Conservation genetics as a tool for conservation and management of the native Japanese freshwater crayfish Cambaroides japonicus (De Haan)
Author Dawkins, Kat, Furse, James
Published 2012
Journal Title Crustacean Research
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Griffith Research Online https://research-repository.griffith.edu.au CRUSTACEAN RESEARCH, SPECIAL NUMBER 7: 35–43, 2012 34
Statzner, B., Fievet, E., Champagne,J. Coughran J.–y., & J. M.Journal Furse of Crustacean Biology, 21: 354–359. Morel, R., & Herouin, E., 2000. Crayfish as Wells, S. M., Pyle, R. M., & Collins, N. M., 1983. Conservation genetics as a tool for conservation and geomorphic agents and ecosystem engineers: The IUCN Invertebrate Red Data Book. The management of the native Japanese freshwater crayfish Biological behavior affects sand and gravel IUCN, Gland, Switzerland. 632 pp. ISBN erosion in experimental streams. Limnology and 2–88032–602–X. Cambaroides japonicus (De Haan) Oceanography, 45: 1030–1040. Statzner, B., Peltret, O., & Tomanova, S., 2003. Crayfish as geomorphic agents and ecosystem Addresses: (JC) Environmental Futures engineers: effect of a biomass gradient on Centre and Griffith School of Environment, baseflow and flood–induced transport of gravel Gold Coast campus, Griffith University, Kathryn L. Dawkins and James M. Furse and sand in experimental streams. Freshwater Queensland 4222, Australia, and Outback Biology, 48: 147–163. Ecology, 1/73 Troy Terrace, Jolimont, Stone, E. L., 1993. Soil burrowing and mixing by Western Australia, 6014, Australia, (JMF) Abstract.—Unprecedented rates of extinction Conservation Monitoring Centre, 1992). a crayfish. Soil Science Society of America Environmental Futures Centre and Griffith have lead to the development of fields, such as While it has been established that terrestrial, Journal, 57: 1096–1099. School of Environment, Gold Coast campus, conservation genetics, in an attempt to better marine, and freshwater ecosystems have all Unestam, T., 1973. Significance of diseases on understand biodiversity and consequently devise been affected, extinction rates are difficult freshwater crayfish. Freshwater Crayfish, 13: Griffith University, Queensland 4222, Australia; conservation programs to maintain the genetic to accurately estimate due to our incomplete 135–150. integrity of species. We discuss the utility and Usio, N., & Townsend, C. R., 2001. The significance understanding of current and past levels Email: (JC) jason.coughran@ application of conservation genetics, using of the crayfish Paranephrops zealandicus as of biodiversity. Nevertheless, previous outbackecology.com, (JMF) j.furse@griffith. examples from Australian freshwater crayfish, shredders in a New Zealand headwater stream. edu.au researchers have identified that biodiversity with a view to applying this methodology to within freshwater ecosystems is at particular the sole native crayfish species found in Japan, risk (e.g., Saunders, 1976; Dudgeon, 1984; Cambaroides japonicus (De Haan). This species Sala et al., 2000; Cook et al., 2008); although is threatened by the non–indigenous crayfish this environment is relatively understudied in Pacifastacus leniusculus (Dana) through comparison to terrestrial and marine systems. displacement, competition pressures, and susceptibility to Aphanomyces astaci Schikora Recent studies employing molecular (the crayfish plague, carried by the invasive techniques have revealed cryptic diversity in species). Examining the genetic diversity within formerly well–studied groups of organisms C. japonicus could allow populations of high (Baker et al., 2004), and cryptic genetic conservation priority to be identified (i.e., divergence in the world’s largest and genetically distinct populations) or, in contrast, arguably most iconic freshwater crayfish show that this species is genetically homogenous. species Astacopsis gouldi Clark (Sinclair et Such genetic continuity may suggest that other al., 2011). These studies highlighted both our conservation methods (e.g., translocations and incomplete understanding of well–known restocking populations) may be suitable for this groups and species, but also that very high species of crayfish. Identifying genetic variability, group– and species–level conservation values or lack thereof, is a key step in dictating the can be easily overlooked. future direction of any conservation measures for This review is concerned with the status C. japonicus. of one particular group of aquatic organisms, freshwater crayfish, which are found on every continent (excluding the African INTRODUCTION (bar Madagascar), Antarctic, and Indian It is apparent that throughout the world, sub–continent) and are widely regarded as population declines and species’ extinctions keystone species (sensu Krebs, 1994) in are occurring at an unprecedented rate areas they inhabit (Hart, 1992; Momot, 1995; (Dirzo & Raven, 2003; DeSalle & Amato, Holdich, 2002c). 2004; May, 2010). Although extinctions are In 2010, the world’s freshwater crayfish natural phenomena, evidence suggests that were assessed for the IUCN Red List of current extinction rates far exceed those Threatened Species, and the evaluation seen in the past and are likely, for the most of 528 species established that freshwater part, attributable to human actions (World crayfish are among the world’s 5 most CONSERvATION OF C. jAPoniCuS USING GENETICS 37 36 36 threatened groups of animals (Dewhurst,K. L. DawKins &Kawai J. M. Furse & Machino, 2010). reproductive success in native species (e.g., reveals many similarities to Europe. Studies 2010 personal communication; IUCN, For example, some iconic, and Söderbäck, 1991; 1994; Westman et al., have compared the biology and ecology of 2011). As first highlighted by Wells et al. comparatively well–studied species are 2002). The United Kingdom (UK) and parts the native C. japonicus and non–indigenous P. (1983), the main threats to invertebrates facing a suite of well known and serious of mainland Europe are good examples of leniusculus and concluded that P. leniusculus and freshwater crayfish are: 1) habitat threats, and are consequently of considerable where entire populations of native species is likely to out–compete C. japonicus for destruction, 2) pollution, 3) non–indigenous conservation concern; yet they are currently have been extirpated, and where surviving habitat and resources, owing to its higher species, and 4) human exploitation (Wells et classified as DD. The World’s 2nd largest native populations are still at threat from the fecundity (and generally higher reproductive al., 1983; IUCN, 2011). Recently identified species of freshwater crayfish, Euastacus invading species (Evans & Edgerton, 2002; capacity), higher growth rates, and resistance emerging threats include changes to climate armatus (von Martens) from Australia is one Jussila et al., 2008; Bjurström et al., 2010; to diseases including the crayfish plague and increasing environmental temperature such species; the species faces numerous Peay et al., 2010). (Yamanaka et al., 1997; Nakata et al., 2004). (Horwitz, 1990a; 2010; Coughran & Furse, serious threats, but a simple lack of quantified Eight native crayfish species are Direct predation and increased success in 2010; Furse & Coughran, 2011a; Furse et al., data on its contemporary distribution and recognised in Europe, and while not all have obtaining shelter/refuges (a limiting variable 2012a) and increased intensity and frequency abundance does not permit assessment versus been assessed for the IUCN Red List (Table for presence and density of crayfish) and of severe weather events (Furse et al., IUCN Red List criteria (Furse & Coughran, 1), the presence of invasive crayfish has been reduced refuge availability have also been 2012b). 