon et al. 2006). release a cloud of eggs that he rapidly fertilizes This species is primarily nocturnal, spending (Yamamoto and Tagawa 2000). The female most of the day burrowed into mud from which produces 1800-15500 eggs per batch with an it emerges at night to feed and mate. However, average survival of 2000 eggs per batch (Berg like their name would suggest, these fish 1949; Suzuki 1983). The eggs are small with a become extremely active right before periods of diameter of 800-850 m and are reddish in color severe weather due to changes in barometric with adhesive properties, however, are broadcast pressure and it is interesting to note that these into the water column upon conception (Suzuki fish were used by ancient Chinese scientists and 1983). Eggs hatch within 30 hours and larvae Victorian era Europeans to predict the weather remain in water column for a short period before (Sterba 1983). settling on the bottom (Wang et al 2008). By combining all of these reproductive characters, Berg (1949) was able to determine the minimum weatherfish devouring the eggs of endemic population doubling time for this species to be fishes, very little evidence has been shown to 1.4-4.4 years, which is fairly short for fish support this and none of the fish collected by species. A vast amount of much more thorough Tabor et al. (2001) had fish eggs in their gut information on the mating habits and cavities. development of M. anguillicaudatus can be found in Chinese literature sources, that the Environmental optima and tolerances author was unable to translate. Some of the characters that make the oriental weatherfish a very successful invader, is Feeding Habits its ability to tolerate and thrive in a very wide Oriental weatherfish are benthic range of temperature and oxygen conditions. omnivores that feed generally on insect larvae, Since M. anguillicaudatus occupies a natural snails, worms, ostracods, cladocerans, fish eggs, range that extends all the way from southern algae and detritus (Sterba 1983). Very little is Siberia to the northern Indochinese Peninsula, known about their feeding habits in their natural they have to be able to tolerate cold winters and range and most of the diet data comes from gut warm summers. In 1996, Logan et al. evaluated content analysis of invasive populations the thermal tolerances of individuals collected in elsewhere in the world. However, Kim et al. Clackamas County, OR by acclimating them to (1994) demonstrated that M. anguillicaudatus 18-20C for six months then the water readily and efficiently consume mosquito larvae temperature was lowered 2C daily until a and have successfully used them to control temperature of 2C was reached. At this mosquito populations in South Korea. Tabor et temperature, the fish were still alive, but al. (2001) found that in their invaded range in required prodding to elicit movement. This Washington, that their diets primarily consisted same study demonstrated that the fish were of crustaceans, such as: amphipods and easily able to tolerate abrupt temperature spikes cladocerans, as well as, chrionomids with from 6C to 18C and within 24h were observed minimal detritus and plant matter. However, in feeding again. Also, their thermal tolerances fit a study by Burchmore et al. (1990), non-native with the average annual temperature fluctuations fish in Australia were found with higher levels of the Snake, Willamette and Sacramento rivers of algae and detritus in their stomachs. Logan et in OR which range from 3.7C to 19.3C (Logan al. (1996) found that these fish did consume et al. 1996). Based on their presence in shallow mosquitofish, Gambusia affinis, fry as well as ponds in tropical regions, it can be assumed that commercially prepared salmon eggs in aquaria. they are also able to tolerate temperatures up to Although there are concerns of non-native and maybe over 30C. This wide thermal tolerance range means that this species is able to external environment allow these fish to survive over winter in many regions and may even be in very small bodies of still water and in hypoxic able to survive in iced over bodies of water swamp and still water environments. This (Fryhof and Korte 2005). means they can also inhabit areas of high In addition to their ability to tolerate a large turbidity and eutrophication and bodies of water thermal range, they are also able to survive in that are affected by algal blooms. low oxygen and hypoxic conditions. This ability These fish are found in freshwater is due in part to the fact that, like other basal environments; however, they have not been teleost fishes, such as, killifish and mudfish, evaluated for their ability to survive in brackish they are able to process atmospheric oxygen. or saltwater. In addition, they have not been They do this by taking air into their mouth and evaluated for their tolerance of variances in pH; diverting it into their anterior alimentary canal of however, since they have been found in stagnant their intestine. There the oxygen is diffused water, rice paddies and swamps, it can be over a membrane of dense capillary beds and is assumed they can inhabit low pH environs (Berg able to diffuse into their blood stream 1949). (McMahon and Burggren 1987). The same paper demonstrated that as the amount of Biotic Associates dissolved oxygen decreased, the amount of air Like almost all fish species, oriental intake by the loaches increase with a similar weatherfish have a number of associated trend found as temperature increased. In parasites and pathogens in their native range. addition to their ability to process atmospheric No commensals are known for this species. oxygen, they are also able to exchange dissolved However, virtually all of the data on their oxygen over their epidermis, with increased parasites and pathogens comes from evaluation rates found while burrowed into moist sediment of individuals in aquaria and collected from their (McMahon and Burggren 1987). Specimens of non-native ranges. In Australia, where most of this group have been collected in exposed mud the research has been conducted, evaluation of and in shallow shore waters with their heads parasites associated with established weatherfish exposed to open air showing that they are able to yielded the presence of three non-native survive for a yet unevaluated period of time out digenean trematode flukes, Echinostoma of water. A specimen (UW113671) that was cinetorchis, E. hortense and Clinostomum collected on the island of Oahu, HI was kept complanatum (Lintermans et al. 1990). These alive in a beaker with 5cm of water in it for two parasites are also not found in the US, and are weeks. These abilities to utilize three different native to Asia. The first two of these parasites methods of extracting oxygen from their can affect humans and cause intestinal problems and C. complanatum affects and rapidly kill Current geographical distribution piscivorous birds by causing lesions to form in their fauces, hindering their feeding behaviors Native range (Aohagi et al. 1992). Lintermans et al (1990) As their common name would suggest, this also found the viral pathogen, birnavirus LV-1 species is naturally distributed in parts of Asia present in some specimens. This viral pathogen from as far north as 53N in southern Siberia, is related to Infectious Pancreatic Necrosis Russia all the way down to Laos and Burma Virus, which as the name suggests, deteriorates around 20S, including Taiwan and the Japanese the pancreas of fishes, eventually killing them archipelago and as far west as India (Berg 1949; and is a major disease of salmonids; therefore, Kapoor et al. 2002). This range is accepted in extremely relevant towards the health of native most publications; however, other publications fish to the state of Washington (Wolf 1988). say it is also native to other countries on the Also in Australia, Dove and Ernst (1998) found Indochinese Peninsula; such as, Cambodia the monogenean Gyrodactylus macracanthus (Tirant 1929), Thailand (Masuda et al. 1984) and attached to the skin of weatherfish. This parasite Vietnam (Kottelat 2001). See Figure 2. attached to fins and skin of the host and causes ulcers and lesions; however, within this genus Global non-native range there is a high level of host-parasite specificity, This species has established breeding so it is unknown if these parasites will affect populations in Australia (Welcomme 1988), other fish in new regions. Germany (Freyhof and Korte 2005), Italy In addition, the parasite Flexibacter columnaris (Razzetti et al. 2001), Mexico (Contreras and that causes lesions on the skin of and can kill Escalante 1984), Palau (Welcomme 1988), many fishes has been found on many of loaches Philippines (Welcomme 1988), Spain (Franch et in Japan and in aquaria (Chowdhury and al. 2008), Turkmenistan (Sal'nikov 1998), and Wakabayashi 1991). This parasite has spread the United States including Hawaii (Yamamoto into the US and affects catfish populations and and Tagawa 2000; Fuller et al. 1999). See could possibly affect other fish populations. As Figure 2. of yet, none of the fishes collected in Washington and many other regions of the US Distribution within the US have not been evaluated for the presence of In the United States, this species was first these pathogens and parasites and whether they established in Hawaii in 1906 (Maciolek 1984), have been transmitted by weatherfish is as of yet then Michigan (Schultz 1960), California (St. undetermined. Amant and Hoover 1969), Florida, Illinois, Idaho, Oregon (Fuller et al. 1999), Washington

