ANTHROPOGENIC DISPERSAL OF DECAPOD CRUSTACEANS IN AQUATIC ENVIRONMENTS

Gilberto Rodríguez and Héctor Suárez

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: grodrigu@oikos.ivic.ve

Héctor Suárez. Biologist, Universidad de Oriente, Venezuela. Research Associate, IVIC.

Resumen

La dispersión antropogénica de crustáceos decápodos es parte de un fenómeno que actualmente adquiere características de un cambio ambiental global. De las 10000 especies conocidas de decápodos se han registrado 58 especies marinas, 8 dulceacuícolas-estuarinas y 21 cangrejos de ríos (Astacura) que se han desplazado de sus áreas originales de distribución. Las migraciones lessepsianas a través de canales, la introducción accidental por transporte marítimo y la importación para acuicultura son los mecanismos responsables de esta dispersión. En Venezuela se han introducido 4 especies de camarones peneídeos, el camarón de río Macrobrachium rosenbergii, el cangrejo de río Procambarus clarkii y 3 especies de brachiuros. Los crustáceos decápodos están particularmente bien adaptados por su anatomía y fecundidad para colonizar nuevas áreas y establecer poblaciones explosivas.

Summary

The anthropogenic dispersal of decapod crustaceans is part of an event that in recent years has acquired characteristics of a global environmental change. From the 10000 species of decapods so far recorded in the world, 58 marine species, 8 freshwater-estuarine species and 21 river crabs (Astacura) have been recorded outside their original areas of distribution. Lessepsian Ms through channels, accidental maritime transport and introduction for aquaculture have been the mechanisms responsible for these dispersals. Four penaeid species, the freshwater shrimp Macrobrachium rosenbergii, the river crab Procambarus clarkii, and 3 brachyuran crabs have been introduced in Venezuela. Due to their characteristic anatomy and high fecundity the decapod crustaceans are particularly apt for the colonization of new areas and the establishment of explosive populations.

KEY WORDS / Crustacea Decapoda / Dispersal / Environmental Impact / Biological Invasion /

Resumo

A dispersão antropogênica de crustáceos decápodes é parte de um fenômeno que atualmente adquire características de uma mudança ambiental global. Das 10000 espécies conhecidas de decápodes foram registradas 58 espécies marinhas, 8 doceacuícolas-estuarinas e 21 caranguejos de rios (Astacura) que foram deslocados de suas áreas originais de distribuição. As migrações lessepsianas através de canais, a introdução acidental por transporte marítimo e a importação para aqüicultura são os mecanismos responsáveis desta dispersão. Na Venezuela foram introduzidas quatro espécies de camarões, peneídeos, o camarão de rio Macrobrachium rosenbergii, o caranguejo de rio Procambarus clarkii e 3 espécies de brachiuros. Os crustáceos decápodes estão particularmente bem adaptados por sua anatomia e fecundidade para colonizar novas áreas e estabelecer populações explosivas.

Recibido: 16/04/2001. Aceptado: 16/05/2001

Biological invasion has been considered by recent authors as equivalent to an environmental global change (Vitousek et al., 1996). This opinion reflects the concern for the increasing list of organisms that cross from an ocean to another, establishing dense populations in the new environments. These authors mention as examples Bermuda, where 65% of the vascular plants are non-native, or California, where 76 species of freshwater fishes are native and 42% are non-native. Although this phenomenon is more noticeable in plants, due to the large number of cultivated species, and in freshwater fishes, object of an active trading for aquaria, other groups of native invertebrates show the same tendency. This is the case of decapod crustaceans, which due to construction of channels, increase in maritime transport and development of aquaculture, have found new opportunities for expanding their original areas of distribution

Lessepsian Migrations

The Suez Canal is an unparalleled situation where two biogeographical provinces, previously totally separated, enter into contact and interpenetrate. The pharaonic connection between the Red and Mediterranean seas from the 13th to the 8th centuries BC, allowed the migration of very few species due to the low salinity there prevailing (Por, 1971). The present canal, projected and built by Ferdinand-Marie de Lesseps (1805-1894) was inaugurated on November 1869. The channel spans 160 km, from the Bay of Suez in the Red Sea to Port Said in the Mediterranean, using the intermediate Manzala, Timsah and Bitter lakes.