2011b; IUCN, 2011). The DD assessment identified as a threat to all native species indicated as threats to the survival prospects The objective of this paper is to review for C. japonicus highlights the need for by the IUCN (2011) and a number of other of C. japonicus (Nakata et al., 2006). the conservation status of the sole native more basic research on its contemporary workers (e.g., Harlioğlu & Harlioğlu, 2006; Japanese species of freshwater crayfish distribution and occupancy of habitat. Peay et al., 2010). Research imperatives and Methods for inves- (Cambaroides japonicus (De Haan)), and In their 2010 assessment, Kawai and There are at least 5 non–native crayfish tigation of C. japonicus introduce and discuss the applications of Machino identified key threats to C. species in Europe, including known carriers Addressing the previously documented existing techniques/methods that could japonicus, the most serious of which was the of the crayfish plague (e.g., P. leniusculus knowledge gaps for C. japonicus should assist in conservation efforts for this species. spread of the non–native and highly invasive and Procambarus Clarkii (Girard)) and as be accorded high priority (and allocated Examples of how these techniques have American signal crayfish, Pacifastacus such are considered as very serious threats to appropriate funding) as the resulting been applied in the context of Australian leniusculus (Dana), which was introduced to the native species (IUCN, 2011). information will allow a thorough evaluation freshwater crayfish will be provided, with the Japanese main Islands in 1928 and has Europeans have a long history of social of the species’ conservation status, and any particular focus on determining levels of been expanding its distribution since that customs and traditions associated with conservation and management measures genetic diversity and how this data can time (Nakata & Goshima, 2003). crayfish (e.g., the annual Swedish “Crayfish that may be required. In particular, it is be used for conservation assessments and There is great deal of concern over the Parties”), as well as industries such as wild imperative that the geographical distribution recovery and management efforts. continued spread of this crayfish, including fisheries and aquaculture (Holdich, 2002b; and taxonomic uncertainties (i.e., any cryptic any human translocations, for a numbers 2002a), and these traditions and industries diversity) of C. japonicus are investigated Current Conservation Status of C. japonicus of reasons: 1) the species is robust and can are threatened by invasive species. As non– and clarified. Cambaroides japonicus has been assessed tolerate a wide range of environmental native species are reasonably resilient and In light of experience elsewhere in the versus IUCN Red List criteria as Data world, genetic techniques can play a central 1 conditions, including near arctic (i.e., latitude robust, quite widely distributed, and typically Deficient (DD) with a declining population 62°N, Heinimaa & Pursiainen, 2008), 2) P. such dominant competitors, it is considered role in facilitating the timely collection of trend (Kawai & Machino, 2010). The DD leniusculus has superior competitive abilities unlikely that native species will be able to information and subsequent conservation assessment was primarily due to a lack of over the smaller C. japonicus, and 3) P. recover to pre–invasion levels (Holdich et evaluations that are required for this species. information regarding its distribution and leniusculus is a known ‘healthy carrier’ of al., 2009). Genetic techniques have been used in a extent of occurrence (EOO) defined by the Aphanomyces astaci Schikora (hereafter, Examination of the situation in Japan number of countries to assess factors such IUCN as “the area contained within the crayfish plague) which is highly contagious shortest continuous imaginary boundary and uniformly fatal to susceptible species which can be drawn to encompass all the (which includes C. japonicus) (Evans & known, inferred or projected sites of present Edgerton, 2002). Table 1. Native European Freshwater Crayfish and Current IUCN Status (Fetzner, 2011; IUCN, 2011) occurrence of a taxon” (IUCN 2010). The The invasive potential, competitive current DD assessment for C. japonicus advantage, and capacity of P. leniusculus Species and Authority IUCN Red List Status and Population Trend should not be interpreted as a stable or to act as a disease vector (i.e., to spread Austropotamobius pallipes (Lereboullet) Endangered (EN), Trend: Decreasing improving conservation status as it is known crayfish plague between waterbodies) are Astacus astacus (Linnaeus) vulnerable (vU), Trend: Decreasing the species faces a number of very serious well documented and reasonably well Astacus leptodactylus (Eschscholtz) Least Concern (LC), Trend: Unknown and ongoing threats (Nakata et al., 2002; understood in regions where the species has Austropotamobius torrentium (Shrank) DD, Trend: Decreasing been introduced, spread, and been intensively Astacus pachypus (Rathke) DD, Trend: Unknown
1 studied. Additional threats to native species Austropotamobius italicus italicus (Faxon) Not Assessed, Trend: Unknown : i.e., “inadequate information to make a direct, or Austropotamobius italicus carsicus M. Karaman Not Assessed, Trend: Unknown indirect, assessment of its risk of extinction based on its are via interspecific competition and distribution and/or population status” (IUCN, 2011). interspecific breeding, resulting in reduced Austropotamobius italicus carinthiacus Albrecht Not Assessed, Trend: Unknown CONSERvATION OF C. jAPoniCuS USING GENETICS 37 36 37 threatened groups of animals (Dewhurst,K. L. DawKins &Kawai J. M. Furse & Machino, 2010). reproductive success in native species (e.g., reveals many similarities to Europe. Studies 2010 personal communication; IUCN, For example, some iconic, and Söderbäck, 1991; 1994; Westman et al., have compared the biology and ecology of 2011). As first highlighted by Wells et al. comparatively well–studied species are 2002). The United Kingdom (UK) and parts the native C. japonicus and non–indigenous P. (1983), the main threats to invertebrates facing a suite of well known and serious of mainland Europe are good examples of leniusculus and concluded that P. leniusculus and freshwater crayfish are: 1) habitat threats, and are consequently of considerable where entire populations of native species is likely to out–compete C. japonicus for destruction, 2) pollution, 3) non–indigenous conservation concern; yet