Alien species have been transported and traded by humans for many centuries. However, with the era of globalization, biological invasions have reached notable magnitudes. Currently, introduction of alien species is one of the major threats to biodiversity and ecosystem functioning. The North American crayfish Procambarus clarkii is one of the most widely introduced freshwater species in the world, especially due to its high economic importance. It is responsible for great modifications in invaded environments causing irreparable ecological and economic damages.

Its impressive ability to successfully colonize a wide range of environments is a consequence of its behavioural and biological characteristics that can adapt to features of the invaded location, conferring to this species a notable ecological plasticity. This review summarizes the available information regarding P. Clarkii's biology and invasive dynamics around the world in order to contribute to the understanding of the threats posed by its establishment, as well as to support management and impact mitigation efforts. Key words: Alien biology; Exotic crayfish; Red swamp crayfish; Invasive features; Impact; Invasion management. Morphological aspects Procambarus clarkii's body, as a typical decapod crustacean, is divided into cephalothorax and abdomen, both parts having appendages following the decapod pattern ( ). The abdominal appendages called pleopods are not always present in decapod males, but in this species, they are present in both males and females ( ). The thoracic appendages (pereiopods) are five in number, as usual, but the first 3 pairs are chelate, which is a characteristic of the infraorder Astacidea (;; ).

Become Our Fan on Social Sites! Thursday, 31 January 2013. Download Winning eleven 2002 (PS1). This paper is divided into eleven sections that cover various aspects of P. Clarkii's systematics, life history, physiology, and ecology, as well as impacts observed in. The thoracic appendages (pereiopods) are five in number, as usual, but the first 3 pairs are chelate, which is a characteristic of the infraorder Astacidea (;;).

The carapace color is dark red, orange or reddish brown, although blue, yellow, white and black varieties are known ( ); chelae are typically red on both surfaces. Juveniles are usually light green with a narrow dark band on either side of the abdomen and a broader lighter band along the dorsal surface. Adult specimens can measure up to 15 centimeters of total length, although most individuals are up to 12 centimeters ( ).

This species presents external sexual dimorphism and sex can be distinguished by the position of the genital pores. The genital openings are located on the coxopodite of the third pair of pereiopods in females and on the fifth pair of pereiopods in males ( ). Moreover, males have a copulatory organ formed by a modification of the first and second pair of pleopods () whereas in females the first pair of abdominal appendages is vestigial and the second has no modification () (; ). Figure 1 Ventral view of Procambarus clarkii individuals. A: Morphotype I male (reproductive form) showing the more calcified copulatory organ and the copulatory hooks on the 3 rd and 4 th pereiopods' ischia. B: Morphotype II male (non-reproductive form) without the copulatory hooks and softener copulatory organ.

C: Female showing the first pair of abdominal appendages which is vestigial and the annulus ventralis. Scale bars = 2 cm Likewise most crayfish species, P.

Clarkii has sexual reproduction. Sexual maturity is reached in approximately three months and, depending on climate, it may produce two or three generations per year ( ). In adult males, two different morphotypes that alternate between each other can be observed: the reproductive form or type I male, with hooks on the ischia of the 3rd and 4th pereiopods and more calcified copulatory organs (), and the non-reproductive form or type II male, in which hooks are lacking () (; ). This morphotype alternation in males is a characteristic of the family Cambaridae ( ). An adult male might remain as type I for up to 9 months in a year.

Procambarus clarkii females, on the other hand, have no morphological alteration during reproductive phase and the only different characteristic is an increase in its receptivity to males, which might happen more than once a year depending on environmental characteristics ( ). Reproduction The knowledge of the reproductive strategy of invasive species is central to the understanding of its invasion ecology as it determines the potential for population increase and range expansion. In Procambarus clarkii, mating period, as well as recruitment and sexual maturation, vary according to hydrographic period and environmental conditions (; ) and therefore, due to the combined effects of these factors, reproduction may change after the species is introduced into different regions. Reproduction is regulated by pheromones perceived by receptors located on the antennae which are responsible for interspecific and intraspecific recognition and behavioural modulation ( ). After sex recognition, male courts female through a specific sequence of movements followed by copulation, when the male turns the female with her dorsal surface against the substrate, holding her chelipeds and both ventral regions remain in contact. Thereafter, male deposits the spermatophore in the annulus ventralis(not in the genital pore) (), which is the female's sexual receptacle, located between the bases of the posterior walking legs (; ).