Migration through the Suez Canal was restrained at the beginning by the hypersaline waters of the Bitter Lakes (»68‰ S), but thereafter a large contingent of Red Sea species have dispersed to the Mediterranean. One of the first to cross was the crab Portunus pelagicus and its trajectory was followed by the Suez Canal Company because of the worth of this species as a food staple. The species became abundant in the canal between 1889 and 1993, reached Port Said in 1898 and four years latter was abundant there. In 1930 was common in Palestine and in 1958 had reached Cyprus and was a common fare in Egypt, with fishing grounds off Port Said, Alexandria and Haifa (Elton, 1958). The active multiplication of other immigrant species has been reflected in the fisheries statistics for the region. Penaeus japonicus, P. semisulcatus, Metapenaeus stebbingi and M. monoceros are now regularly trawled in the Eastern Mediterranean by Turkish, Israeli and Egyptian fishermen (Gorgy, 1966; Holthuis, 1980).

There are at present 40 species of decapods in the Mediterranean accounted for as Lessepsian migrants (Table I) while, on the contrary, the number of species dispersed in the area through ships ballast water is considered negligible. The occasional presence of the lobster Thennus orientalis, first recorded in 1896 in Fiume, Italy (Elton, 1958), can be explained by transport on ships’ hulls.

The unidirectional dispersal from the Red Sea to the Mediterranean (with the exception of a few fish species) has been ascribed to the prevalence of a northward current in the canal. More recently Por (1971) considered the Mediterranean as a zoogeographical cul-de-sac, a tropical sea non-saturated by the temperate Atlantic fauna because of its high salinity and temperature, and consequently "pre-adapted" to receive immigrant species. On the other hand Ben-Tuvia (1966) considered that the Red Sea contains a larger number of species than the Mediterranean due to the adaptive diversity in the tropical and subtropical biotopes. It can be expected that the more vigorous Indo-Pacific species can successfully compete with the native Mediterranean species, while it is less probable than the smaller Atlanto-Mediterranean populations could adapt to the Red Sea conditions. Aron and Smith (1971) considered that the Eastern Mediterranean is still in an unstable equilibrium and that the competitive pressures will lead to a more efficient use of the energy available, which not necessarily will accord to human interests.

The Kiel Canal, built between 1887 and 1895 in an extent of 98 Km, links the North Sea with the Baltic Sea, from the mouth of the Elbe River to the Kiel Bay, bypassing the detour along the Danish peninsula. It is possible that the estuarine mud crab Rhithropanopeus harrisii found its way to the Baltic through the Kiel Canal, after its introduction in the Netherlands, since its first record in that sea, in 1936, was from the Baltic end of the canal (Wolff, 1954). In other respects, this Canal is of little biogeographical relevance.

The Panama Canal spans 64 Km from coast to coast. Although its lake-lock design and the presence of a wide freshwater zone supplied by the Chagres River precludes any Lessepsian migration, at both ends of the present canal several fouling and perforating organisms (pholadid bivalves, teredos, cirripeds, bryozoans, and other) usually considered as natives of the opposing ocean, can be observed. These are euryhaline forms that can withstand transport through the freshwater section attached to the hulls of local vessels that have been moored for a long time (Carlton, 1985).

It has been extensively debated whether it is possible that the ballast water discharged at opposite sides of the Canal, since its aperture in 1914, could be an "active" mechanism for the transport. Carlton (1985) recorded 9 invertebrates and 4 fishes for which this mechanism is possible and 5 invertebrates and 2 fishes for which it is probable. Among the decapod species, transport in ballast water is considered a possible mechanism for Rhithropanopeus harrisii and the freshwater crab Neorhynchus alcocki found in Pedro Miguel lock. Eurypanopeus dissimilis, found in the third lock, ranges from Florida to Brazil and its presence in the Panama Canal is not unusual.

Accidental Introduction

Ship hulls

Several species of crabs, such as Pachygrapsus transversus, Plagusia chabrus, P. depressa, P. tuberculata, Planes minutus, Carcinides maenas, Menippe convexa, etc., have been observed attached to ship’s hulls (Wolff, 1954). This was a convenient means of dispersal for crabs when the old wooden hulls were in use, but it became unavailable with the modern metallic hulls, antifouling paints and reduced stowage time. No decapods were detected during a two-year survey of 89 vessels of different types that moored at New Zealand ports, although 45 species of sessile invertebrates were found (Skerman, 1960).

Ballast water and cargo

Ballast water offers the most effective mechanism for the introduction of exotic decapod species, although it has been confirmed only in a few instances (Carlton, 1985). Larval stages of decapod crustaceans have been recovered from ballast tanks in viable conditions (Chu et al., 1997). Rees and Catley (1949) proposed this mechanism for the introduction of the larvae of Processa equimana, living in the North Sea plankton, into the Mediterranean and Red Sea, but Carlton (1985) considered this unlikely. Furthermore, the North Sea form has been described as a different species, P. modica, with the subspecies P. modica carolii in the Mediterranean.