Fig. 2: The global distribution of Misgurnus anguillicaudatus, blue indicating native range and red indicating countries with established non-native populations

Fig 3: Invaded range of Misgurnus anguillicaudatus within the United States as of Nov 2008, courtesy of USGS Nonindigenous Aquatic Species database. bait and food. They were first recorded in the wild in Hawaii in 1906 (Welcomme 1988) and were found to have established by the 1930s and 1940s, with a specimen at the University of Washington Fish Collection being collected on Oahu in 1945 in the drainages off of farm irrigation (UW113671) (Maciolek 1984). The fish were first imported on to the mainland in 1939 in Michigan and were found to be established in the wild soon there after in the

Fig. 4: The invasive range of Misgurnus anguillicaudatus 1950s (Schultz 1960). Within a few years, in the state of Washington as indicated in red, courtesy of established populations were also found in USGS Nonidiginous Aquatic Species database southern California (St. Amant and Hoover 1969). Tabor et al. 2001) and Indiana (Simon et al. In 1978, a single specimen was collected in 2006). There are also tentative records in Jovita Creek, a tributary of the White River near Maryland, New Jersey, Louisiana and Tennessee Puyallup; however, it was determined by USGS (NAS). See Figure 3. that this failed to establish since another specimen was never collected at the site Distribution in Washington (Wydoski and Whitney 2003). However, this is In the state of Washington, populations have not the case because five more fish were been established in Lake Washington and its drainages Tabor et al. 2001), the Snohomish drainage Tabor et al. 2001) and on Puget Sound drainages on the Tulalip Indian Reservation (NAS). Individual specimens have also been collected in the Sammamish River and White River (Tabor pers. comm.). See Figure 4.