they are currently have been extirpated, and where surviving habitat and resources, owing to its higher species, and 4) human exploitation (Wells et classified as DD. The World’s 2nd largest native populations are still at threat from the fecundity (and generally higher reproductive al., 1983; IUCN, 2011). Recently identified species of freshwater crayfish, Euastacus invading species (Evans & Edgerton, 2002; capacity), higher growth rates, and resistance emerging threats include changes to climate armatus (von Martens) from Australia is one Jussila et al., 2008; Bjurström et al., 2010; to diseases including the crayfish plague and increasing environmental temperature such species; the species faces numerous Peay et al., 2010). (Yamanaka et al., 1997; Nakata et al., 2004). (Horwitz, 1990a; 2010; Coughran & Furse, serious threats, but a simple lack of quantified Eight native crayfish species are Direct predation and increased success in 2010; Furse & Coughran, 2011a; Furse et al., data on its contemporary distribution and recognised in Europe, and while not all have obtaining shelter/refuges (a limiting variable 2012a) and increased intensity and frequency abundance does not permit assessment versus been assessed for the IUCN Red List (Table for presence and density of crayfish) and of severe weather events (Furse et al., IUCN Red List criteria (Furse & Coughran, 1), the presence of invasive crayfish has been reduced refuge availability have also been 2012b). 2011b; IUCN, 2011). The DD assessment identified as a threat to all native species indicated as threats to the survival prospects The objective of this paper is to review for C. japonicus highlights the need for by the IUCN (2011) and a number of other of C. japonicus (Nakata et al., 2006). the conservation status of the sole native more basic research on its contemporary workers (e.g., Harlioğlu & Harlioğlu, 2006; Japanese species of freshwater crayfish distribution and occupancy of habitat. Peay et al., 2010). Research imperatives and Methods for inves- (Cambaroides japonicus (De Haan)), and In their 2010 assessment, Kawai and There are at least 5 non–native crayfish tigation of C. japonicus introduce and discuss the applications of Machino identified key threats to C. species in Europe, including known carriers Addressing the previously documented existing techniques/methods that could japonicus, the most serious of which was the of the crayfish plague (e.g., P. leniusculus knowledge gaps for C. japonicus should assist in conservation efforts for this species. spread of the non–native and highly invasive and Procambarus Clarkii (Girard)) and as be accorded high priority (and allocated Examples of how these techniques have American signal crayfish, Pacifastacus such are considered as very serious threats to appropriate funding) as the resulting been applied in the context of Australian leniusculus (Dana), which was introduced to the native species (IUCN, 2011). information will allow a thorough evaluation freshwater crayfish will be provided, with the Japanese main Islands in 1928 and has Europeans have a long history of social of the species’ conservation status, and any particular focus on determining levels of been expanding its distribution since that customs and traditions associated with conservation and management measures genetic diversity and how this data can time (Nakata & Goshima, 2003). crayfish (e.g., the annual Swedish “Crayfish that may be required. In particular, it is be used for conservation assessments and There is great deal of concern over the Parties”), as well as industries such as wild imperative that the geographical distribution recovery and management efforts. continued spread of this crayfish, including fisheries and aquaculture (Holdich, 2002b; and taxonomic uncertainties (i.e., any cryptic any human translocations, for a numbers 2002a), and these traditions and industries diversity) of C. japonicus are investigated Current Conservation Status of C. japonicus of reasons: 1) the species is robust and can are threatened by invasive species. As non– and clarified. Cambaroides japonicus has been assessed tolerate a wide range of environmental native species are reasonably resilient and In light of experience elsewhere in the versus IUCN Red List criteria as Data world, genetic techniques can play a central 1 conditions, including near arctic (i.e., latitude robust, quite widely distributed, and typically Deficient (DD) with a declining population 62°N, Heinimaa & Pursiainen, 2008), 2) P. such dominant competitors, it is considered role in facilitating the timely collection of trend (Kawai & Machino, 2010). The DD leniusculus has superior competitive abilities unlikely that native species will be able to information and subsequent conservation assessment was primarily due to a lack of over the smaller C. japonicus, and 3) P. recover to pre–invasion levels (Holdich et evaluations that are required for this species. information regarding its distribution and leniusculus is a known ‘healthy carrier’ of al., 2009). Genetic techniques have been used in a extent of occurrence (EOO) defined by the Aphanomyces astaci Schikora (hereafter, Examination of the situation in Japan number of countries to assess factors such IUCN as “the area contained within the crayfish plague) which is highly contagious shortest continuous imaginary boundary and uniformly fatal to susceptible species which can be drawn to encompass all the (which includes C. japonicus) (Evans & known, inferred or projected sites of present Edgerton, 2002). Table 1. Native European Freshwater Crayfish and Current IUCN Status (Fetzner, 2011; IUCN, 2011) occurrence of a taxon” (IUCN 2010). The The invasive potential, competitive current DD assessment for C. japonicus advantage, and capacity of P. leniusculus Species and Authority IUCN Red List Status and Population Trend should not be interpreted as a stable or to act as a disease vector (i.e., to spread Austropotamobius pallipes (Lereboullet) Endangered (EN), Trend: Decreasing improving conservation status as it is known crayfish plague between waterbodies) are Astacus astacus (Linnaeus) vulnerable (vU), Trend: Decreasing the species faces a number of very serious well documented and reasonably well Astacus leptodactylus (Eschscholtz) Least Concern (LC), Trend: Unknown and ongoing threats (Nakata et al., 2002; understood in regions where the species has Austropotamobius torrentium (Shrank) DD, Trend: Decreasing been introduced, spread, and been intensively Astacus pachypus (Rathke) DD, Trend: Unknown
