Assessing mate size in the red swamp crayfish Procambarus clarkii: effects of visual versus chemical stimuli

1. Mate assessment requires an efficient system of information exchange between the sexes and often relies on several forms of communication. Both sexes of the crayfish, Procambarus clarkii , select a mate on the basis of its body size, independently of chelar size/symmetry and dominance status, large size being a proxy for a high quality partner. Here, we explored the effects that visual and chemical stimuli, emitted alone or in combination, exert on the assessment of size of a potential mate.
2. We followed a binary choice test paradigm, in which two male or female 'targets', of either large or small size, were simultaneously presented to a female or a male 'chooser'. We recorded the first target visited, the total duration of the visits per target, and the behaviour of the chooser when approaching the target area.
3. Our results show that females require a combination of visual and chemical stimuli to select the larger male. A more complex pattern was found when males were the choosers. While visual and chemical cues together rendered them willing to mate with the larger female, their initial choice and the total time spent near her depended only on smell. In mate assessment by P. clarkii , visual and chemical information seem to act as 'non-redundant signals'.     

Mutual mate choice in crayfish: large body size is selected by both sexes, virginity by males only

Mate choice is often assumed to be a prerogative of females because of their putatively larger reproductive investment than males. However, recent evidence suggests that spermatogenesis is far from being limitless and that males show a high selectivity towards their mates, thus maximizing their reproductive success. We investigated mutual mate choice in the crayfish Procambarus clarkii through two experiments. The first experiment explored the effects of body size, chelar size and chelar symmetry and social status of a potential partner. In the second experiment, we asked whether this species can discriminate between partners of the same body size but with different mating status. We used a binary choice test paradigm, in which two 'targets' with opposing characteristics were simultaneously presented to a test animal, the 'chooser'. The results showed that P. clarkii males are more selective than expected. Similar to the other sex, they were significantly attracted by targets with large body sizes, but not by individuals with larger and symmetric chelae or with a dominant status. An inter-sexual difference was found in the second experiment, in which only males seemed to select virgin potential mates. The several adaptive explanations for these preferences, still under debate, are finally discussed.     

Habitat use and dispersal of the invasive crayfish Procambarus clarkii in ephemeral water bodies of Portugal

We used radio-telemetry to analyze habitat use and dispersal of the invasive crayfish Procambarus clarkii in the extreme environmental conditions of a temporary river in southern Portugal. The rationale of this study was that an understanding of the properties that make this species a successful invader can be of help for the conservation of freshwater biodiversity. The results showed that: (1) crayfish displaced at a similar rate (ranging 2.5-38 m d^-1) as other European populations; (2) movement was faster at night; (3) crayfish movement had no clear environmental correlates, except for water temperature; (4) peaks of more intense locomotion were intercalated with longer periods of slow or null speed; and (5) P. clarkii did not excavate burrows in the study habitat, but took refuge under boulders and most often occupied complex microhabitats (e.g., vegetated sections of the river). The conclusion was that, notwithstanding their extreme conditions, ephemeral water bodies in southern Europe are highly susceptible to invasion by P. clarkii.     

Eleven polymorphic microsatellites isolated from red swamp crayfish, Procambarus clarkii

Eleven microsatellites were isolated from a genomic DNA library enriched for CA- and GA-repeats, and were characterized in 48 individuals of red swamp crayfish Procambarus clarkii. All 11 microsatellites were polymorphic with an average of allele number of 6.6 per locus. Genotypic distributions of these 11 markers were found to be in conformance with the expectations of Hardy-Weinberg equilibrium. All 11 markers were unlinked. These markers are being used to study the invasion routine, genetic diversity and population structure of the species P. clarkii.     

Intensive trapping and increased fish predation cause massive population decline of an invasive crayfish

1. Invasive species frequently have adverse impacts on native communities and ecosystems. Management options are often limited. Our goal is to evaluate the effect of intensive trapping and fish predation on the population dynamics of an invasive crayfish. 2. From 2001 to 2005, we removed invasive rusty crayfish (Orconectes rusticus) by trapping in Sparkling Lake in northern WI. In addition, the Wisconsin Department of Natural Resources restricted harvest of fish species known to consume crayfish, thereby increasing predation on crayfish that are too small to trap. 3. After an initial increase, catch rates of rusty crayfish declined by approximately 95%, from 11 crayfish per trap per day in 2002 to 0.65 in 2004. The catch rate in 2005 remained low at 0.5 crayfish per trap. Females comprised nearly 50% of the catch from 2002 to 2004. Unlike rusty crayfish in Sparkling Lake, catch rates of O. rusticus and Orconectes propinquus in three nearby lakes increased or remained relatively constant over the 5‐year removal period. 4. We also examined the influence of habitat and temperature on crayfish catch rates. Catch rates were highest at water temperatures between 20 and 25 °C and on cobble, log or macrophyte habitats that may serve as refuge from fish predation. 5. Five summers of intensive trapping and fisheries management practices reduced abundances, but did not extirpate rusty crayfish in Sparkling Lake. To determine the potential of trapping as a management option for invasive crayfishes, these methods must be tested in other systems.     