History of invasiveness

The oriental weatherfish was first imported into Fig. 5: Misgurnus anguillicaudatus specimen regions outside its native range in the 19th UW113571 collected 1994 in pothole near White century, mainly being raised in fish farms for River; Sumner, WA. collected in the lower White River during the fish have been introduced via intentional electrofishing sampling in 1994, one of which is aquarium release only, since there are no fish in the University of Washington Fish Collection farms that stock weatherfish in the state Tabor et as well (Fig 5. UW111106) (Tabor pers. al. 2001). These fish are an extremely popular comm.). No established populations were aquarium fish due to their hardiness and minimal observed in Washington until 1998 when Tabor husbandry requirements and are imported in et al. (2001) found them in Lake Washington. great numbers, with quantifiable numbers as of The next year in 1999, an established population yet undetermined for the US, but were estimated was found in Tulalip Creek, drainage of the as high as 50,000 per year in Australia in the Snohomish and Puget Sound and in 2004, 1980s (Koster et al. 2002), into the United States populations were observed in the Sammamish from Asia or raised in aquaculture facilities and River (Tabor pers. comm.). distributed to many pet stores (Logan et al. 1996). Invasion process For example, a probable vector for this species introduction into Lake Washington can be traced Pathways, vectors and routes of introduction to the University; there is a large pet store that In many invaded regions, the pathway stocks M. anguillicaudatus less than 0.5 mi from for introduction of this species has been escape the Lake were the fish only cost $3.99 each or intentional release from fish farms and more (Nov 2008). It can be assumed that many recently the intentional or accidental release students living in the dormitories will purchase from home aquaria Tabor et al. 2001). In these fish and at the end of the school year when Michigan and California, established they move out they may not be able to continue populations were readily traced back to fish keeping their aquaria, instead of giving the fish farms from which the fish had been released away or euthanizing them, many students will after the farm stopped selling them (Schultz think a humane method of disposal is releasing 1960) or from escape due to flooding events (St. them into Lake Washington which is less than Amant and Hoover 1969). In Hawaii and other 0.1 mi from some of the dormitories. This regions of the world, such as Turkmenistan, this vector is purely hypothetical; however, is a very species has been introduced by Asian likely explanation for their initial introduction immigrants through its use as live bait for into the lake, since, it is known that these fish as fishing (Sal'nikov, 1998) and as a food source well as goldfish Carassius and even piranhas (Maciolek 1984); however, they have not been Piaractus? have been found in fountains on the found being used as live bait in the continental University of Washington campus (Grujon, pers. United States. In Washington, it is believed that comm.). Factors influencing establishment and spread also calculated their average minimum Oriental weatherfish are very well suited population doubling time to just 1.4-4.4 years, for establishment and spread in many locations which is extremely rapid for an invasive species. due to their physiological characteristics and Their main method of dispersal from their have many of the characteristics typical of a establishment sites appears to be via irrigation successful invader (Taylor et al 1984). As systems, stream, river and lake drainages, discussed previously, they have a very wide flooding events and flood control practices thermal tolerance range (2C-30C) which (Logan et al. 1996). In Australia, rapid allows them to survive in the cold winters of expansion of populations has been shown using much of the northern United States and parts of these methods of dispersal (Burchmore et al. Southern Canada, as well as, the hot summers of 1990). In areas of the United States, such as in Southern California, Hawaii and Florida (Logan Michigan and California, their ranges have been et al. 1996). In addition to this characteristic, fairly restricted by limited irrigation channels their ability to utilize atmospheric oxygen can and flooding events (Logan et al. 1996). allow them to survive in low oxygen and However, in Michigan, M. anguillicaudatus was hypoxic environments, such as, areas of still one of three fish species that has a high probably water and high eutrophication, swamps and of spread into the Great Lakes and it appears to under mats of macrophytes (McMahon and be only a matter of time (Rixon et al. 2005). In Burggren 1987). Another factor is that they are Washington, rapid spread via this mechanism generalist omnivores with a diverse and flexible appears to be occurring Tabor et al. 2001). The diet, which allows them to consume many novel specimen that was collected in 1994 near the prey items with ease (Watanabe and Hidaka White River was actually collected in a pothole, 1983). On the other side of this, they also have a located over 500m from the river, which was low vulnerability to predation due to their formed after flood control procedures fossorial habits, such as habitats of choice and (UW113671). Tabor (pers. comm.) found that nocturnal behaviors (Logan et al. 1996). within just two years, oriental weatherfish Their reproductive habits and r-selected populations have spread more than 35km in characteristics also contribute greatly to their Washington. abilities to establish in non-native ranges. They mature somewhat rapidly (1-2 years), have a Potential ecological and/or economic impacts long spawning season and are able to disperse The potential ecological and economic upwards of 15500 eggs in a