Interciencia ISSN: 0378-1844 [email protected] Asociación Interciencia Venezuela

Rodríguez, Gilberto; Suárez, Héctor Anthropogenic dispersal of decapod in aquatic environments Interciencia, vol. 26, núm. 7, julio, 2001, pp. 282-288 Asociación Interciencia Caracas, Venezuela

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How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative ANTHROPOGENIC DISPERSAL OF DECAPOD CRUSTACEANS IN AQUATIC ENVIRONMENTS

GILBERTO RODRÍGUEZ and HÉCTOR SUÁREZ

iological invasion has The present canal, projected and built by ber of species dispersed in the area been considered by re- Ferdinand-Marie de Lesseps (1805-1894) through ships ballast water is considered cent authors as equiva- was inaugurated on November 1869. The negligible. The occasional presence of the lent to an environmental global change channel spans 160 km, from the Bay of lobster Thennus orientalis, first recorded (Vitousek et al., 1996). This opinion re- Suez in the to Port Said in the in 1896 in Fiume, Italy (Elton, 1958), can flects the concern for the increasing list Mediterranean, using the intermediate be explained by transport on ships’ hulls. of organisms that cross from an ocean to Manzala, Timsah and Bitter lakes. The unidirectional dis- another, establishing dense populations in Migration through the persal from the Red Sea to the Mediter- the new environments. These authors was restrained at the begin- ranean (with the exception of a few fish mention as examples Bermuda, where ning by the hypersaline waters of the species) has been ascribed to the preva- 65% of the vascular plants are non-na- Bitter Lakes (»68‰ S), but thereafter a lence of a northward current in the canal. tive, or California, where 76 species of large contingent of Red Sea species have More recently Por (1971) considered the freshwater fishes are native and 42% are dispersed to the Mediterranean. One of Mediterranean as a zoogeographical cul- non-native. Although this phenomenon is the first to cross was the Portunus de-sac, a tropical sea non-saturated by more noticeable in plants, due to the pelagicus and its trajectory was followed the temperate Atlantic fauna because of large number of cultivated species, and in by the Suez Canal Company because of its high salinity and temperature, and freshwater fishes, object of an active the worth of this species as a food staple. consequently “pre-adapted” to receive im- trading for aquaria, other groups of na- The species became abundant in the ca- migrant species. On the other hand Ben- tive invertebrates show the same ten- nal between 1889 and 1993, reached Port Tuvia (1966) considered that the Red Sea dency. This is the case of decapod crus- Said in 1898 and four years latter was contains a larger number of species than taceans, which due to construction of abundant there. In 1930 was common in the Mediterranean due to the adaptive di- channels, increase in maritime transport Palestine and in 1958 had reached versity in the tropical and subtropical and development of aquaculture, have Cyprus and was a common fare in , biotopes. It can be expected that the found new opportunities for expanding with fishing grounds off Port Said, Alex- more vigorous Indo-Pacific species can their original areas of distribution andria and Haifa (Elton, 1958). The ac- successfully compete with the native tive multiplication of other immigrant Mediterranean species, while it is less Lessepsian Migrations species has been reflected in the fisheries probable than the smaller Atlanto-Medi- statistics for the region. Penaeus japoni- terranean populations could adapt to the The Suez Canal is an cus, P. semisulcatus, Metapenaeus steb- Red Sea conditions. Aron and Smith unparalleled situation where two biogeo- bingi and M. monoceros are now regu- (1971) considered that the Eastern Medi- graphical provinces, previously totally larly trawled in the Eastern Mediterra- terranean is still in an unstable equilib- separated, enter into contact and interpen- nean by Turkish, Israeli and Egyptian rium and that the competitive pressures etrate. The pharaonic connection between fishermen (Gorgy, 1966; Holthuis, 1980). will lead to a more efficient use of the the Red and Mediterranean seas from the There are at present 40 energy available, which not necessarily 13th to the 8th centuries BC, allowed the species of decapods in the Mediterranean will accord to human interests. migration of very few species due to the accounted for as Lessepsian migrants The Kiel Canal, built low salinity there prevailing (Por, 1971). (Table I) while, on the contrary, the num- between 1887 and 1895 in an extent of

KEY WORDS / Crustacea / Dispersal / Environmental Impact / Biological Invasion / Recibido: 16/04/2001. Aceptado: 16/05/2001

Gilberto Rodríguez. Marine Biologist. M.Sc., University of Miami, USA. Ph.D., University of Wales, UK. Emeritus Researcher, Instituto Venezolano de Investigaciones Científicas (IVIC). Address: Centro de Ecología, IVIC, Apartado 21827, Caracas 1020-A, Venezuela. e-mail: [email protected] Héctor Suárez. Biologist, Universidad de Oriente, Venezuela. Research Associate, IVIC.