1 studied. Additional threats to native species Austropotamobius italicus italicus (Faxon) Not Assessed, Trend: Unknown : i.e., “inadequate information to make a direct, or Austropotamobius italicus carsicus M. Karaman Not Assessed, Trend: Unknown indirect, assessment of its risk of extinction based on its are via interspecific competition and distribution and/or population status” (IUCN, 2011). interspecific breeding, resulting in reduced Austropotamobius italicus carinthiacus Albrecht Not Assessed, Trend: Unknown CONSERvATION OF C. jAPoniCuS USING GENETICS 39 38 38 as: 1) levels of genetic diversityK. withinL. DawK ains &2008). J. M. Furse Due to extensive translocations and such extensive and expansive movements coastal regions of Australia are becoming species, allowing prioritisation of additional hybridisation (Bunn et al., 2008), the genetic of C. destructor may have resulted in increasingly urbanised (ABS, 2009), data collection for conservation assessments, integrity of this species has been raised as a genetic homogenisation throughout the threatening many of the remaining habitat 2) effective population sizes, 3) need to serious conservation concern. Maintaining species. Examination of the mtDNA 16S fragments that these species occupy (Davie, manage captive populations of threatened genetic diversity (and eliminating hybrids) is gene region by Nguyen et al. (2004) found 2007; Coughran et al., 2008). three major lineages within this species fauna, 4) identifying the extent of disease a management priority for both commercial Conservation implications of identifying susceptibility, and 5) identifying introduction and wild populations (see Bunn et al., 2008). and consequently recommended them as ESUs. Cherax destructor populations cryptic diversity points of invasive organisms for policing Nguyen et al. (2002), examined the An example of the implications of purposes (Avise, 1989; O'Brien, 1994; genetic variation within the mtDNA gene are under considerable pressure from a large and widespread recreational fishery, identifying cryptic diversity in a reasonably Frankham, 1995b; 1995a; Amos & Balmford, region 16S between populations throughout well–known and widespread species 2001; DeSalle & Amato, 2004). Western Australia as well as translocated aquaculture activities, and ongoing rural development (e.g., agriculture)(Austin et is provided by the case of Euastacus As potential uses of genetic techniques populations located in South Australia sulcatus Riek (IUCN Status: vU). The cover such wide areas of application, a series and victoria. Two distinct genetic groups al., 2003; Nguyen et al., 2004). However, the close geographic proximity of the three species contains a number of small and of case studies are provided below to outline were identified (one from Margaret River geographically isolated populations, and applicability and utility of evaluating genetic in southern Western Australia, the other discrete lineages is an additional concern for maintaining the genetic integrity of wild C. Coughran & Furse (2010) and Furse & diversity for conservation purposes. consisting of all other populations that were Coughran (2011b) identified the possibilty evaluated); it was recommended that both destructor populations (Austin et al., 2003). Genetic diversity in Australian crayfish spe- that genetic analysis could identify a number groups be recognised as ESUs, with the Tenuibranchiurus glypticus Riek of genetically distinct populations (or distinct cies Margaret River ESU represented by only Examining genetic diversity within The sole species in this genus is the species). If assessed versus IUCN criteria a single population. The extremely high smallest freshwater crayfish species in at the population level, as opposed to the populations for the purpose of conservation conservation value of the Margaret River is part of the relatively new field of Australia: IUCN status EN (IUCN, 2011). current species–level assessment, some E. population (versus the other populations) was Due to its very small size with a total sulcatus populations would warrant listing as conservation genetics. Conservation highlighted in this study, as it was “restricted genetics combines genetic information with length (rostrum – telson) < 35 mm, and Critically Endangered or EN (Coughran & to this single river system and represents highly fragmented distribution, this crayfish Furse 2010). There is potential for a similar ecological and distributional information the only significant genetic diversity within to identify geographic regions or localities has effectively no dispersal capabilities situation to apply in the case of C. japonicus the species so far recorded” (Nguyen et al., (Coughran et al., 2010; Dawkins et al., in Japan. where species are most at risk (O'Brien, 2002). 1994). Identifying geographical areas where 2010). The species is restricted to coastal Without identification of this genetic Melaleuca swamp habitat along the central Application of conservation genetics in unique and/or discrete genetic populations diversity (i.e., distinct genetic structure), japan exist effectively prioritises these areas as eastern coast of Australia, with habitat consequences for the Margaret River patchiness and biological restrictions (e.g., As the geographical range of P. ‘of conservation concern’. One common population could have been dire, including; leniusculus appears to be increasing in Japan, method for delineating such populations small size) thought to be the cause of the translocations, potential cross–breeding, high levels of genetic divergence between and, as no successful methods of eradicating is by identifying them as evolutionarily and quite possibly extirpation of the unique this species are known, the primary focus significant units (ESUs, Moritz, 1994), isolated populations (Dawkins et al., 2010). genetic stock in the population. The Nguyen Examination of both mtDNA (Dawkins et of conservation efforts for C. japonicus using information obtained from both et al. (2002) study also identified negative should include identifying any populations mitochondrial (mtDNA) and nuclear DNA al., 2010) and nuDNA (authors unpublished genetic outcomes such as introgression and data) from the various geographically that possess unique genetic information (i.e., (nuDNA) (as outlined in Moritz, 1994). outbreeding depression, which can lead to cryptic diversity), identifying them as ESUs In the Australian context, ESUs have been isolated populations show very high levels decreased population fitness. of genetic diversity, and suggests recognition and assigning them critical conservation recognised in a variety of freshwater crayfish, status. Cherax destructor Clark of at least 5 ESUs is warranted based on both widespread and with highly restricted By identifying any priority areas of Cherax destructor h a s t h e m o s t mtDNA data (Dawkins et al., 2010). Genetic and/or fragmented distributions, and these critical habitat (i.e., areas containing ESUs), widespread natural geographic distribution data also indicates the presence of at least are outlined in the following sections. conservation management can first focus on in Australia and extends into the semi–arid one additional previously unidentified species those populations that are most ‘genetically Cherax tenuimanus (Smith) zone: IUCN Status vU (IUCN, 2011). (Dawkins et al., 2010). Such high levels valuable’ (and potentially saveable) and Cherax tenuimanus is endemic to Western Cherax destructor has a history of of previously unknown genetic divergence then efforts can be extended to remaining Australia and is one of the world’s largest widespread and long–range translocations highlight the utility of genetic methods/ populations of lesser priority. Any “invasion– freshwater crayfish species: IUCN Status both pre– and post–European settlement techniques as powerful conservation tools. In proof” areas of suitable C. japonicus habitat Critically Endangered (CR) (IUCN, 2011). (Horwitz, 1990b). These translocations are this particular case it has allowed a potential should also be identified. Isolated areas of This is an important species for on–going (Coughran et al., 2009; Coughran new species to be identified, and genetically habitat suitable for C. japonicus (i.e., without aquaculture (Imgrund et al., 1997) and for & Daly, 2012) and are of considerable unique populations to be considered and non–indigenous crayfish, and not threatened cultural reasons due to a large and well concern for conservation of native species. treated as extremely high conservation by invasion by non–indigenous crayfish) established recreational fishery (Bunn et al., Horwitz & Knott (1995) suggested that priorities. This is particularly important as CONSERvATION OF C. jAPoniCuS USING GENETICS 39 38 39 as: 1) levels of genetic diversityK. withinL. DawK ains &2008). J. M. Furse Due to extensive translocations and such extensive and expansive movements coastal regions of Australia are becoming species, allowing prioritisation of additional hybridisation (Bunn et al., 2008), the genetic of C. destructor may have resulted in increasingly urbanised (ABS, 2009), data collection for conservation assessments, integrity of this species has been raised as a genetic homogenisation throughout the threatening many of the remaining habitat 2) effective population sizes, 3) need to serious conservation concern. Maintaining species. Examination of the mtDNA 16S fragments that these species occupy (Davie, manage captive populations of threatened genetic diversity (and eliminating hybrids) is gene region by Nguyen et al. (2004) found 2007; Coughran et al., 2008). three major lineages within this species fauna, 4) identifying the extent of disease a management priority for both commercial Conservation implications of identifying susceptibility, and 5) identifying introduction and wild populations (see Bunn et al., 2008). and consequently recommended them as ESUs. Cherax destructor populations cryptic diversity points of invasive organisms for policing Nguyen et al. (2002), examined the An example of the implications of purposes (Avise, 1989; O'Brien, 1994; genetic variation within the mtDNA gene are under considerable pressure from a large and widespread recreational fishery, identifying cryptic diversity in a reasonably Frankham, 1995b; 1995a; Amos & Balmford, region 16S between populations throughout well–known and widespread species 2001; DeSalle & Amato, 2004). Western Australia as well as translocated aquaculture activities, and ongoing rural development (e.g., agriculture)(Austin et is provided by the case of Euastacus As potential uses of genetic techniques populations located in South Australia sulcatus Riek (IUCN Status: vU). The cover such wide areas of application, a series and victoria. Two distinct genetic groups al., 2003; Nguyen et al., 2004). However, the close geographic proximity of the three species contains a number of small and of case studies are provided below to outline were identified (one from Margaret River geographically isolated populations, and applicability and utility of evaluating genetic in southern Western Australia, the other discrete lineages is an additional concern for maintaining the genetic integrity of wild C. Coughran & Furse (2010) and Furse & diversity for conservation purposes. consisting of all other populations that were Coughran (2011b) identified the possibilty evaluated); it was recommended that both destructor populations (Austin et al., 2003). Genetic diversity in Australian crayfish spe- that genetic analysis could identify a number groups be recognised as ESUs, with the Tenuibranchiurus glypticus Riek of genetically distinct populations (or distinct cies Margaret River ESU represented by only Examining genetic diversity within The sole species in this genus is the species). If assessed versus IUCN criteria a single population. The extremely high smallest freshwater crayfish species in at the population level, as opposed to the populations for the purpose of conservation conservation value of the Margaret River is part of the relatively new field of Australia: IUCN status EN (IUCN, 2011). current species–level assessment, some E. population (versus the other populations) was Due to its very small size with a total sulcatus populations would warrant listing as conservation genetics. Conservation highlighted in this study, as it was “restricted genetics combines genetic information with length (rostrum – telson) < 35 mm, and Critically Endangered or EN (Coughran & to this single river system and represents highly fragmented distribution, this crayfish Furse 2010). There is potential for a similar ecological and distributional information the only significant genetic diversity within to identify geographic regions or localities has effectively no dispersal capabilities situation to apply in the case of C. japonicus the species so far recorded” (Nguyen et al., (Coughran et al., 2010; Dawkins et al., in Japan. where species are most at risk (O'Brien, 2002). 1994). Identifying geographical areas where 2010). The species is restricted to coastal Without identification of this genetic Melaleuca swamp habitat along the central Application of conservation genetics in unique and/or discrete genetic populations diversity (i.e., distinct genetic structure), japan exist effectively prioritises these areas as eastern coast of Australia, with habitat consequences for the Margaret River patchiness and biological restrictions (e.g., As the geographical range of P. ‘of conservation concern’. One common population could have been dire, including; leniusculus appears to be increasing in Japan, method for delineating such populations small size) thought to be the cause of the translocations, potential cross–breeding, high levels of genetic divergence between and, as no successful methods of eradicating is by identifying them as evolutionarily and quite possibly extirpation of the unique this species are known, the primary focus significant units (ESUs, Moritz, 1994), isolated populations (Dawkins et al., 2010). genetic stock in the population. The Nguyen Examination of both mtDNA (Dawkins et of conservation efforts for C. japonicus using information obtained from both et al. (2002) study also identified negative should include identifying any populations mitochondrial (mtDNA) and nuclear DNA al., 2010) and nuDNA (authors unpublished genetic outcomes such as introgression and data) from the various geographically that possess unique genetic information (i.e., (nuDNA) (as outlined in Moritz, 1994). outbreeding depression, which can lead to cryptic diversity), identifying them as ESUs In the Australian context, ESUs have been isolated populations show very high levels decreased population fitness. of genetic diversity, and suggests recognition and assigning them critical conservation recognised in a variety of freshwater crayfish, status. Cherax destructor