Days, weeks or months after mating, depending on environmental conditions, the female safeguards herself in a burrow and starts oviposition; this process may occur in open water but this is very uncommon ( ). The number of eggs per brood may reach up to 700 (; ) although it depends on female size and is also related to water temperature, population density and the length of the hydroperiod ( ). The embryonic development time depends on atmospheric temperature and may be inhibited under 10 ºC ( ). After hatching, juveniles are kept under the female's abdomen for three weeks ( ). Trophic ecology Procambarus clarkii is a generalist omnivore species whose opportunistic diet favours its own successful establishment in different types of water bodies ( ).

Despite the fact that its diet in natural habitats is not widely studied, much information has been collected in its invasive range (;;;;; ). They are reported to feed on plant and animal detritus, macrophytes and live animals such as molluscs, insects, annelids, nematodes, platyhelminthes, tadpoles and fingerlings () (;;;;;;;; ). Figure 2 Trophic ecology and biotic relationships of Procambarus clarkii regarding food items, predators, pathogens and parasites. Clarkii's predators, the most widely cited in literature are fishes, birds and mammals like otters and capybaras (; ); juveniles can also be eaten by odonate nymphs, coleopteran larvae and aquatic hemipterans () ( ). Invasive species might bring exotic diseases that can reach much higher severity than in its native distribution area (). By far, the most known and studied pathogen of P.

Clarkii is the the parasitic Oomycete Aphanomyces astaci, the causative agent of crayfish plague, which is lethal to many species of crayfish (; ). This pathogen initially infects the exocuticle and after the endocuticle; in more susceptible species or individuals, it penetrates the basal lamina, underneath the epidermis cell layer and spreads throughout the body, invading connective tissue and blood vessels ( ). Aphanomyces astaci does not produce sexual structures, with transmission occurring via zoospores released from infected animals and it is able to survive for several days in water and several weeks in mud (;; ).

Astaci, another parasitic fungus that can infests P. Clarkii carapace and constitute a threat to other crustaceans is Saprolegnia parasitica ( ). The great threat posed by the introduction of A. Astaci, was the trigger to initiate studies on freshwater crayfish pathology in the mid-1900s and this organism has become one of the most intensively studied of all invertebrate infectious diseases ( ).

Clarkii may host many other parasites, pathogens and symbionts that are not as much studied as A. Astaci, although some of them may affect human health. Among viruses, the White Spot Syndrome Virus (WSSV) is perhaps the most devastating of all crustacean viruses.

WSSV has a very wide host range, including P. Clarkii (; ). Bacterial infections are also very common and can be developed by a variety of Gram positive and Gram negative forms that can inhabit their external exoskeleton, gut and frequently the haemolymph ( ). One of the most studied bacteria that affects P. Clarkii is Vibrio mimicus ( ) and it is recognized as a cause of gastroenteritis in humans when feeding on contaminated raw crayfish ( ).

Regarding Platyhelminthes, digeneans are the most common parasites of P. Within Paragonimidae, species from Paragonimus,highly evolved parasites with a complex life cycle that involves at least three different hosts, i.e., snails, crustaceans, and mammals, are known to use crayfish species, including P. Clarkii, as intermediate hosts ( ); the adults of Paragonimus reside and mate in the lungs of a variety of mammalian hosts, wild and domestic animals, as well as humans, causing a disease named paragonimiasis (; ). Some digeneans also may use P. Clarkii as intermediate host, for example: some species of Microphallus (Microphallidae), whose adults parasitize bird and fish species, being larvae cercariae maybe found in the hepatopancreas of P.clarkii ( ); Maritrema obstipum (Van Cleave and Mueller, 1932) (Microphallidae), whose definitive host are birds and mammals and intermediate host maybe freshwater crustaceans and snails, being larvae cercariae found in the gills and hepatopancreas of P. Clarkii (; ); Sogandaritrema progeneticum (Sogandares-Bernal, 1962) (Microphallidae) that parasitizes some crayfishes, including P. Clarkii (; ); Gorgodera amplicava (Looss, 1899) (Gorgoderidae), whose definitive host are some anuran species, may use freshwater bivalves from de family Sphaeriidae and crustaceans as intermediate host.