282 0378-1844/01/07/282-07 $ 3.00/0 JUL 2001, VOL. 26 Nº 7 98 Km, links the North Sea with the Bal- TABLE I tic Sea, from the mouth of the Elbe MARINE DECAPOD CRUSTACEANS DISPERSED FROM THEIR NATURAL River to the Kiel Bay, bypassing the de- DISTRIBUTION AREAS tour along the Danish peninsula. It is possible that the estuarine mud crab Rhithropanopeus harrisii found its way Family Species From To Date Mechanism to the Baltic through the Kiel Canal, af- Penaidae Fenneropenaeus indica Red Sea Mediterranean 19811 M ter its introduction in the Netherlands, Marsupenaeus japonicus Red Sea Mediterranean 19272 M since its first record in that sea, in 1936, Japan Brazil, Tahiti, others. 19753,4,5 Cult Penaeus monodon Indo-Pacific Hawaii, Tahiti, USA, was from the Baltic end of the canal Brazil, England 19753 Cult (Wolff, 1954). In other respects, this Ca- Penaeus semisulcatus Red Sea Mediterranean 19286 M nal is of little biogeographical relevance. Metapenaeus monoceros Red Sea Mediterranean 19272 M The Panama Canal spans Metapenaeus stebbingi Red Sea Mediterranean 19272 M 64 Km from coast to coast. Although its Trachysalambria curvirostris Red Sea Mediterranean 19297 M lake-lock design and the presence of a Metapenaeopsis aegyptia Red Sea Mediterranean 19908 M wide freshwater zone supplied by the Metapenaeopsis Chagres River precludes any Lessepsian mogiensis mogiensis Indo-Pacific, Mediterranean 19979 M migration, at both ends of the present ca- Red Sea 2 nal several fouling and perforating organ- Sergestidae Lucifer hanseni Red Sea Mediterranean 1927 M Leptochela isms (pholadid bivalves, teredos, cirripeds, aculeocaudata Red Sea Mediterranean 193610 M bryozoans, and other) usually considered Leptochela pugnax Red Sea Mediterranean 195811 M as natives of the opposing ocean, can be Processidae Processa aequimana Red Sea Mediterranean 194612 M observed. These are euryhaline forms that North Sea BW 11 can withstand transport through the fresh- Palemonidae Palaemonella rotumana Red Sea Mediterranean 1958 M Periclimenes calami Red Sea Mediterranean 19272 M water section attached to the hulls of local Palemon adspersus Eastern Atlantic Aral Sea 195413 IT vessels that have been moored for a long Caspian Sea 195613 time (Carlton, 1985). 198014 It has been extensively Palemon elegans Eastern Atlantic Aral Sea 195413 AI Caspian Sea 195613 debated whether it is possible that the 198014 ballast water discharged at opposite sides Alphaeidae Automate branchialis Red Sea Mediterranean 195811 M of the Canal, since its aperture in 1914, edwardsi Red Sea Mediterranean 192415 M could be an “active” mechanism for the Alpheus inopinatus Red Sea Mediterranean 195811 M transport. Carlton (1985) recorded 9 in- Alpheus lobidens Red Sea Mediterranean 193610 M Alpheus migrans Red Sea Mediterranean 197816 M vertebrates and 4 fishes for which this Alpheus rapacida Red Sea Mediterranean 196417 M mechanism is possible and 5 inverte- Synalpheus hululensis Red Sea Mediterranean 196417 M brates and 2 fishes for which it is prob- Pandalidae Pandalus kessleri North Pacific Black Sea 195914 IT able. Among the decapod species, trans- Ogyrididae Ogyrides mjobergi Red Sea Mediterranean 195811 M 18,19 port in ballast water is considered a pos- Scyllaridae Thenus orientalis Red Sea Mediterranean 1896 AI Palinuridae Panulirus ornatus Red Sea Mediterranean 198920 M sible mechanism for Rhithropanopeus Lithodidae Paralithodes North Pacific Barents Sea 199621 IT harrisii and the freshwater crab Neorhyn- camtschaticus chus alcocki found in Pedro Miguel lock. Raninidae Notopus dorsipes Red Sea Mediterranean 196417 M Eurypanopeus dissimilis, found in the Calappidae Matuta banksi Red Sea Mediterranean 19908 M 11 third lock, ranges from Florida to Brazil Leucosidae monodi Red Sea Mediterranean 1958 M Leucosia signata Red Sea Mediterranean 19908 M and