Clark of at least 5 ESUs is warranted based on both widespread and with highly restricted By identifying any priority areas of Cherax destructor h a s t h e m o s t mtDNA data (Dawkins et al., 2010). Genetic and/or fragmented distributions, and these critical habitat (i.e., areas containing ESUs), widespread natural geographic distribution data also indicates the presence of at least are outlined in the following sections. conservation management can first focus on in Australia and extends into the semi–arid one additional previously unidentified species those populations that are most ‘genetically Cherax tenuimanus (Smith) zone: IUCN Status vU (IUCN, 2011). (Dawkins et al., 2010). Such high levels valuable’ (and potentially saveable) and Cherax tenuimanus is endemic to Western Cherax destructor has a history of of previously unknown genetic divergence then efforts can be extended to remaining Australia and is one of the world’s largest widespread and long–range translocations highlight the utility of genetic methods/ populations of lesser priority. Any “invasion– freshwater crayfish species: IUCN Status both pre– and post–European settlement techniques as powerful conservation tools. In proof” areas of suitable C. japonicus habitat Critically Endangered (CR) (IUCN, 2011). (Horwitz, 1990b). These translocations are this particular case it has allowed a potential should also be identified. Isolated areas of This is an important species for on–going (Coughran et al., 2009; Coughran new species to be identified, and genetically habitat suitable for C. japonicus (i.e., without aquaculture (Imgrund et al., 1997) and for & Daly, 2012) and are of considerable unique populations to be considered and non–indigenous crayfish, and not threatened cultural reasons due to a large and well concern for conservation of native species. treated as extremely high conservation by invasion by non–indigenous crayfish) established recreational fishery (Bunn et al., Horwitz & Knott (1995) suggested that priorities. This is particularly important as CONSERvATION OF C. jAPoniCuS USING GENETICS 41 40 40 K. L. DawKins &already J. M. Furse scarce resources to conservations Macroinvertebrates of Large Boreal Lakes: Sampled Red List Index Team, The Zoological that are able to be maintained in the long A Pilot Study in Lake Päijänne, Finland. Society of London, England. term could prove invaluable for conservation efforts and guide decision making processes Freshwater Crayfish, 17: 177–182. Dirzo, R., & Raven, P. H., 2003. Global State of this native Japanese species (sensu ‘Ark towards the best possible outcomes. Bunn, J., Koenders, A., Austin, C. M., & Horwitz, P., of Biodiversity and Loss. Annual Review of Sites’ in the UK) (Holdich et al., 2009). 2008. Identification of hairy, smooth and hybrid Environment and Resources, 28: 137–167. If ESUs are not indicated and populations Acknowledgements.—We are grateful for marron (Decapoda: Parastacidae) in the Margaret Dudgeon, D. 1984. The importance of streams in are identified as genetically homogenous, this Dr Tadashi Kawai for kindly inviting us to River: Morphology and allozymes. Freshwater tropical rain–forest systems. Tropical Rain– also provides other avenues for conservation attend and present a Lecture on this topic Crayfish, 16: 113–121. Forest: The Leeds Symposium, Leeds, England, at the “Conservation of freshwater crayfish Cook, B. D., Page, T. J., & Hughes, J. M., 2008. pp. 71–82. Special publication by The Leeds efforts. For instance, one of the primary Importance of cryptic species for identifying Philosophical and Literary Society. concerns for genetically distinct populations Symposium” at the Sapporo Maruyama ‘representative’ units of biodiversity Evans, L. H., & Edgerton, B. F., 2002. General (as previously outlined) is the potential Zoo, Hokkaido, Japan in 2011. We are for freshwater conservation. Biological Biology: Pathogens, Parasites and Commensals. for deleterious effects occurring through especially thankful to Mr Tanaka, Ms Notani, Conservation, 141: 2821–2831. In: D. M. Holdich (Ed.) Biology of Freshwater translocations. For example, if C. japonicus Dr Kawai, Dr Nonaka, Mr Takaesu, Mr Coughran, J., Dawkins, K. L., & Furse, J. M., 2008. Crayfish. Blackwell Science, Oxford, England. is genetically homogenous then it may be Miyamoto, and our under–ice diving team: An assessment of genus Tenuibranchiurus pp 377–438. ISBN 0–632–05431–X. possible and feasible for areas experiencing Mr Ishikawa, Mr Kitajima and Mrs. Ohtsuka, (1 species) versus IUCN Red List criteria. Fetzner, J. W., Jr, 2011. The crayfish and lobster population declines to be ‘restocked’ to for arranging and hosting us on our field trip unpublished report prepared for the for the taxonomy browser. Available online via: http:// to Lake Shikaribetsu. Thanks are extended global species conservation assessment of iz.carnegiemnh.org/crayfish/NewAstacidea/ mitigate any such reductions. crayfishes for the IUCN Red List of Threatened infraorder.asp?io=Astacidea Furthermore, if successful eradication to our editors and 2 anonymous reviewers Species. A report jointly prepared by the School Frankham, R., 1995a. Inbreeding and Extinction: methods are developed for P. leniusculus, for providing useful comments on an earlier of Environmental Science and Management A Threshold Effect. Conservation Biology, 9: areas previously occupied by the native version of this paper. This research project (Southern Cross University), and Griffith School 792–799. species could possibly be repopulated. Such was supported by the Australian Rivers of Environment (Griffith University), 2 pp. Frankham, R., 1995b. Conservation Genetics. methods would of course require careful Institute, and Environmental Futures Centre, Coughran, J., McCormack, R. B., & Daly, G., 2009. Annual Review of Genetics, 29: 305–327. consideration of other factors, including and the Griffith School of Environment, Translocation of the Yabby Cherax destructor Furse, J. M., & Coughran, J., 2011a. An assessment Griffith University, Gold Coast campus, into eastern drainages of New South Wales, of the distribution, biology, threatening the potential of inadvertently translocating Australia. Australian Zoologist, 35: 100–103. processes and conservation status of the diseases and organisms such as commensals Queensland, Australia. Coughran, J., Dawkins, K. L. & Furse, J. M. 2010. freshwater crayfish, genus Euastacus (Decapoda: and parasites. Tenuibranchiurus glypticus. In: IUCN 2010. Parastacidae), in Continental Australia. II. By simply examining the genetic diversity LITeRaTURe CITeD IUCN Red List of Threatened Species. version Threats, Conservation Assessments and Key within C. japonicus, a number of potential 2010.4.