Peters and Panning (1933) reviewed the available evidence for the first implant of the Chinese crab Eriocheir sinensis in the North Sea, from 1912, via ballast water. Subsequently large populations of this crab have colonized the European brackish waters and have extended to California and Canada, apparently using the same transport mechanism (Cohen and Carlton, 1997).

The original range of Rhithropanopeus harrisii was in the Atlantic coast of America, from the St. Lawrence Gulf in Canada to Veracruz in Mexico. It was introduced into the Netherlands before 1874 and described as a new species by Maitland (1874). It invaded the Baltic through the Kiel Canal, but the low temperatures of this sea in winter halted its advance (Green, 1961). Latter it has been reported from brackish waters throughout Europe, California and Tropical America (Table II). Although Wolff (1954) attributed its first dispersal to ballast water, this is not possible because this sort of structure were not in use at the time, but could account for its further invasion of California and the Black Sea (Carlton, 1985). It was introduced in California in shipments of commercial oysters, and perhaps through the same mechanism along the Atlantic coasts.

The dispersal of the European green crab Carcinus maenas has been traced since the beginning of the 19th century when it was introduced into the Atlantic coast of the United States (1817) and Australia (1901). They possibly traveled as adults attached to the wooden hulls of ships (Cohen et al., 1995), since ballast tanks came into use only after 1870 (Carlton, 1985). In recent times it has continued its advance to South Africa (1983) and California (1989) in ballast water. They could have been introduced also as adults by research laboratories and schools that import these crabs for experiments and demonstrations, and afterward release them into the environment (Carlton, 1985), or in the algae used as packing for marine products. Japanese zoologists have stated that the invading species in Japan is Carcinus aestuarii (=C. mediterraneus), but Geller et al. (1977) determined the differences between the two sibling species of Carcinus and established that in the Japanese and South African populations concur the Atlantic (C. maenas) and Mediterranean (C. aestuarii) haplotypes, while in Eastern United States, California and Australia only the Atlantic haplotype (C. maenas) is present. The mixture of both haplotypes is explained by multiple invasions of the Japanese and South African localities. The introduction of Hemigrapsus sanguineus into the Atlantic coast of the United States, according to McDermott (1998), was through the discharge of ballast water into one or several major estuaries between New England and North Carolina.

Ship cargo also seems to play a role in the transport of organisms. The saber crab Platychirograpsus spectabilis was transported in a cargo of cedar logs from Tabasco State, Mexico, to Hillsborough (St. Petersburg), Florida, where it has become permanently established (Marchand, 1946).

Oil platforms

These and similar structures can be slowly towed across the ocean, with the submersed parts unprotected by antifouling paints, and stay anchored for extended periods of time in different parts of the world. In this way was transported the Japanese shore crab Plagusia dentipes, observed alive in California after a trans-Pacific cruise of 61 days on a self-powered drilling platform, which had been working for four years between Japan and Malaysia (Benech, 1978). Similar circumstances are mentioned for the transport of Plagusia tuberculata, together with an encrusting community of cirripeds, on an oil platform built in Japan and transported to New Zealand, after a voyage of 68 days (Foster and Williams, 1979).

Fishery products

According to Zenkevitch (1963) two Mediterranean shrimps, Palaemon adspersus and P. elegans, were accidentally introduced into the Aral Sea with species of mullets. We have already mentioned the possible introduction of Rhithropanopeus harrisii along the Atlantic coast of the United States, with oysters, and of Carcinus maenas, into San Francisco Bay, with algae used for packing of living bait (Cohen and Carlton, 1997). The dispersal of several species of crayfish throughout the United States, Orconectes rusticus for instance (Table III), is attributed to their escape from containers of living bait used by sport fishermen (Taylor and Redmer, 1996).

Food and aquaria

Many decapod crustaceans are important food staples. They are frequently processed alive, with the implied risk of escape to natural environments. The original area of Callinectes sapidus is in the Atlantic coast of America, where it is the object of an important fishery, but since the beginning of the 20th century it has been successively reported from numerous localities; to those recorded in Table I it should be added the Gulf of Genoa, the north Adriatic, the occidental Black Sea, the eastern Mediterranean, Burma, and other localities. Although living specimens have been recovered alive in San Francisco Bay (Cohen and Carlton, 1997), Denmark, the Netherlands, and France (Wolff, 1954), the species is not established permanently in these localities. On the other hand, it is fished intensively in the Bitter Lakes and adjacent sea in Egypt, and in Greece.