its presence in the Panama Canal is Myrax fugax Red Sea Mediterranean 193022 M not unusual. Majidae Pyromaia tuberculata Eastern Pacific Japan 197023 AI New Zealand 197824 Accidental Introduction Platimaia wywillethompsoni Red Sea Mediterranean 196725,26 M Hyastenus hilgendorfi Red Sea Mediterranean 196417 M Ship hulls Libinia dubia Atlantic USA, Mediterranean 2000 27 AI Cuba Several species of , Hymenoso- such as Pachygrapsus transversus, Plagu- matidae Elamenopsis kempi Indo-Pacific Panama Canal 196928 AI Cancridae Cancer magister NE Pacific Japan 197629 AI sia chabrus, P. depressa, P. tuberculata, Portunidae Carcinus maenas Europe East Coast, USA 181730 AI Planes minutus, Carcinides maenas, Cape Cod, USA 187731 Menippe convexa, etc., have been ob- New Jersey, USA 190032 served attached to ship’s hulls (Wolff, San Francisco, USA 198533 1954). This was a convenient means of 198929 California, USA 198334 dispersal for crabs when the old wooden South Africa 190035 hulls were in use, but it became unavail- Port Phillip, Victoria able with the modern metallic hulls, anti- (Australia) 197836 fouling paints and reduced stowage time. Australia S Australia, Tasmania 199037 38 No decapods were detected during a two- Carcinus aestuarii Mediterranean Japan 1989 AI Charybdis helleri Indo-Pacific, Mediterranean 19297 M year survey of 89 vessels of different Red Sea Cuba 198739 types that moored at New Zealand ports, Colombia 198840 although 45 species of sessile inverte- Venezuela 199541 brates were found (Skerman, 1960). Florida 199542

JUL 2001, VOL. 26 Nº 7 283 TABLE I (continued) vasion of California and the Black Sea MARINE DECAPOD CRUSTACEANS DISPERSED FROM THEIR NATURAL (Carlton, 1985). It was introduced in Cali- DISTRIBUTION AREAS fornia in shipments of commercial oysters, and perhaps through the same mechanism Family Species From To Date Mechanism along the Atlantic coasts. The dispersal of the Eu- Charybdis longicollis Red Sea Mediterranean 196143 M ropean green crab Carcinus maenas has Thalamita poissonii Red Sea Mediterranean 195811 M been traced since the beginning of the 19th 44 Callinectes sapidus American France 1901 AI century when it was introduced into the Atlantic Holand 195145 Denmark 195146 Atlantic coast of the United States (1817) Japan 197545 and Australia (1901). They possibly trav- Hawaii 198545 eled as adults attached to the wooden Scylla serrata Indo-Pacific New Zealand 196447 Curr hulls of ships (Cohen et al., 1995), since Portunus pelagicus Red Sea Mediterranean 192448 M Xanthidae Atergatis roseus Red Sea Mediterranean 196417 M ballast tanks came into use only after Neopane sayi American Atlantic Wales, UK 195949 AI 1870 (Carlton, 1985). In recent times it Eurypanopeus dissimilis American Atlantic Panama Canal 197150 AI has continued its advance to South Africa Pilumnopeus vauquelini Red Sea Mediterranean 192751 M (1983) and California (1989) in ballast Sphaerozius nitidus Red Sea Mediterranean 197252 M water. They could have been introduced Pilumnus hirsutus Red Sea Mediterranean 193610 M Goneplacidae crenata Red Sea Mediterranean 192751 M also as adults by research laboratories and Grapsidae Plagusia tuberculata Red Sea Mediterranean 196725,26 M schools that import these crabs for experi- Hemigrapsus sanguineus NW Pacific Massachusetts- 198853,54 AI ments and demonstrations, and afterward North Carolina release them into the environment (Carl- 55 sexdentatus Mediterranean Wales, UK 1957 AI ton, 1985), or in the algae used as pack- Abbreviations: AI, Accidental introduction; BW, Ballast Water; Curr, Currents; Cult, Culture; IT, Intro- ing for marine products. Japanese zoolo- duction; M, Migration. gists have stated that the invading species Sources: 1Abdel-Razek et al., 1981; 2Balss, 1927; 3Liao et al., 1982; 4Hanson, 1977; 5Lucien-Brun, in Japan is Carcinus aestuarii (=C. medi- 1997; 6Gruvel, 1928; 7Steinitz, 1929 (cited by Por, 1971); 8Galil and Golani, 1990; 9Galil, 1997; terraneus), but Geller et al. (1977) deter- 10 11 12 13 14 Balss, 1936; Holthuis and Gottlieb, 1958; Rees and Catley, 1949; Zenkevitch, 1963; Holthuis, mined the differences between the two 1980; 15Fox, 1926; 16Lewinsohn and Holthuis, 1978; 17Lewinsohn and Holthuis, 1964; 18Elton, 1958; 19Por, 1971; 20Galil et al., 1989; 21Kuzman et al., 1996 (cited by Jamieson et al., 1998); 22Monod, sibling species of Carcinus and estab- 1930; 23Sakai, 1976 (cited by Cohen and Carlton, 1997); 24Weber and Wear, 1981(cited by Cohen and lished that in the Japanese and South Afri- Carlton, 1997); 25Steinitz, 1967; 26Galil, 1992; 27Einzenross and Einzenross, 2000; 28Abele, 1972; can populations concur the Atlantic (C. 29Abe, 1981 (cited by Cohen and Carlton 1997; 30Say, 1817 (cited by Cohen et al., 1995); 31Smith, maenas) and Mediterranean (C. aestuarii) 32 1879 (cited by Lemaitre, 1995); Dow and Wallace, 1952 (cited by Grodholz and Ruiz, 1996); haplotypes, while in Eastern United States, 33Cohen et al., 1995; 34LeRoux et al., 1990 (cited by Groholz and Ruiz, 1996); 35Fulton and Grant, 1900 (cited by Cohen et al., 1995); 36Zidler, 1978 (cited by Grosholz and Ruiz, 1996); 37Jamieson et California and Australia only the Atlantic al., 1998; 38Geller et al., 1997; 39Gómez and Martínez Iglesias, 1990 (cited by Lemaitre, 1996); 40Cam- haplotype (C. maenas) is present. The pos and Türkay (cited by Lemaitre, 1996); 41Hernández and Bolaños, 1995; 42Lemaitre, 1996; mixture of both haplotypes is explained 43Holthuis, 1961; 44Bouvier, 1901(cited by Wolff, 1954); 45Eldredge, 1995 (cited by Cohen and Carlton, by multiple invasions of the Japanese and 1997); 46Wolff, 1954; 47Dell, 1964 (cited by Foster and Willan, 1979); 48Fox, 1924 (cited by Galil, 1992); 49Naylor, 1960; 50McCosker and Dawson, 1975; 51Calman, 1927 (cited by Galil, 1992); South African localities. The introduction 52Ramadan and Dowidar, 1972; 53Williams and McDermott, 1990; 54McDermott, 1998; 55Naylor, 1957. of Hemigrapsus sanguineus into the At- lantic coast of the United States, accord- ing to McDermott (1998), was through the discharge of ballast water into one or sev- eral major estuaries between New England Ballast water and cargo 1912, via ballast water. Subsequently and North Carolina. large populations of this crab have colo- Ship cargo also seems to Ballast water offers the nized the European brackish waters and play a role in the transport of organisms. most effective mechanism for the intro- have extended to California and Canada, The saber crab Platychirograpsus spect- duction of exotic decapod species, al- apparently using the same transport abilis was transported in a cargo of cedar though it has been confirmed only in a mechanism (Cohen and Carlton, 1997). logs from Tabasco State, Mexico, to few instances (Carlton, 1985). Larval The original range of Hillsborough (St. Petersburg), Florida, stages of decapod crustaceans have been Rhithropanopeus harrisii was in the At- where it has become permanently estab- recovered from ballast tanks in viable lantic coast of America, from the St. lished (Marchand, 1946). conditions (Chu et al., 1997). Rees and Lawrence Gulf in Canada to Veracruz in Catley (1949) proposed this mechanism Mexico. It was introduced into the Nether- Oil platforms for the introduction of the larvae of lands before 1874 and described as a new Processa equimana, living in the North species by Maitland (1874). It invaded the These and similar struc- Sea plankton, into the Mediterranean and Baltic through the Kiel Canal, but the low tures can be slowly towed across the Red Sea, but Carlton (1985) considered temperatures of this sea in winter halted ocean, with the submersed parts unpro- this unlikely. Furthermore, the North Sea its