streams: the importance of species interactions,K. L. DawKins & J. M.Project Furse in the River Pyhäjoki, Western Finland: 5755. Dana. Nordic Journal of Freshwater Research, physical factors, and chance. Oecologia, 91: A Case Study. Freshwater Crayfish, 16: 51–56. Peay, S., Holdich, D. M., & Brickland, J., 2010. Risk 69: 137–143. 220–228. Kawai, T., & Machino, Y., 2010. Cambaroides Assessments of Non–Indigenous Crayfish in Wells, S. M., Pyle, R. M., & Collins, N. M., 1983. Heinimaa, S., & Pursiainen, M., 2008. Signal japonicus. In: IUCN 2011. IUCN Red List Great Britain. Freshwater Crayfish, 17: 109–122. The IUCN Invertebrate Red Data Book. The Crayfish Pacifastacus leniusculus at Northerly of Threatened Species. version 2011.1., The Sala, O. E., Chapin, F. S., Armesto, J. J., Berlow, IUCN, Gland, Switzerland. 632 pp. ISBN Latitudes: A Search for the Distribution Limits. International Union for Conservation of Nature E., Bloomfield, J., Dirzo, R., Huber–Sanwald, 2–88032–602–X. Freshwater Crayfish, 16: 37–41. and Natural Resources. Gland, Switzerland. E., Huenneke, L. F., Jackson, R. B., Kinzig, Westman, K., Savolainen, R., & Julkunen, M., 2002. Holdich, D. M., 2002a. Part 3: Conclusions. In: D. M. Krebs, C. J., 1994. Ecology: the experimental A., Leemans, R., Lodge, D. M., Mooney, H. Replacement of the native crayfish Astacus Holdich (Ed.) Biology of Freshwater Crayfish. analysis of distribution and abundance. A., Oesterheld, M., Poff, N. L., Sykes, M. T., astacus by the introduced species Pacifastacus Blackwell Science, Oxford, England. pp 673– HarperCollins, New York, USA. 801 pp. ISBN Walker, B. H., Walker, M., & Wall, D. H., 2000. leniusculus in a small, enclosed Finnish lake: a 682. ISBN 0–632–05431–X. 0–06–500410–8. Global Biodiversity Scenarios for the Year 2100. 30–year study. Ecography, 25: 53–73. Holdich, D. M., 2002b. General Biology: May, R. M., 2010. Ecological science and Science, 287: 1770–1774. World Conservation Monitoring Centre, 1992. Background and Functional Morphology. In: tomorrow's world. Philosophical Transactions of Saunders, G. W., 1976. Decomposition in freshwater. Global Biodiversity: Status of the Earth's living D. M. Holdich (Ed.) Biology of Freshwater the Royal Society B, 365: 41–47. In: J. M. Anderson & A. Macfadyen (Eds), The resources. Chapman & Hall, London. 614 pp. Crayfish. Blackwell Science, Oxford, England. Momot, W. T., 1995. Redefining the role of crayfish Role of Terrestrial and Aquatic Organisms in Yamanaka, K., Kuwabara, R., & Shio, T., 1997. pp 3–29. ISBN 0–632–05431–X. in aquatic ecosystems. Reviews in Fisheries Decomposition Processes. Blackwell Scientific Larval development of a Japanese crayfish, Holdich, D. M., 2002c. Background and Functional Science, 3: 33–63. Publications, Oxford, England.: pp 341–373. Cambaroides japonicus (De Haan). Bulletin of Morphology. In: D. M. Holdich (Ed.) Biology of Moritz, C., 1994. Defining ‘Evolutionarily ISBN 0–632–00018–X. Marine Science, 61: 165–175. Freshwater Crayfish. Blackwell Science, Oxford. Significant Units’ for conservation. Trends in Sinclair, E. A., Madsen, A., Walsh, T., Nelson, J., & pp 3–29. Ecology and Evolution, 9: 373–375. Crandall, K. A., 2011. Cryptic genetic divergence Holdich, D. M., Reynolds, J. D., Souty–Grosset, Nakata, K., Hamano, T., Hayashi, K., & Kawai, in the giant Tasmanian freshwater crayfish Addresses: (KLD) Australian Rivers C., & Sibley, P. J., 2009. A review of the ever T., 2002. Lethal limits of high temperature for Astacopsis gouldi (Decapoda: Parastacidae): Institute and Griffith School of Environment, increasing threat to European crayfish from non– two crayfishes, the native species Cambaroides implications for conservation. Animal Gold Coast campus, Griffith University, indigenous crayfish species. Knowledge and japonicus and the alien species Pacifastacus Conservation, 14: 87–97. Queensland 4222, Australia, (JMF) Management of Aquatic Ecosystems, (394–395): leniusculus in Japan. Fisheries Science, 68: Söderbäck, B., 1991. Interspecific dominance relationship and aggressive interactions in the Environmental Futures Centre and Griffith 46 pp. 763–767. School of Environment, Gold Coast campus, Horwitz, P., 1990a. The conservation status of Nakata, K., & Goshima, S., 2003. Competition for freshwater crayfishes Astacus astacus (L.) and Australian freshwater crayfish with a provisional shelter of preferred sizes between the native Pacifastacus leniusculus (Dana). Canadian Griffith University, Queensland 4222, list of threatened species, habitats and potentially crayfish species Cambaroides japonicus and the Journal of Zoology, 69: 1321–1325. Australia; threatening processes. Report series No. 14. alien crayfish species Pacifastacus leniusculus Söderbäck, B., 1994. Reproductive interference Email addresses: (KLD) k.dawkins@ Australian National Parks and Wildlife Service, in Japan in relation to prior residence, sex between two co–occurring crayfish species, Astacus astacus L. and Pacifastacus leniusculus griffith.edu.au, (JMF) [email protected] Canberra, Australia. 121 pp. difference and body size. Journal of Crustacean – Horwitz, P., 1990b. The Translocation of Freshwater Biology, 23: 897–907. Crayfish in Australia: Potential Impact, the Need Nakata, K., Tanaka, A., & Goshima, S., 2004. for Control and Global Relevance. Biological Reproduction of the alien crayfish species Conservation, 54: 291–305. Pacifastacus leniusculus in Lake Shikaribetsu, Horwitz, P., & Knott, B., 1995. The distribution Hokkaido, Japan. Journal of Crustacean Biology, and spread of the yabby Cherax complex in 24: 496–501. Australia: speculations, hypotheses and the need Nakata, K., Tstutsumi, K., Kawai, T., & Goshima, for research. Freshwater Crayfish, 10: 81–91. S., 2006. Coexistence of two North American Horwitz, P., 2010. The Conservation Status of invasive crayfish species, Pacifastacus Freshwater Crayfish: The Basis for Concern, leniusculus (Dana, 1852) and Procambarus Listing and Recovery Processes, and Community clarkii (Girard, 1852) in Japan. Crustaceana, 78: Involvement. Freshwater Crayfish, 17: 1–12. 1389–1394. Imgrund, J., Groth, D., & Wetherall, J., 1997. Nguyen, T. T. T., Meewan, M., Ryan, S., & Austin, Genetic analysis of the freshwater crayfish C. M., 2002. Genetic diversity and translocation Cherax tenuimanus. Electrophoresis, 18: 1660– in the marron, Cherax tenuimanus (Smith): 1665. implications for management and conservation. IUCN, 2010. Guidelines for Using the IUCN Red Fisheries Management and Ecology, 9: 163–173. List Categories and Criteria: version 8. Prepared Nguyen, T. T. T., Austin, C. M., Meewan, M. by the Standards and Petitions Working Group M., Schultz, M. B., & Jerry, D. R., 2004. for the IUCN SSC Biodiversity Assessments Phylogeography of the freshwater crayfish Sub–Committee 2010. 60 pp. Cherax destructor Clark (Parastacidae) in inland IUCN, 2011. The IUCN Red List of Threatened Australia: historical fragmentation and recent Species (version 2011.1), The International range expansion. Biological Journal of the Union for Conservation of Nature and Natural Linnean Society, 83: 539–550. Resources. Gland, Switzerland. O'Brien, S. J., 1994. A role for molecular genetics Jussila, J., Ojala, K., & Mannonen, A., 2008. Noble in biological conservation. Proceedings of the Crayfish (Astacus astacus) Reintroduction National Academy of Sciences USA 91: 5748– CONSERvATION OF C. jAPoniCuS USING GENETICS 43 42 43 streams: the importance of species interactions,K. L. DawKins & J. M.Project Furse in the River Pyhäjoki, Western Finland: 5755. Dana. Nordic Journal of Freshwater Research, physical factors, and chance. Oecologia, 91: A Case Study. Freshwater Crayfish, 16: 51–56. Peay, S., Holdich, D. M., & Brickland, J., 2010. Risk 69: 137–143. 