single spawning, impacts of invasion by this species are not as with an average survival rate of 2000 eggs per drastic or detrimental as they are for other hatch (Berg 1949; Suzuki 1983). Berg (1949) invaders. There are a few main ecological concerns for the invasion of M. blooms and eutrophication of many bodies of anguillicaudatus. The first concern is that they water. However, as before this has not been can predate on the eggs, larvae and fry on native recorded or demonstrated in the wild and species in their established ranges and have been therefore has not been observed in Washington. found to ingest Gambusia fry and salmonid eggs All three of these factors, if realized could be in the laboratory (Logan et al. 1996; Simon et al. economically detrimental, but this has also not 2006). However, none of the specimens been analyzed. collected in Washington showed any fish Despite these negative impacts, a study by Kim material in their gut content Tabor et al. 2001). et al. (1994) demonstrated a few positive The second ecological concern is their ability to impacts that M. anguillicaudatus could transport non-native pathogens and parasites. potentially have. Kim et al. (1994) As stated earlier, weatherfish in Australia have demonstrated in South Korea that weatherfish been found to have introduced and help establish could be effectively used to control and the monogenean Gyrodactylus macracanthus eradicate mosquito populations by preying on which contributed greatly to Australia’s decision mosquito larvae and they could be a potential to ban the species from import (Dove and Ernst biocontrol for diseases such as, malaria and the 1998). Of primary concern in Washington, West Nile virus. In this study, they also however, is their ability to carry the viral demonstrated that these fish were implemented pathogen birnavirus LV-1, which is linked to in rice paddies as weed control because they Infectious Pancreatic Necrosis Virus, which as easily uprooted the shallow roots of pest the name suggests, deteriorates the pancreas of vegetation in rice fields. However, this second fishes, eventually killing them and is a major positive impact is not applicable in the state of disease of salmonids (Wolf 1988; Lintermans et Washington since the climate does not allow for al. 1990). Again, this pathogen has not been rice paddies. Lastly, these fish can be and have found on weatherfish in the state of Washington been used as a food source for humans; (Tabor pers. comm.). The third and final major however, their appeal and market value would ecological concern is that they may disrupt the probably be minimal and not economically benthic structure and facilitate eutrophication advantageous (Maciolek 1984). A good deal of and increase turbidity. Keller and Lake (2007) substantiating evidence appears to be necessary demonstrated in the laboratory that oriental before any real ecological and economic impacts weatherfish did indeed cause significant levels of this species can be demonstrated and of increased turbidity and water column evaluated. nitrogen. Both of these factors could contribute heavily in the facilitation of harmful algal Management strategies, control methods and the establishment of any clearly negative future directions ecological or economic impacts may be necessary to promote active monitoring and In Australia, this species has been control of this species in the United States and in eradicated in watersheds in Queensland by the state of Washington. electrofishing and benthic suction removal and have been completely banned from import since Literature Cited 1986 (Lintermans et al. 1990). However, in all other invaded countries, including the United Aohagi Y, Shibahara T, Machida N et al (1992) States, this species has been largely unmanaged Natural infections of Clinostomum or controlled through any methods or by any complanatum (Trematoda, organized bodies. Many states have monitored Clinostomatidae) in wild heron and egrets, their established populations periodically; Tottori Prefecture, Japan. J Wild Dis however, no long-term and consistent 28:470-471 monitoring programs have been instituted (Rixon et al. 2005). In many states they are Berg LS (1949) Freshwater fishes of the listed as species of concern, but very little U.S.S.R. and adjacent countries, funding is provided for their study and Smithsonian, Washington DC Vol. 2, 4th management. There are also no restrictions on edition, p. 467-468. Translated from their import or sales via the aquarium trade, even Russian though this is their main pathway of introduction. Currently in Washington state, Burchmore J, Faragher R, Thorncraft G (1990) they are not being managed, collected, Occureances of the introduced oriental controlled or researched by any government weather loach (Misgurnus body or research institution and all funding of anguillicaudatus) in the Wingecarribee this nature has been cut as of 2004 (Tabor pers. River, New South Wales. Australian comm.) and their populations have not even Government Publishing Services, Canberra. been officially evaluated since 2001. Most in: Introduced and translocated fishes and current data on their spread within the state their ecological effects, p. 38-46 comes from them being caught as by catch in other surveys. These are also difficult to sample Chowdhury MBR, Wakabayashi H (1991) A due to their habitat preferences, which also does study of Flexibacter columnaris infection not make them advantageous to consistent in loach, Misgurnus anguillicaudatus monitoring Tabor et al. 2001). It appears that (Bleeker, Gunther). J Fish Dis 14:389-394 Dove ADM, Ernst I (1998) Concurrent invaders- predators, Moroco oxycephalus and four exotic species of Monogenea now Misgurnus anguillicaudatus in the established on exotic freshwater fishes in laboratory and semi-field rice paddy. Kor J Australia. Int J Parasitology 28:1755-1764 Entomol 24:269-284