The immature form of G. Amplicava can be found in the stomach of P. Another example is Macroderoides typicus (Winfield, 1929) (Macroderoididae) that uses the freshwater fish bowfin, Amia calva Linnaeus, 1766, as definitive hosts and freshwater gastropods and crayfishes as intermediate hosts. Clarkii, cysts have been observed in the cephalothorax and antennae ( ). Acanthocephalans are obligate parasites that require usually two hosts, an invertebrate and a vertebrate. Typically, the invertebrate host is an insect or a crustacean and the vertebrate hosts can include mammals, birds and fish. These helminths are not considered significant pathogens of freshwater crayfish.

However, when there is a high prevalence of infection, crayfishes might become stressed and more susceptible to other diseases ( ). Southwellina dimorpha (Schmidt, 1973) (Polymorphidae) is the most studied acanthocephalan observed in P. Clarkii and its cysts are in the anterior portion of the crayfish abdomen, usually attached along the intestine (; ). Procambarus clarkii is also vulnerable to microsporidians, intracellular parasites belonging to the Phylum Microspora ( ). One example is Thelohania contejeani (Henneg, 1892), that has caused mass mortalities of crayfish in Europe (;; ). Another known disease is psorospermiasis, caused by Psorospermium(Mesomycetozoea) that colonizes gills, hepatopancreas, antennal gland, connective and neural tissues, ovary membranes and cardiac and skeletal muscle (;; ).

Additionally, a range of ectocommensals or ectosymbionts from a number of different Phyla can infest the crayfish exoskeleton, including gills. Most of them are temnocephalans (Platyhelminthes, Temnocephalida) that usually are not pathogenic to crayfish, however, they may impair proper gill functioning if present in such high numbers as to impede water flow in the gill cavity ( ). No temnocephalans were reported on P. Clarkii but considering their wide range of occurrence among crayfishes (; ), it is believed that the Red Swamp Crayfish might also be infested by them. Branchiobdellidans (Annelida, Clitellata) may also be observed on crayfishes, including P. Clarkii, but infestations usually take place only when individuals are already ill; infested healthy animals are rarely found ( ). Temnocephalids are predominantly found in the southern hemisphere and no branchobdellid species occur naturally in the southern hemisphere ( ) although both groups have been found outside their native range, as well as many other crayfish diseases, due to anthropogenic movements of the crayfish host (; ).

Behaviour Different behavioural characteristics may contribute to the invasive potential of species; aggressive behaviour, for exemple, may influence competitive displacement of native species (;; Usio et al., 2000), locomotion and dispersal hability can influence rates of spread and spatial patterns of invasion (; ), burrowing behaviour might help coping with environmental stress ( ) and parental care can improve reproductive fitness by increasing brood survival ( ). Procambarus clarkii is a social animal with formation of social dominance hierarchies in adults and juveniles (; ). Little is known about how the hierarchical relationship forms over time and is subsequently maintained, despite some evidences shown by where dominance seems to be size-based. High levels of aggressiveness, experience of winning and attacking or approaching first also may contribute to the formation and maintenance of a dominant hierarchy (;;; ).

This species is defined as nocturnal ( ) although some individuals were observed performing long displacements both by day and by night (; ). Regarding its locomotion, home-range faithfulness and dispersal abilities, some findings have been controversial, which might indicate that behaviour varies according to environmental characteristics.

According to, P. Clarkii specimens are capable of major usage of space, moving up to 4 km.day -1. Found that locomotory speed is significantly correlated with crayfish size and the extent of locomotion is variable, ranging from 1 to 11 m.day -1; in this study, neither speed nor distance walked were related to gender., on the other side, showed that both sexes disperse although males and females use space differently, females being more nomadic than males. The movement pattern seems to be complex, with one or more short peaks of intense locomotion ('wandering' phases) alternated with periods of scarce mobility, with slow speeds or no movement ('stationary' phases) (;;;; ). Like many crayfish species, P. Clarkii is an efficient digger wich uses a combination of tactile and visual information, together with environmental use of cues (i.e. Humidity cues) to orientate its burrowing behaviour; burrows are used as refuge to avoid predation, dehydration and environmental stress as well as to nest (; ).