advance (Green, 1961). Latter it has tected by antifouling paints, and stay an- form has been described as a different been reported from brackish waters chored for extended periods of time in species, P. modica, with the subspecies throughout Europe, California and Tropi- different parts of the world. In this way P. modica carolii in the Mediterranean. cal America (Table II). Although Wolff was transported the Japanese shore crab Peters and Panning (1954) attributed its first dispersal to bal- Plagusia dentipes, observed alive in Cali- (1933) reviewed the available evidence last water, this is not possible because this fornia after a trans-Pacific cruise of 61 for the first implant of the Chinese crab sort of structure were not in use at the days on a self-powered drilling platform, Eriocheir sinensis in the North Sea, from time, but could account for its further in- which had been working for four years

284 JUL 2001, VOL. 26 Nº 7 between Japan and Malaysia (Benech, TABLE II 1978). Similar circumstances are men- FRESHWATER AND ESTUARINE DECAPOD CRUSTACEANS DISPERSED tioned for the transport of Plagusia FROM THEIR NATURAL DISTRIBUTION AREAS tuberculata, together with an encrusting community of cirripeds, on an oil platform Family Species From To Date Mechanism built in Japan and transported to New Zealand, after a voyage of 68 days (Foster Atydae Potimirin potimirin Brazil Florida 19711 AI and Williams, 1979). Palaemo- Macrobrachium lar Indo-Pacific, Hawaii 19802 IT nidae Australia Fishery products M. rosenbergii Indo-Pacific, America 19702 IT Australia Middle East According to Zenkevitch Japan (1963) two Mediterranean shrimps, Polynesia Palaemon adspersus and P. elegans, were New Caledonia accidentally introduced into the Aral Sea Palaemon with species of mullets. We have already macrodactylus Indo-Pacific, San Francisco- 19572,3 AI mentioned the possible introduction of Australia USA Rhithropanopeus harrisii along the Atlan- Hymenoso- Neorhynchoplax tic coast of the United States, with oys- matidae kempi Iraq Panama Canal 19691 AI ters, and of Carcinus maenas, into San (locks) Francisco Bay, with algae used for pack- Grapsidae Eriocheir sinensis China, Korea Germany 19124 S ing of living bait (Cohen and Carlton, France 19305 1997). The dispersal of several species of Denmark 19356 crayfish throughout the United States, Denmark 19367 Orconectes rusticus for instance (Table Czechoslovakia 19388 III), is attributed to their escape from con- Belgium 19439 tainers of living bait used by sport fisher- Finland, Sweden 196310 11 men (Taylor and Redmer, 1996). Lake Erie, Canada 1963 British Islands 198612 13 Food and aquaria Portugal 1988 California, USA 199214 Many decapod crusta- Platychirograpsus spectabilis Mexico Florida 194615 S ceans are important food staples. They Xanthidae Rhithropanopeus are frequently processed alive, with the harrisii East coast, Netherlands 187416 AI implied risk of escape to natural environ- USA Germany 193617 ments. The original area of Callinectes California 193718 sapidus is in the Atlantic coast of Ameri- Black Sea 193919 ca, where it is the object of an important Denmark 195320 fishery, but since the beginning of the Venezuela 195621 20th century it has been successively re- Caspian Sea 195822 ported from numerous localities; to those Panama Canal 196923 recorded in Table I it should be added Portugal 199124 the Gulf of Genoa, the north Adriatic, the Spain 199125 occidental Black Sea, the eastern Medi- terranean, Burma, and other localities. Abbreviations: AI, Accidental introduction; IT, Introduction; S, Ships. Although living specimens have been re- Sources: 1Abele, 1972; 2Holthuis, 1980; 3Newman, 1963; 4Peters and Panning, 1933; 5Otto and Kamps, 1935 (cited by Cabral and Costa, 1999); 6Jensen, 1936 (cited by Cabral and Costa, 1999); 7Panning, covered alive in