Cohen and Carlton (1997) recorded 16 cases of interception of living specimens of Eriocheir sinensis at San Francisco airport between 1989 and 1995, with the confiscation of 10 to 50 specimens on each occasion. This species is sold alive in Asian food markets in the United States (Lemaitre, 1995).

The trade of organisms for aquaria is another means of dispersal. Living specimens of Atya scabra and Procambarus clarkii can be observed frequently at the pet shops in Caracas.

Aquaculture

Penaeid shrimps

The species of Peneaidae most frequently cultured are Marsupenaeus japonicus and Penaeus monodon, but the possibility exists of an increase in the culture of nine other species, P. semisulcatus, Farfantepenaeus aztecus, Fenneropenaeus indicus, F. penicillatus, Sergestes orientalis, Litopenaeus schmitti, L. setiferus, L. stylirostris and L. vannamei (Liao and Huang, 1982). Farfantepenaeus aztecus is already under experimental culture in AQUACOP station, in Tahiti, since 1975, and L. stylirostris in Corpus Christy, Texas, and Crystal River, Florida, from approximately the same date. M. japonicus was the first species cultured in the world after the research by Hudinaga in 1934. By 1942 its full culture had been achieved, but only after World War II it was established on a commercial basis. In the United States shrimp culture began by the native species L. setiferus and F. aztecus in 1963, and F. duorarum in 1968. This same year Liao began the culture of P. monodon in Taiwan.

Although the penaeids have being cultured in shrimp farms for an extended period of time, there is little information in the literature on the escape of these species and its effects on the natural ecosystems. The information available on the dispersal of M. japonicus in the Mediterranean Sea indicates that, at least under conditions of biological poverty, the species is able to propagate rapidly into the recipient communities.

Several diseases produced by microorganisms and invertebrate ectoparasites have been detected in penaeid shrimps. Those produced by viruses offer the greater environmental risks since, not being specific, they could extend to the shrimp stocks exploited commercially. Numerous viruses have been identified in penaeids in the last 25 years, including 3 baculoviruses, 1 picornavirus, 1 parvovirus and 1 rheovirus (Dall et al., 1990). The first baculovirosis in shrimp, produced by Baculovirus penaei, was detected in 1974 in natural populations of F. duorarum from the Gulf of Mexico; the second was described in P. monodon from shrimp cultured in the laboratory in the United States and transmissible to L. stylirostris and F. californiensis; the third was described in M. japonicus cultured in Japan. The virus responsible for the infectious hypodermic and hematopoietic necrosis (IHHN) was detected in Hawaii, in L. stylirostris from South American cultures, and it is transmissible to P. monodon and L. vannamei. The parvovirosis was found in several species of Indo Pacific Penaeidae, and the rheovirus was detected in M. japonicus from the Mediterranean and P. monodon cultured in Malaysia. In 1992 a new virosis was detected in shrimp farms from Ecuador which produced the Taura syndrome (TSU), with a high mortality rate; it has extended along the Pacific coast from Perú to California, and the Atlantic coast from Texas to Mexico, seriously affecting the production (Lucien-Brun, 1997).

A vibriosis producing bacterial hemorragic septicemia is frequently found in penaeid cultures in Asia and South America. The ethiological agent, Vibrio harveyi, has been detected in shrimp farms in Venezuela, infesting L. vanamei vanamei and L. stylyrostris, but it is also present in feral populations of L. schmitti and several species of marine and estuarine fishes (Alvarez et al., 1998).

Palaemonid shrimps

Macrobrachium rosenbergii, a freshwater-estuarine shrimp occasionally found at sea (Holthuis, 1980) is the palaemonid species most widely cultured in the world (Table II). It has been introduced through all the American continent, except in Nicaragua, Belize, Chile, Paraguay and Bolivia. In the Caribbean it is farmed in the Dominican Republic, Puerto Rico, Dominica, Guadalupe, Martinica, Saint Lucia and Trinidad-Tobago (FAO, 1995).

Crayfishes

The crayfishes of the families Astacidae and Parastacidae are strictly freshwater organisms. According to Hobbs et al. (1989) it would be impossible to mention all the introductions of astacids since in many cases, or in the majority, they have not been recorded (Table III). The largest diversity of species occurs in North America, where Hobbs (1989) reported 271 species and subspecies native to Canada, USA and Mexico. Several species have been transplanted within North America itself. This is the case of Orconectes rusticus, which has displaced two other native species of Orconectes in Illinois (Taylor and Redmer, 1996). These displacements can affect the ecosystem through the reduction in the density of macrophytes (Lodge and Lorman, 1987) and benthonic macroinvertebrates (Houghton et al., 1998).