However, despite the great importance of these shelters, individuals seem not to return to previously occupied burrows at the end of their foraging excursions despite the time and energy expended to excavate them ( ). This species, like most freshwater crayfishes, shows relatively complex parental care (;; ). Mothers often remain in their burrow for many weeks as they execute few cleaning and feeding acts; hatchlings and young remain attached to mother's abdomen for 3 to 4 months (; ). This behaviour, despite being costly, increases reproductive fitness by augmenting brood survivorship and therefore improving the colonization capability of the species.

Causes of introduction Shipping and aquacultural activities are the main agents of introduction of invasive alien crustaceans around the world ( ). Regarding alien freshwater crayfishes, shipping and ballast water are not important pathways but aquaculture and activities associated with the aquarium and bait industry are (; ). Indeed, the aquaculture of Procambarus clarkii is the most important vector of introduction, this species being one of the most important freshwater decapods farmed for consumption (; ). Furthermore, this species has also been introduced as food for fishes and for other edible species like bullfrogs (; ). In Africa, P.

Clarkii has also been introduced as a biological control agent to reduce snail populations, which are intermediate hosts of schistosomiasis (Bilharzia) ( ). Besides, escapes from garden ponds and the pet trade are also important introduction pathways (;; ). Furthermore, especially in Europe, the red swamp crayfish has also been introduced to replace indigenous species, like Austropotamobius pallipes Lereboullet, 1858 for example, which was nearly extinct (; ). Invasion range and history Despite being native to the central southern United States and northeastern Mexico (), Procambarus clarkiihas been cultured extensively through the USA and was introduced by humans into different parts of the northern region, currently being found in Arizona, California, Georgia, Hawaii, Idaho, Indiana, Maryland, Nevada, New Mexico, New York, North Carolina, Ohio, Oklahoma, Oregon, South Carolina and Utah (;;; ). In northwestern Mexico, it has been successfully introduced in the states of Baja California and Sonora (; ). Impacts Procambarus clarkii occupies an important position in the trophic structure of invaded environments, interacting with different trophic levels and changing the whole ecosystem functioning (;;; ). Its flexible feeding strategy affects both lower and higher trophic levels by grazing on macrophytes and algae and preying on macroinvertebrates, fish fingerlings and tadpoles (;;; ).

Its efficiency on grazing macrophytes and extensive burrowing activity can alter freshwater environments, modifying them from macrophytedominated areas with clear water to phytoplankton dominated turbid areas (;; ). Macrophytes are particularly important to aquatic environments because they function as service providers and ecosystem engineers (;; ) avoiding erosion, facilitating nutrient cycling and providing habitat to associated faunal communities (;; ), and serious changes may occur in aquatic environments if submersed plant species are overgrazed. Additionally, the burrowing behaviour might also cause river or channel bank erosion and increase water turbidity (; ). These changes in water characteristics alter aquatic ecosystems and are believed to induce cyanobacterial blooms ( ). Furthermore, P. Clarkii is one of the vectors of the crayfish plague, which is mostly asymptomatic in North American crayfish species such as P. Clarkii, Orconectes limosus ( ) and Pacifastacus leniusculus (Dana, 1852), but lethal to crayfish from other regions (;; ).

This disease, caused by Aphanomyces astaci, constitutes a remarkable threat to indigenous crayfish species, thus being one of the leading causes of native crayfish population decline in Europe (;; ). Clarkii may carry many other pathogens, parasites, epibionts and diseases (see 'Trophic ecology') that can affect other species, man included. In addition to its influence on biodiversity, P. Clarkii can also have a considerable economic impact. Primarily, costs of ecological damage and control measures can be highlighted ( ). Costs of damage mainly occur in the agricultural, forestry, and fishery sectors.