San Francisco Bay 1939 (cited by Cohen and Carlton, 1997); 8Leloup, 1943 (cited by Cabral and Costa, 1999); 9Hoestland, (Cohen and Carlton, 1997), Denmark, the 1959 (cited by Cabral and Costa, 1999); 10Haatela, 1963 (cited by Cohen and Carlton, 1997); 11Nepszy Netherlands, and France (Wolff, 1954), and Leach, 1973 (cited by Cohen and Carlton, 1997); 12Ingle, 1986 (cited by Cohen and Carlton, 1997); the species is not established permanently 13Cabral and Costa, 1999; 14Cohen and Carlton, 1997; 15Marchand, 1946 (cited by Cohen and Carlton, in these localities. On the other hand, it 1997); 16Maitland, 1874; 17Schubert, 1936 (cited by Wolff, 1954); 18Jones, 1940 (cited by Wolff, 1954); 19Makarov, 1939 (cited by Wolff, 1954); 20Wolff, 1954; 21Rodríguez, 1963; 22Nebolsina, 1959 (cited by is fished intensively in the Bitter Lakes Gonçalves et al., 1995); 23Dawson, 1973 (cited by Carlton, 1985); 24Gonçalves et al., 1995; 25Mariscal et and adjacent sea in Egypt, and in Greece. al., 1991 (cited by Gonçalves et al., 1995). Cohen and Carlton (1997) recorded 16 cases of interception of liv- ing specimens of Eriocheir sinensis at Aquaculture Litopenaeus schmitti, L. setiferus, L. styl- San Francisco airport between 1989 and irostris and L. vannamei (Liao and Huang, 1995, with the confiscation of 10 to 50 Penaeid shrimps 1982). Farfantepenaeus aztecus is already specimens on each occasion. This species under experimental culture in AQUACOP is sold alive in Asian food markets in the The species of Peneaidae station, in Tahiti, since 1975, and L. styl- United States (Lemaitre, 1995). most frequently cultured are Marsupenaeus irostris in Corpus Christy, Texas, and The trade of organisms japonicus and Penaeus monodon, but the Crystal River, Florida, from approximately for aquaria is another means of dispersal. possibility exists of an increase in the cul- the same date. M. japonicus was the first Living specimens of Atya scabra and ture of nine other species, P. semisulcatus, species cultured in the world after the re- Procambarus clarkii can be observed fre- Farfantepenaeus aztecus, Fenneropenaeus search by Hudinaga in 1934. By 1942 its quently at the pet shops in Caracas. indicus, F. penicillatus, Sergestes orientalis, full culture had been achieved, but only

JUL 2001, VOL. 26 Nº 7 285 after World War II it was established on a TABLE III commercial basis. In the United States CRAYFISH SPECIES DISPERSED FROM THEIR NATURAL DISTRIBUTION shrimp culture began by the native species AREAS (BASED IN HOBBS et al., 1989, HOBBS, 1989, FAO, 1995) L. setiferus and F. aztecus in 1963, and F. duorarum in 1968. This same year Liao began the culture of P. monodon in Tai- Family Species From To wan. Although the penaeids Parastacidae Cherax destructor Central, E, S Australia SW Australia, have being cultured in shrimp farms for Samoa, WA an extended period of time, there is little Cherax quadricarinatus Samoa, Zambia information in the literature on the escape Cherax tenuimanus Australia of these species and its effects on the Astacidae Astacus astacus France, Germany, Italy Finland, Great natural ecosystems. The information avail- Britain, Spain able on the dispersal of M. japonicus in Astacus leptodactylus Affluents of Black, Austria, Poland, the indicates that, at Caspian and Aral Seas Spain least under conditions of biological pov- Austropotamobius pallipes France England, Ireland, erty, the species is able to propagate rap- Corcega idly into the recipient communities. Pacifastacus leniusculus British Columbia, CA, Austria, Poland, Several diseases pro- ID, NV, OR, UT, WA Finland, Great duced by microorganisms and invertebrate Britain, Ireland, ectoparasites have been detected in Sweden, Japan penaeid shrimps. Those produced by vi- Cambaridae Cambarellus patzcuarensis Mexico Japan ruses offer the greater environmental risks