Behavioural and growth differences between experienced and naïve populations of a native crayfish in the presence of invasive rusty crayfish

1. Species invasions are a leading threat to native species and ecosystems. How populations of native species respond to the presence of invasive species will ultimately determine their long-term persistence.
2. In this study, we capitalise on a unique opportunity to compare the behaviour and growth of naïve and experienced virile crayfish ( Orconectes virilis Hagen) populations in the presence of invasive rusty crayfish ( O. rusticus Girard ) . In behavioural trials, experienced O. virilis (coexisted with O. rusticus for >30 years) showed more aggressive behaviour than their naïve counterparts. Naïve O . virilis retreated from aggressive interactions with O. rusticus 38% more often than experienced O . virilis. Experienced O. virilis spent 39% more time occupying shelter than naïve O. virilis.
3. There were also differences in O. rusticus behaviour: O. rusticus spent 24% more time occupying shelter with naïve O. virilis relative to experienced O. virilis .
4. In field mesocosm experiments with O. rusticus , naïve O . virilis declined in body mass by 1% while experienced O. virilis' body mass increased by 6%, thus highlighting the potential population-level implications of the previously observed behavioural effects.
5. Our work demonstrates significant behaviour and growth differences between naïve and experienced O. virilis in the presence of invasive O. rusticus. Whether this difference is the result of phenotypic plasticity or evolution by natural selection remains to be determined. Either way, this area of inquiry has implications for managing native populations and species in an increasingly invaded world.     

Invasive species of crayfish use a broader range of predation-risk cues than native species

The responses of invasive and native species of crayfish to conspecific and heterospecific alarm odors were recorded in the laboratory. Individuals of the North American invasive Procambarus clarkii responded just as strongly to odors from crushed Austropotomobius pallipes as they did to crushed conspecifics. The North American invasive Orconectes limosus also responded as strongly to P. clarkii odor as to conspecific odor. The native Italian species A. pallipes responded more strongly to conspecific alarm than to heterospecific alarm from P. clarkii. The pattern of invasive species of crayfish using a broader range of danger signals than displaced native species appears to be robust.     

Invasive crayfish in Europe: the impact of Procambarus clarkii on the littoral community of a Mediterranean lake

1. Despite the growing number of non‐indigenous crayfish species introduced worldwide, little scientific attention has been given to their effects on the recipient communities. Here, we quantified the impact of adult red swamp crayfish (Procambarus clarkii) on key components of the littoral community of a mesotrophic lake in central Italy. We used two densities of crayfish plus a no‐crayfish control, and two complementary methods –in situ enclosures and a laboratory study. 2. Both experiments showed that P. clarkii strongly affects the community even at a low density (4 m⁻²), although a high crayfish density (8 m⁻²) showed a more pronounced impact in a shorter time. 3. In enclosures, P. clarkii quickly consumed the pond snail Haitia acuta, notwithstanding its anti‐predator behaviour, but not the mosquitofish Gambusia affinis. The biomass of the hydrophytes Nymphoides peltata and Potamogeton spp. was strongly reduced by crayfish grazing, coupled with their non‐consumptive plant clipping and uprooting, which were particularly intense in the case of Potamogeton crispus. In contrast, Utricularia australis was avoided by P. clarkii, even when other resources were exhausted. 4. The introduction of P. clarkii could reduce biodiversity and increase biotic homogenisation in a short time.     

Exploration of potential microbial control agents for the invasive crayfish,Orconectes virilis

The introduced crayfish, Orconectes virilis (Cambarinae; Hagen 1870), has become a serious invasive species in Arizona (USA), altering stream ecosystems and contributing to the decline of native species. But because it is native to the eastern US, and related crayfish including endangered species inhabit nearby states, introduction of a biological control agent presents a unique challenge. This represents a feasibility study, in which we explored bacteria, nematodes, and a virus as potential biological control agents for O. virilis while avoiding harm to native species. White Spot Syndrome Virus (WSSV) from shrimp was found to be highly pathogenic and readily passed by cannibalistic behavior but not by water transmission. Potential activity of WSSV against representative non-target arthropods that may be found in the same ecosystems was also explored.     