Regarding agricultural economic impacts specifically, crayfish infestation has caused serious damage to drainage systems as a consequence of its burrowing activities, causing important losses of rice yield (;;;; ). The red swamp crayfish is also a problem for fisheries once it spoils valuable fish caught in gill-nets, damages fish nets and is considered a pest in many fish ponds (de Moor, 2002; ). Management and control Procambarus clarkii invasion management options include the elimination or reduction of populations employing physical, chemical or biological methods and the use of legislation to prohibit the transport and release of specimens.

Removal campaigns using traps, fyke or seine nets and electro-fishing are commonly utilized as physical control, although they are often biased by crayfish size and sex (; ). These methods are effective for population reduction but eradication is unlikely if populations are not restricted in range and size ( ). Nevertheless, when physical control is to be used, it is better to invest in continued trapping then short-term intensive trapping, which can cause a feedback response in the population by stimulating faster maturation of juveniles and larger offspring per brood (; ). Drainage of ponds is also extensively used, especially in water bodies with dense populations, as well as diversion of rivers and construction of barriers; nonetheless, the efficiency of these methods is not yet confirmed, especially for pond drainage, since P.

Clarkii is resistant to drought due to its burrowing capacity (; ). Manugistics User Manual. Another common practice to eradicate or control crayfish populations is with biocide, the most used being the application of xenobiotics, organophosphate, organochlorine, and pyrethroid insecticides ( ). Chemical methods, however, were found to be ineffective because of their selective efficiency, with individual crayfish being differentially affected depending on size. In Italy, a laboratory test using the synthetic pyrethroid ciflutrin was found to be relatively effective ( ). Chemical control of crayfish activity, aiming to induce temporary inactivity, was also tested without success in rice field ecosystems ( ).

Besides being expensive, especially when applied to large areas, chemical control methods may have devastating impacts on native species and affect a wide range of organisms (; ). In fact, there are no selective biocides for crayfish or even crustaceans and development of resistance is frequent. Furthermore, the possibility of bioaccumulation and biomagnification cannot be discounted.

Biological control methods were also employed worldwide, including the use of fish predators, disease-causing organisms and microbes that produce toxins (; ) but the only method that has been successful so far is the use of predaceous fish like eels, burbots, perches and pikes (;; ). Nevertheless, biocontrol might be risky since it may lead to new species introductions and it is not specific to the target organism, possibly also affecting native organisms as well. All methods mentioned above present environmental costs that can overcome their benefits. In fact, no single method for eradication is apparently successful. Thus, a combination of methods should be considered, such as trapping and the introduction of predatory fish species ( ).

Final remarks Species moved beyond the limits of their normal geographic ranges by human actions usually have strong ecological impacts (;;; ) and the effects of biological invasion in freshwater habitats seem to be greater than in terrestrial ecosystems, especially because freshwater invasive species have a greater tendency to disperse (; ). Aqua Aquarius Скачать Mp3 here. Additionally, the importance of freshwater environments to humankind is enormous and modifications on its services will have a strong impact on human welfare. Crayfish species have social, economic and ecological significance in several regions around the world, favouring their introduction into allochthonous areas ( ). Procambarus clarkii is among these successfully and widely translocated species, and its importance is mainly associated with aquaculture and the aquarium trade, being the most harvested crayfish species in the world and thus, the most intentionally introduced (; ). The great concern regarding this species intensive introduction is that P. Clarkii is a successful colonizer which has specific features that increase its invasive ability and favours its colonization success across the world, in different climatic and geographic areas; these features are its ecological plasticity, resistant gene pool to population changes, adaptation of its biology and life cycle to changing environmental conditions, high tolerance to salinity, oxygen and temperature variations, high somatic growth and reproductive output, short development time and flexible feeding strategy (;; ). Therefore, after establishment, this species may quickly become a keystone species and cause serious changes in native plant and animal communities, altering water quality and sediment characteristics ( ).

In many countries, legislation designed to prevent crayfish spread is unsuccessful and causes conflicts due to the strong relationship between humans and crayfish that results from its recreational or commercial importance. However, once introduced into favorable habitats, P. Clarkii is difficult to eliminate ( ). Different management and control methods were cited before although their applicability and efficiency is site-specific. Unsuccessful population control is frequent and apparently solutions cannot be standardized. Therefore, the most economically and environmentally effective technique is prevention of introduction and range expansion.

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