Cambarellus shufeldtii LA, Ms, TN GA since, not being specific, they could ex- Faxonella clypeata AR and LA. to FL GA tend to the shrimp stocks exploited com- Orconectes immunis Mississippi River Canada, CO, NY mercially. Numerous viruses have been Orconectes juvenilis Mississippi River VA identified in penaeids in the last 25 years, Orconectes limosus USA Austria, France, including 3 baculoviruses, 1 picornavirus, Germany, 1 parvovirus and 1 rheovirus (Dall et al., Poland, 1990). The first baculovirosis in shrimp, Netherlands, ME, NH produced by Baculovirus penaei, was de- Orconectes neglectus White and Arkansas OR tected in 1974 in natural populations of F. rivers duorarum from the Gulf of Mexico; the Orconectes obscurus NY, PA, MD, VA Canada, VT, second was described in P. monodon from ME, MA, NY shrimp cultured in the laboratory in the Orconectes propinquus Ontario, Central USA WI United States and transmissible to L. styl- Orconectes rusticus OH, KY, IN Canada, CT, IL, irostris and F. californiensis; the third was ME, MA, NH, NM described in M. japonicus cultured in Ja- Orconectes virilis Canada, CO Francia, Sweden, pan. The virus responsible for the infec- Mexico, CA, MN, tious hypodermic and hematopoietic ne- MD, MA, NY, VT crosis (IHHN) was detected in Hawaii, in Procambarus clarkii USA All continents L. stylirostris from South American cul- except Australia tures, and it is transmissible to P. mon- and Antarctica odon and L. vannamei. The parvovirosis Procambarus acutus MS GA, ME, NY was found in several species of Indo Pa- Abbreviations: AR, Arkansas; CA, California; CO, Colorado; CT, Connecticut, FL, Florida; GA, Georgia; cific , and the rheovirus was de- ID, Idaho; IL, Illinois; IN, Indiana; KY, Kentucky; LA, Louisiana; MA, Massachusetts; MD, Maryland; tected in M. japonicus from the Mediterra- ME, Maine; MS, Mississippi; NV, Nevada; NH, New Hampshire; NM, New Mexico; NY, New York; OH, nean and P. monodon cultured in Malay- Ohio; OR, Oregon; PA, Pennsylvania; TN, Tennessee; UT, Utah; VA, Virginia; VT, Vermont; WA, Wash- sia. In 1992 a new virosis was detected in ington; WI, Wisconsin. shrimp farms from Ecuador which pro- duced the Taura syndrome (TSU), with a high mortality rate; it has extended along the Pacific coast from Perú to California, Palaemonid shrimps Crayfishes and the Atlantic coast from Texas to Mexico, seriously affecting the production Macrobrachium rosen- The crayfishes of the (Lucien-Brun, 1997). bergii, a freshwater-estuarine shrimp occa- families Astacidae and Parastacidae are A vibriosis producing sionally found at sea (Holthuis, 1980) is strictly freshwater organisms. According to bacterial hemorragic septicemia is fre- the palaemonid species most widely cul- Hobbs et al. (1989) it would be impos- quently found in penaeid cultures in Asia tured in the world (Table II). It has been sible to mention all the introductions of and South America. The ethiological agent, introduced through all the American conti- astacids since in many cases, or in the Vibrio harveyi, has been detected in nent, except in Nicaragua, Belize, Chile, majority, they have not been recorded shrimp farms in Venezuela, infesting L. Paraguay and Bolivia. In the Caribbean it (Table III). The largest diversity of species vanamei vanamei and L. stylyrostris, but it is farmed in the Dominican Republic, occurs in North America, where Hobbs is also present in feral populations of L. Puerto Rico, Dominica, Guadalupe, Mar- (1989) reported 271 species and subspe- schmitti and several species of marine and tinica, Saint Lucia and Trinidad-Tobago cies native to Canada, USA and Mexico. estuarine fishes (Alvarez et al., 1998). (FAO, 1995). Several species have been transplanted