Disturbance and species displacement: different tolerances to stream drying and desiccation in a native and an invasive crayfish

1. Crayfish are among the most threatened taxa in the world and invasive crayfish are the primary cause of the decline of native crayfish. Most research has emphasised biotic interactions as the mechanism by which native crayfish are displaced by invasives, although crayfish occupy variable environments and the role of disturbance in facilitating crayfish invasion and displacement is understudied. 2. We compared tolerance to a disturbance, stream drying, in a native and invasive crayfish as a potential mechanism to explain their distribution. Our experiments and observations were conducted across scales, from laboratory environmental chambers to stream mesocosms to field sampling. We hypothesised that the invasive crayfish would be more tolerant of desiccation than the native, and that this physiological distinction between the two would be reflected in their distribution in relation to stream drying. 3. In the laboratory, the native crayfish Orconectes eupunctus was less tolerant of desiccation than the invasive Orconectes neglectus chaenodactylus, with all native crayfish dying within 2 days without water, while some of the invasive crayfish survived for nearly 2 weeks. Under simulated stream drying in mesocosms, only the native O. eupunctus survived less well than in a control. Field sampling demonstrated a significant negative relationship between O. eupunctus density and low summer flows, while O. neglectus density was positively associated with low summer flows. The greater resistance of O. neglectus to drying could, through priority effects, inhibit recolonisation by O. eupunctus once flow resumes. 4. Abiotic disturbances are potentially important to the displacement of native by invasive crayfish. Disturbance mediated displacement of aquatic species provides both an opportunity to conserve native species by maintaining or restoring habitat and disturbance regimes and is also a challenge due to increasing human water demand, flow regime alteration and global climate change.     

Invasion of the shelter snatchers: behavioural plasticity in invasive red swamp crayfish,Procambarus clarkii

1. In many freshwater systems, competition for shelter plays an important role in determining the persistence of both native and alien species. The red swamp crayfish, Procambarus clarkii, is currently invading the native habitat of the signal crayfish, Pacifastacus leniusculus, in southern Oregon, and interspecific competition for shelter may be driving the species replacement in this region. 2. We designed a 2 × 3 factorial mesocosm experiment, with shelter density and species combination as factors, to investigate shelter occupancy and resource competition. Contrary to our predictions, the two crayfish species are equal competitors for shelter. Further, the invasive P. clarkii modified its shelter occupancy behaviour in the presence of the native P. leniusculus and has broader microhabitat preferences. 3. Specifically, we found that P. clarkii alters shelter occupancy and space use patterns when the two species occurred together, such that shelter use was identical between P. clarkii and P. leniusculus in mixed‐species treatments. In such treatments, both species increased their use of shelters when shelter density increased. When P. clarkii was alone, however, individuals did not alter shelter use as a function of shelter density, whereas P. leniusculus exhibited similar density‐dependent behaviour in both mixed‐ and single‐species treatments. 4. In a complementary field survey, we employed an ‘epicentre‐based’ design to sample two field sites. We observed patterns of microhabitat use and breadth for each species similar to those in our mesocosm experiment: the invasive P. clarkii was more abundant across different habitats and used a broader range of microhabitats than the native P. leniusculus. As such, we found that P. clarkii was more abundant across both field sites than the P. leniusculus, occupying microhabitats within and beyond the preferred range of P. leniusculus. Both field sites were affected by urban development and agriculture. 5. The use of microhabitats by both species was similar in the laboratory and the field. This study confirms that P. clarkii individuals can, and do, successfully occupy microhabitats preferred by P. leniusculus in the Willamette Valley. The results from our study may be relevant to other freshwater systems inhabited by P. clarkii and contribute to the understanding of ‘niche opportunity’, a concept which defines the environmental conditions that promote biological invasions.     

Crayfish invading Europe: the case study of Procambarus clarkii

The red swamp crayfish, Procambarus clarkii, native to northeastern Mexico and southcentral USA, is today the dominant macroinvertebrate in several European countries. While the first introduction of this species into Spain is well-documented, little is known about its pathways of invasion and the reason for its rapid spread in several European countries. Study of the biology of the species has revealed a number of properties that makes this crayfish a successful invader. Procambarus clarkii exhibits properties characteristic of an r-selected species, including early maturity at small body size, rapid growth rates, large numbers of offspring at a given parental size, and relatively short life spans. It is also plastic in its life cycle, able to disperse widely in the habitat and to tolerate environmental extremes. It displays generalist and opportunistic feeding habits, consuming macrophytes and preying on amphibians. Procambarus clarkii can also replace indigenous crayfish by a combination of mechanisms, including competitive exclusion and transmission of the fungus-like Aphanomyces astaci, responsible for the crayfish plague. Finally, this species shows a wide behavioral flexibility when coping with new types of predators. The results of these studies, combined with the increasing information available in the scientific literature on this and other crayfish species, will help us understand invasions in this taxon and make predictions about the identity of future crayfish invaders.     