Another species that has extended its intracontinental distribution is Cherax destructor, from Australia, which has furthermore reached intercontinentally to Washington State. The most notable case of intercontinental expansion is presented by Procambarus clarkii, which has spread out to all continents, except Australia and Antarctica. Laurent and Forest (1979) summarized the damages produced by this species as follows: "In Japan it was introduced since 1930 (from 18 specimens). A short time afterwards it pullulated, destroying rice fields and ditches, producing populations of 2000 k/hectare and not been used by the people [as food]… In Kenya it produced rapidly enormous populations not exploited by the natives… breaking the fishing gears and interfering with the breeding of tilapia".

Species Introduced in Venezuela

Penaeid shrimps

Authorization for the introduction of Marsupenaeus japonicus, P. monodon, Litopenaeus stylirostris and L. vannamei was granted by the Ministry of Agriculture of Venezuela on November 1984. The original areas of distribution of the two former species are in the Indian Ocean and the Western Pacific, and of the other two on the Pacific coast of America.

There is a high risk that the introduced species would escape into the nearby coastal zones, where there are already considerable stocks of native penaeids under exploitation by the local fishing industry. However, each exotic species present different levels of risk according to their peculiar environmental requirements. The life cycle of Marsupenaeus japonicus takes place entirely in the open sea. Penaeus monodon, Litopenaeus stylirostris and L. vannamei spawn in the sea, but larval stages penetrate the estuaries with different timings and to different distances, according to the species (Garcia and Le Reste, 1981). Consequently, M. japonicus would compete only within the sublitoral communities in the Gulf of Venezuela and the continental platform in the Miranda and Anzoategui States. The other three species will compete as well with the estuarine communities, particularly in the Maracaibo System and several coastal lagoons where the breeding grounds of the native L. schmitti are located.

Macrobrachium rosenbergii

This species was introduced in 1980, into a small shrimp farm in Margarita Island, by La Salle Foundation. Although its culture in the country has been unsuccessful, Pereira et al. (1996) found a wild population in the Orinoco Delta in 1996. They considered that this species passed into the natural environment between 1991 and 1993.

Procambarus clarkii

One specimen of this crayfish (total length 8.2 cm, IVIC Reference Collection) was captured in a pond of the Officers Club (Circulo Militar) in Caracas. As mentioned before, the species is available at pet shops in the city.

Brachyuran crabs

Charibdis helleri, a species from the IndoPacific and Red Sea regions that has migrated to the Mediterranean and several localities in the Caribbean, has been recorded by Hernández and Bolaños (1995) from eastern Venezuela. (2) Callinectes arcuatus, recorded by Hernández and Bolaños (1995) from Eastern Venezuela, had not been previously observed outside its original area of distribution, between California and Peru, and a misidentification by these authors cannot be ruled out. (3) Rhithropanopeaus harrisii was observed by the first time in 1957 in several localities of the Tablazo and Strait of Maracaibo (Rodríguez, 1963; Rodríguez and Morales, 2000). As the species was already very abundant in this estuary in 1957, it should have been introduced several years before, possibly in the ballast water of oil tankers involved in the intense traffic between the local oil terminals and the eastern coast of the United States.

General Remarks

From approximately 10000 species of decapod crustaceans described in the world, 88 are recorded in the present contribution as migrants -58 from marine environments, 8 from freshwater-estuarine habitats and 21 are freshwater crayfishes- but still there may be others not yet recorded in the literature. These species have migrated through navigation channels, have been accidentally transported by ships, inadvertently imported with fishery products or introduced for aquaculture. It might be expected that the list would increase in the future.

Decapod crustaceans are particularly well adapted for long distance Migration and occupation of new localities. Their exoskeleton partially protects them from desiccation or osmotic differences, since water and electrolyte transport only occurs through the gills and arthrodial membranes (Green, 1961). Their fecundity is remarkable, being able to produce numerous larvae (hundreds of thousands in some cases), which under native conditions are regulated by predation or loss in the environment, but in new localities could have a high rate of survival.

The effect of invading species on recipient communities could be direct, by displacement of native species or predation on other members of the community. Indirectly they could affect the native species through introduction of diseases. Up to the present severe damages have been rarely observed, for instance the destruction of rice fields in Japan by Procambarus clarkii (Laurent and Forest, 1979). But as the invasion into coastal zones, rivers and lakes progresses, we can expect the gradual displacement of native forms, with results impossible to foresee at present.

ACNOWLEDGEMENTS

The authors are grateful to Jesus Eloy Conde and Jon Paul Rodríguez for critically reading the manuscript.

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