Franch N, Calvero M, Garrido M, et al (2008) Koster WM, Raadik TA, Clunie P (2002) On the establishment and range expansion Scoping study of the potential spread and of oriental weatherfish (Misgurnus impact of the exotic fish Oriental anguillicaudatus) in NE Iberian Peninsula. weatherloach in the Murray-Darling Basin, Bio Invas 10:1327-1331 Australia: a resource document. Arthur Rylah Institute for Environmental Freyhof J, Korte E (2005) The first record of Research, Melbourne 78 pp. Misgurnus anguillicaudatus in Germany. J Fish Biol 66:568–571 Kottelat M (1998) Fishes of the Nam Theun and Xe Bangfai basins, Laos, with diagnoses of Fuller PL, Nico LG, Williams JD (1999) twenty-two new species (Teleostei: Nonindiginous fishes introduced into inland Cyprinidae, Balitoridae, Cobitidae, Coiidae waters of the United States. Amer J Fish and Odontobutidae). Ichthyol Explor Special Publication 21, Bethesda, MD Freshwat 9:1-128

Kapoor D, Dayal R, Ponniah AG (2002) Fish Kottelat M (2001) Freshwater fishes of northern biodiversity of India. National Bureau of Vietnam. A preliminary check-list of the Fish Genetic Resources Lucknow, India. p. fishes known or expected to occur in 230-231 northern Vietnam with comments on systematics and nomenclature. Keller RP, Lake PS (2007) Potential impacts of Environment and Social Development Unit, a recent and rapidly spreading colonizer of East Asia and Pacific Region. The World Australian freshwaters: oriental Bank. Freshwater Fish. Vietnam 123pp. weatherloach (Misgurnus anguillicaudatus). Eco Fres Fish 16:124- Lei F, Wang B (1990) Studies on the 132 reproduction and growth of loach. Acta Biol Sin 14:60-67 Kim H, Kim M, Yu H (1994) biological control of vector mosquitoes by the use of fish Lintermans M, Rutzou T, Kukolic K (1990) The History, New York, NY. Natural history of status, distribution and possible impacts of Central Asia vol. 9, p. 204-205 the oriental weatherloach Misgurnus anguillicaudatus in the Ginninberra Creek Razzetti E, Nardi PA, Strosselli S, Bernini F catchment. Research report 2. ACT Parks (2001) Prima segnalazione di Misgurnus and Conservation Service Tuggeranong, anguillicaudatus (Cantor, 1842) in acque Australia. interne italiane (First citation of Misgurnus anguillicaudatus in Italian freshwaters). Logan DJ, Bibles EL, Markle DF (1996) Recent Ann Mus Civ Stor Nat Genova 93:559–563 collections of exotic aquarium fishes in the (in Italian) freshwaters of Oregon and thermal tolerance of oriental weatherfish and Rixon CAM, Duggan IC, Bergeron NMN et al pirapatinga. Calif Fish Game 82:66–80 (2005) Invasion risks posed by the aquarium trade and live fish markets on the Maciolek JA (1984) Exotic fishes in Hawaii and Laurentian Great Lakes. Biodiv Conserv other islands of Oceania. The Johns 14:1365-1381 Hopkins University Press, Baltimore, MD. In: W. R. Courtenay, Jr., and J. R. Stauffer, Sal'nikov VB (1998) Anthropogenic migration Jr., editors. Distribution, biology, and of fish in Turkmenistan. J Ichthyol 38:591- management of exotic fishes p.131-161 602