Release of Sterile Mediterranean Fruit Flies, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae), Using an Automated Ground Release Vehicle

A trial was conducted in Tampa, Florida to test an automated ground release machine as an alternative to aerial methods for release of sterile Mediterranean fruit flies, Ceratitis capitata (Wiedemann). Twenty-four Jackson traps baited with trimedlure at three release sites yielded lower than expected numbers of ground released Medflies. Recaptures per trap ranged from 2.8 (±0.7 S.E.) to 6.8 (±1.7 S.E.). Average recovery of sterile flies at each site for all releases combined (144 observations) was similar, 3.4 (±0.5 S.E.), 4.4 (±0.5 S.E.), 4.5 (±0.5 S.E.) for sites 1, 2 and 3 respectively. There was a significant reduction in flight ability from 67.5 (±2.1 S.E.) to 53.6 (±2.9 S.E.)%, from the time of pre-chilling to when immobilized flies were released after 2.5 h in the release container. Further tests are needed to determine what mortality factors may influence recovery and how the automated ground release machine compares to the aerial release method.     

The invasive red swamp crayfish as a predictor of Eurasian bittern density in the Camargue, France

Few data exist on the relationships between food levels and breeding density of the Eurasian bittern Botaurus stellaris , a vulnerable species of high-priority concern in Europe. Concurrent data were obtained on male bittern density and relative food abundance over a 3-year period in two wetlands totalling 2500 ha of Mediterranean reed marsh enclosing 25% of the French bittern population. Food abundance was estimated by sampling up to 25 hydrological units using a beach seine in early June of 2002, 2003 and 2004. The density of booming males in each hydrological unit was obtained by point counts and acoustic triangulation in May of the same years. The impact of food abundance on male bittern density was assessed by general regression models using a forward stepwise procedure with mosquitofish Gambusia affinis , carp Cyprinus carpio , other fish species, amphibians, red swamp crayfish Procambarus clarkii and other invertebrates as prey groups. Of these, only to crayfish abundance was bittern density related, contributing to 45% of the variance observed. When the impact of water level was taken into account, the relative abundance of crayfish explained 56% of the inter-annual differences in bittern density. Because crayfish are rich in calcium and well adapted to fluctuating hydroperiods alternating with drought intervals, they provide a good food source for the bitterns throughout the breeding season at the study sites. The loss of diversity and degradation reported from macrophyte-dominated marshes following crayfish invasion does not seem to apply to reed-dominated wetlands. It is further suggested that the recent increase in bittern numbers in the Camargue, while other French populations were decreasing, could in part be related to red swamp crayfish abundance.     

Absence of the crayfish plague pathogen (Aphanomyces astaci) facilitates coexistence of European and American crayfish in central Europe

1. Most European crayfish species are strongly threatened, mainly as a result of the introduced pathogen, Aphanomyces astaci, transmitted by invasive North American crayfish. Long‐term coexistence of American and European crayfish species is therefore regarded as almost impossible, even though some coexisting populations have been observed. 2. In this study, crayfish were collected from presently coexisting populations of the introduced spiny‐cheek crayfish (Orconectes limosus) and the native noble crayfish (Astacus astacus) from nine standing waters in central Europe. Our aim was to resolve whether the coexistence resulted from reduced virulence in local strains of A. astaci, increased immunity in the native crayfish or an absence of the pathogen in these populations. We used highly sensitive A. astaci‐specific real‐time PCR to evaluate the crayfish latent carrier status, combined with transmission experiments to further validate the molecular results. 3. From the total of 523 crayfish tested (490 spiny‐cheek crayfish, 33 noble crayfish), none positive for A. astaci was detected. Transmission experiments confirmed these results: No abnormal mortality or behavioural changes were seen in noble crayfish kept together with American crayfish from the coexisting populations. If we assume a very low prevalence of A. astaci of 10% in a carrier population, there is a 98% probability of disease being absent in five of the nine coexisting populations tested. Hence, a consistent absence, or an extremely low prevalence, of A. astaci seems to allow the coexistence of European and American crayfish in these central European populations. 4. The results are important for native crayfish conservation and management and demonstrate that disease transmission risk may vary substantially between the different populations of spiny‐cheek crayfish in central Europe.     