220–228. Kawai, T., & Machino, Y., 2010. Cambaroides Assessments of Non–Indigenous Crayfish in Wells, S. M., Pyle, R. M., & Collins, N. M., 1983. Heinimaa, S., & Pursiainen, M., 2008. Signal japonicus. In: IUCN 2011. IUCN Red List Great Britain. Freshwater Crayfish, 17: 109–122. The IUCN Invertebrate Red Data Book. The Crayfish Pacifastacus leniusculus at Northerly of Threatened Species. version 2011.1., The Sala, O. E., Chapin, F. S., Armesto, J. J., Berlow, IUCN, Gland, Switzerland. 632 pp. ISBN Latitudes: A Search for the Distribution Limits. International Union for Conservation of Nature E., Bloomfield, J., Dirzo, R., Huber–Sanwald, 2–88032–602–X. Freshwater Crayfish, 16: 37–41. and Natural Resources. Gland, Switzerland. E., Huenneke, L. F., Jackson, R. B., Kinzig, Westman, K., Savolainen, R., & Julkunen, M., 2002. Holdich, D. M., 2002a. Part 3: Conclusions. In: D. M. Krebs, C. J., 1994. Ecology: the experimental A., Leemans, R., Lodge, D. M., Mooney, H. Replacement of the native crayfish Astacus Holdich (Ed.) Biology of Freshwater Crayfish. analysis of distribution and abundance. A., Oesterheld, M., Poff, N. L., Sykes, M. T., astacus by the introduced species Pacifastacus Blackwell Science, Oxford, England. pp 673– HarperCollins, New York, USA. 801 pp. ISBN Walker, B. H., Walker, M., & Wall, D. H., 2000. leniusculus in a small, enclosed Finnish lake: a 682. ISBN 0–632–05431–X. 0–06–500410–8. Global Biodiversity Scenarios for the Year 2100. 30–year study. Ecography, 25: 53–73. Holdich, D. M., 2002b. General Biology: May, R. M., 2010. Ecological science and Science, 287: 1770–1774. World Conservation Monitoring Centre, 1992. Background and Functional Morphology. In: tomorrow's world. Philosophical Transactions of Saunders, G. W., 1976. Decomposition in freshwater. Global Biodiversity: Status of the Earth's living D. M. Holdich (Ed.) Biology of Freshwater the Royal Society B, 365: 41–47. In: J. M. Anderson & A. Macfadyen (Eds), The resources. Chapman & Hall, London. 614 pp. Crayfish. Blackwell Science, Oxford, England. Momot, W. T., 1995. Redefining the role of crayfish Role of Terrestrial and Aquatic Organisms in Yamanaka, K., Kuwabara, R., & Shio, T., 1997. pp 3–29. ISBN 0–632–05431–X. in aquatic ecosystems. Reviews in Fisheries Decomposition Processes. Blackwell Scientific Larval development of a Japanese crayfish, Holdich, D. M., 2002c. Background and Functional Science, 3: 33–63. Publications, Oxford, England.: pp 341–373. Cambaroides japonicus (De Haan). Bulletin of Morphology. In: D. M. Holdich (Ed.) Biology of Moritz, C., 1994. Defining ‘Evolutionarily ISBN 0–632–00018–X. Marine Science, 61: 165–175. Freshwater Crayfish. Blackwell Science, Oxford. Significant Units’ for conservation. Trends in Sinclair, E. A., Madsen, A., Walsh, T., Nelson, J., & pp 3–29. Ecology and Evolution, 9: 373–375. Crandall, K. A., 2011. Cryptic genetic divergence Holdich, D. M., Reynolds, J. D., Souty–Grosset, Nakata, K., Hamano, T., Hayashi, K., & Kawai, in the giant Tasmanian freshwater crayfish Addresses: (KLD) Australian Rivers C., & Sibley, P. J., 2009. A review of the ever T., 2002. Lethal limits of high temperature for Astacopsis gouldi (Decapoda: Parastacidae): Institute and Griffith School of Environment, increasing threat to European crayfish from non– two crayfishes, the native species Cambaroides implications for conservation. Animal Gold Coast campus, Griffith University, indigenous crayfish species. Knowledge and japonicus and the alien species Pacifastacus Conservation, 14: 87–97. Queensland 4222, Australia, (JMF) Management of Aquatic Ecosystems, (394–395): leniusculus in Japan. Fisheries Science, 68: Söderbäck, B., 1991. Interspecific dominance relationship and aggressive interactions in the Environmental Futures Centre and Griffith 46 pp. 763–767. School of Environment, Gold Coast campus, Horwitz, P., 1990a. The conservation status of Nakata, K., & Goshima, S., 2003. Competition for freshwater crayfishes Astacus astacus (L.) and Australian freshwater crayfish with a provisional shelter of preferred sizes between the native Pacifastacus leniusculus (Dana). Canadian Griffith University, Queensland 4222, list of threatened species, habitats and potentially crayfish species Cambaroides japonicus and the Journal of Zoology, 69: 1321–1325. Australia; threatening processes. Report series No. 14. alien crayfish species Pacifastacus leniusculus Söderbäck, B., 1994. Reproductive interference Email addresses: (KLD) k.dawkins@ Australian National Parks and Wildlife Service, in Japan in relation to prior residence, sex between two co–occurring crayfish species, Astacus astacus L. and Pacifastacus leniusculus griffith.edu.au, (JMF) [email protected] Canberra, Australia. 121 pp. difference and body size. Journal of Crustacean – Horwitz, P., 1990b. The Translocation of Freshwater Biology, 23: 897–907. Crayfish in Australia: Potential Impact, the Need Nakata, K., Tanaka, A., & Goshima, S., 2004. for Control and Global Relevance. Biological Reproduction of the alien crayfish species Conservation, 54: 291–305. Pacifastacus leniusculus in Lake Shikaribetsu, Horwitz, P., & Knott, B., 1995. The distribution Hokkaido, Japan. Journal of Crustacean Biology, and spread of the yabby Cherax complex in 24: 496–501. Australia: speculations, hypotheses and the need Nakata, K., Tstutsumi, K., Kawai, T., & Goshima, for research. Freshwater Crayfish, 10: 81–91. S., 2006. Coexistence of two North American Horwitz, P., 2010. The Conservation Status of invasive crayfish species, Pacifastacus Freshwater Crayfish: The Basis for Concern, leniusculus (Dana, 1852) and Procambarus Listing and Recovery Processes, and Community clarkii (Girard, 1852) in Japan. Crustaceana, 78: Involvement. Freshwater Crayfish, 17: 1–12. 1389–1394. Imgrund, J., Groth, D., & Wetherall, J., 1997. Nguyen, T. T. T., Meewan, M., Ryan, S., & Austin, Genetic analysis of the freshwater crayfish C. M., 2002. Genetic diversity and translocation Cherax tenuimanus. Electrophoresis, 18: 1660– in the marron, Cherax tenuimanus (Smith): 1665. implications for management and conservation. IUCN, 2010. Guidelines for Using the IUCN Red Fisheries Management and Ecology, 9: 163–173. List Categories and Criteria: version 8. Prepared Nguyen, T. T. T., Austin, C. M., Meewan, M. by the Standards and Petitions Working Group M., Schultz, M. B., & Jerry, D. R., 2004. for the IUCN SSC Biodiversity Assessments Phylogeography of the freshwater crayfish Sub–Committee 2010. 60 pp. Cherax destructor Clark (Parastacidae) in inland IUCN, 2011. The IUCN Red List of Threatened Australia: historical fragmentation and recent Species (version 2011.1), The International range expansion. Biological Journal of the Union for Conservation of Nature and Natural Linnean Society, 83: 539–550. Resources. Gland, Switzerland. O'Brien, S. J., 1994. A role for molecular genetics Jussila, J., Ojala, K., & Mannonen, A., 2008. Noble in biological conservation. Proceedings of the Crayfish (Astacus astacus) Reintroduction National Academy of Sciences USA 91: 5748–