Masuda, HK, Amaoka C, Araga, T et al editors. Schultz EE (1960) Establishment and early (1984) The fishes of the Japanese dispersal of a loach, Misgurnus Archipelago. Tokai University Press, anguillicaudatus (Cantor), in Michigan. Tokai, Japan. 437pp. Trans Amer Fish Soc 89:376-377

McMahon BR, Burggren WW (1987) Simon TP, Bright G, Veraldi F et al (2006) New Respiratory physiology of intestinal air- record for the alien oriental weatherfish breathing in the teleost fish Misgurnus Misgurnus aguillicaudactus, in the Lake anguillicaudatus. J Exp Biol 133:371-393 Michigan Basin, Indiana (Cypriniformes: Cobitidae). Proc Indiana Acad Sci 115:32- Nichols JT (1943) The freshwater fishes of 36 China. American Museum of Natural St. Amant JA, Hoover FG (1969) Addition of Wang Y, Hu M, Cao L et al (2008) Effects of Misgurnus anguillicaudatus (Cantor) to the daphnia (Moina micurura) plus chlorella California fauna. Calif Fish Game 57:330- (Chlorella pyrenoidosa) or microparticle 331 diets on growth and survival of larval loach (Misgurnus anguillicaudatus). Aquacult Int Sterba G (1973) Freshwater fishes of the world. 16:361-368 Tropical Fish Hobbyist Publications, Inc., Neptune City, NJ. English translation and Watanabe K, Hidaka T (1983) Feeding behavior revision from German. Two volumes of the Japanese loach, Misgurnus anguillicaudatus (Cobitididae). J Ethol Suzuki R (1983) Multiple spawning of the 1:86-90 cyprinid loach, Misgurnus anguillicaudatus. Aquaculture 31:233-243 Welcomme RL (1988) International introductions of inland aquatic species. Tabor RA, Warner E, Hager S (2001) An Food and Agriculture Organization of the oriental weatherfish (Misgurnus United Nations (FAO), Rome, Italy. FAO anguillicaudatus) population established in Fisheries Technical Paper 294 318 pp. Washington state. Northwest Sci 75:72–76 Wolf K (1988) Fish viruses and fish viral Taylor JN, Courtenay WR Jr, McCann JA diseases. Comstock Publishing Associates, (1984) Known impacts of exotic fishes in Ithaca, NY the continental United States. The Johns Hopkins University Press, Baltimore, MD Wydoski RS, Whi|ney RR (2003) Inland Fishes p. 322-273. in: Distribution, biology and of Washington. Second Edition. University management of exotic fishes. of Washington Press, Seattle, WA

Tirant G (1929) Oeuvre ichtyologique de G. Yamamoto MN, Tagawa AW (2000) Hawai'i's Tirant. Réimpression par le Service native and exotic freshwater animals. Océanographique des pêches de Mutual Publishing, Honolulu, Hawaii. 200 l'Indochine, Gouvernement Général de p. l'Indochine, Saïgon(First citation of presence in Cambodia). 6° Note, 175 p. (In French)

Other Resources

FishBase Article on Misgurnus anguillicaudatus: http://fishbase.org/Summary/speciesSummary.p hp?ID=3016&genusname=Misgurnus&speciesn ame=anguillicaudatus

USGS Nonindiginous Aquatic Species Database Entry for Misgurnus anguillicaudatus: http://nas.er.usgs.gov/queries/FactSheet.asp?spe ciesID=498

Fact sheet for Misgurnus anguillicaudatus (Cantor, 1824): http://nis.gsmfc.org/nis_factsheet.php?toc_id=19 2

Expert contacts

Roger Tabor , [email protected] 360-753- 9541. Supervisory Fish and Wildlife Biologist U.S. Fish and Wildlife Service, First to document established Misgurnus anguillicaudatus population in WA state in Lake Washington (1999). No longer studying weatherfish, however, keeps a personal database of present records in the state.