Detoxification enzymes of freshwater crayfish Procambarus clarkii fed a diet enriched in selenium: Preliminary results

This study aimed at investigating the extent of accumulation of selenium in the hepatopancreas of the crayfish Procambarus clarkii and its effects on the enzymatic activities of catalase, glutathione peroxidase, and glutathione reductase. Crayfish were fed for 15 days either a diet enriched in Se, ED (1.21 mg kg^-1) or a standard diet, SD (0.30 mg kg^-1). Results showed that Se accumulation is higher in the sample subject to ED rather than to SD. A sexual difference was found for the response of the analyzed enzymes in the ED sample only. Catalase activity and glutathione peroxidase activity decreased in females and males, respectively, whereas glutathione reductase activity showed significant reductions in both sexes. These preliminary data might provide information about prooxidant effects of Se on P. clarkii when fed a Se enriched diet.     

Movement and dispersal of the invasive signal crayfish Pacifastacus leniusculus in upland rivers

1. The American signal crayfish Pacifastacus leniusculus, an invasive species widely introduced throughout Europe, is a major threat to native European crayfish species and is causing increasing concern because of its wide impact on aquatic ecosystems. 2. Whilst various control and management methods have been proposed, very little is known about the factors influencing dispersal and movements of signal crayfish. 3. Sixty‐four adult signal crayfish (carapace length 31.9–63.8 mm) were radiotagged in upland rivers in northern England, during four periods. Tracking was carried out at two sites, a low‐density establishing population and a high‐density established population. Tracking was carried out at both sites concurrently during midsummer (June to August 2002), during late summer (August to September 2001) at the low‐density population site and during autumn to winter (October to February 2000/01) at the high‐density population site. 4. Maximum movement occurred during midsummer. Temperature appeared to be a major factor influencing the timing and extent of movements between tracking periods. 5. The frequency distribution of the maximum distance moved upstream and downstream by radiotagged crayfish showed an inverse power relationship. The median maximal upstream and downstream distances moved were 13.5 m (range 0–283 m) and 15 m (range 0–417 m), respectively. There was a significant difference between the distributions of upstream and downstream ranges, with greater distances moved downstream. 6. All downstream movements made by crayfish appeared to be active movements and not the result of passive movement during periods of high discharge. There was no apparent influence of size, sex or density on the amount of movement recorded. 7. The study provides important information on the spatial and temporal behaviour of introduced crayfish in upland lotic systems. In contrast to lowland rivers, our results suggest that flow or gradient may influence the invasive potential of signal crayfish in an upstream direction in upland rivers.     

From introduction to fishery dominance: the initial impacts of the invasive carp Cyprinus carpio in Lake Naivasha, Kenya, 1999 to 2006

Following the accidental introduction of the carp Cyprinus carpio into Lake Naivasha during 1999, a sustainable population became rapidly established and in early 2004 became the principal species exploited in the commercial fishery. Over 9000 kg of carp were harvested from the lake between October 2005 and 2006, when fish were captured between fork lengths ( L _F) 200 and 800 mm (>8 kg). Diet of carp <100 mm L _F was dominated by zooplankton, >100 mm L _F there was a shift to benthic macro-invertebrates, with these carp feeding principally upon food resources previously unexploited by the fish community. Contrary to predictions and despite the increasing carp abundance, there has been macrophyte regeneration in littoral areas since 2004. There have been substantial increases in areal cover, with coverage in 2006 at levels not observed since the late 1980s, and significant increases in species richness. Possible reasons for this, and the significance of this carp introduction, are discussed.     

Invasive American crayfish <Emphasis Type="Italic">Pacifastacus leniusculus</Emphasis> (Decapoda: Astacidae) in the Morava River (Slovakia)

The signal crayfish, Pacifastacus leniusculus (Dana, 1852), was recorded in the Slovak-Austrian stretch of the Morava (March) River in the Záhorie region in August 2006. This is the first confirmed record of a non-indigenous crayfish species of American origin in Slovakia. Most likely, signal crayfish were intentionally released to the river by Austrian fishermen, or invaded from adjacent water bodies in Austria. As this species has a substantial invasive capability and is a potential vector of the crayfish plague pathogen, it represents a threat to the native crayfish species and its spread should be monitored. Its invasion upstream to the territory of the Czech Republic is not unlikely in near future.