286 JUL 2001, VOL. 26 Nº 7 within North America itself. This is the lagoons where the breeding grounds of the Decapod crustaceans are case of Orconectes rusticus, which has native L. schmitti are located. particularly well adapted for long distance displaced two other native species of Migration and occupation of new locali- Orconectes in Illinois (Taylor and Macrobrachium rosenbergii ties. Their exoskeleton partially protects Redmer, 1996). These displacements can them from desiccation or osmotic differ- affect the ecosystem through the reduction This species was intro- ences, since water and electrolyte transport in the density of macrophytes (Lodge and duced in 1980, into a small shrimp farm only occurs through the gills and arthro- Lorman, 1987) and benthonic macroinver- in Margarita Island, by La Salle Founda- dial membranes (Green, 1961). Their fe- tebrates (Houghton et al., 1998). tion. Although its culture in the country cundity is remarkable, being able to pro- Another species that has has been unsuccessful, Pereira et al. duce numerous larvae (hundreds of thou- extended its intracontinental distribution is (1996) found a wild population in the sands in some cases), which under native Cherax destructor, from Australia, which Orinoco Delta in 1996. They considered conditions are regulated by predation or has furthermore reached intercontinentally that this species passed into the natural loss in the environment, but in new locali- to Washington State. The most notable environment between 1991 and 1993. ties could have a high rate of survival. case of intercontinental expansion is pre- The effect of invading sented by Procambarus clarkii, which has Procambarus clarkii species on recipient communities could be spread out to all continents, except Aus- direct, by displacement of native species or tralia and Antarctica. Laurent and Forest One specimen of this predation on other members of the commu- (1979) summarized the damages produced crayfish (total length 8.2 cm, IVIC Refer- nity. Indirectly they could affect the native by this species as follows: “In Japan it ence Collection) was captured in a pond species through introduction of diseases. was introduced since 1930 (from 18 speci- of the Officers Club (Circulo Militar) in Up to the present severe damages have mens). A short time afterwards it pullu- Caracas. As mentioned before, the species been rarely observed, for instance the de- lated, destroying rice fields and ditches, is available at pet shops in the city. struction of rice fields in Japan by Procam- producing populations of 2000 k/hectare barus clarkii (Laurent and Forest, 1979). and not been used by the people [as Brachyuran crabs But as the invasion into coastal zones, riv- food]… In Kenya it produced rapidly ers and lakes progresses, we can expect the enormous populations not exploited by the Charibdis helleri, a spe- gradual displacement of native forms, with natives… breaking the fishing gears and cies from the IndoPacific and Red Sea re- results impossible to foresee at present. interfering with the breeding of tilapia”. gions that has migrated to the Mediterra- nean and several localities in the Carib- ACNOWLEDGEMENTS Species Introduced in Venezuela bean, has been recorded by Hernández and Bolaños (1995) from eastern Venezuela. (2) The authors are grateful Penaeid shrimps Callinectes arcuatus, recorded by Hernán- to Jesus Eloy Conde and Jon Paul Rodrí- dez and Bolaños (1995) from Eastern Ve- guez for critically reading the manuscript. Authorization for the in- nezuela, had not been previously observed troduction of Marsupenaeus japonicus, P. outside its original area of distribution, be- monodon, Litopenaeus stylirostris and L. tween California and Peru, and a misi- vannamei was granted by the Ministry of dentification by these authors cannot be REFERENCES Agriculture of Venezuela on November ruled out. (3) Rhithropanopeaus harrisii 1984. 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