Do introduced crayfish affect benthic fish in stony littoral habitats of large boreal lakes?

Invasive crayfish are spreading rapidly across Europe, where they are replacing the native crayfish species and impacting negatively on some other biota. Freshwater crayfish and many benthic fishes share similar habitat and food requirements and hence potentially compete for resources. In this study, we investigated impacts of the introduced signal crayfish (Pacifastacus leniusculus) on fish in stony littoral habitats of two large boreal lakes. We compared the littoral fish community composition and the densities of two common benthic fish species between sites with and without crayfish. To evaluate whether signal crayfish share the same food resources as benthic littoral fish or change their feeding habits, we used mixing models and trophic niche estimates based on analyses of stable isotopes of carbon and nitrogen. Both the community composition of littoral fish and the densities of benthic fish species were similar at sites with and without signal crayfish. Even though stable isotope signatures indicated strong dietary overlap between crayfish and benthic fish, the use of food sources and trophic niche widths of fish were not noticeably different between crayfish sites and non-crayfish sites. Our results suggest that, at current densities, the non-native signal crayfish does not have significant impacts on benthic fish in the stony littoral habitats of large boreal lakes.     

Are fish populations in temperate streams affected by crayfish? – A field survey and prospects

Synopsis Fish populations may be affected by predation and competition from various types of organisms, among which crayfish have been suggested as important actors. We here present results from stream surveys, suggesting that neither native noble, Astacus astacus, nor introduced signal crayfish, Pacifastacus leniusculus, necessarily affect fish population densities in temperate stream communities. Comparisons of fish densities within stream sites between years with absence and presence of crayfish showed no effect of either crayfish species. A further analysis of changes in fish densities between periods without and with crayfish in low, intermediate and high densities revealed that crayfish density did neither have an effect on fish densities. Our study is one of exceptionally few that consider the above aspects in long-term perspectives in natural systems, and we discuss that previously reported divergent results of crayfish effects on fish may be highly dependent on specific species and methods used, and that the effects of crayfish on fish populations deserve further attention to enable reliable predictions of community processes in streams.     

Non‐lethal predator effects on the performance of a native and an exotic crayfish species

1. I tested the hypothesis that the potential for non‐lethal effects of predators are more important for overall performance of the fast‐growing exotic signal crayfish (Pacifastacus leniusculus Dana) than for the slower growing native noble crayfish (Astacus astacus L.). I further tested if omnivorous crayfish switched to feed on less risky food sources in the presence of predators, a behaviour that could reduce the feeding costs associated with predator avoidance. 2. In a 2 month long outdoor pool experiment, I measured behaviour, survival, cheliped loss, growth, and food consumption in juvenile noble or signal crayfish in pools with either a caged predatory dragonfly larvae (Aeshna sp.), a planktivorous fish that do not feed on crayfish (sunbleak, Leucaspius delineatus Heckel), or predator‐free controls. Crayfish had access to multiple food sources: live zooplankton, detritus and periphyton. Frozen chironomid larvae were also supplied ad libitum outside crayfish refuges, simulating food in a risky habitat. 3. Crayfish were mainly active during hours of darkness, with signal crayfish spending significantly more time outside refuges than noble crayfish. The proportion of crayfish outside refuges varied between crayfish species, time and predator treatment, with signal crayfish spending more time in refuges at night in the presence of fish. 4. Survival in noble crayfish was higher than in signal crayfish, and signal crayfish had a higher frequency of lost chelipeds, indicating a high level of intraspecific interactions. Crayfish survival was not affected by the presence of predators. 5. Gut‐contents analysis and stable isotope values of carbon (δ¹³C) and nitrogen (δ¹⁵N) indicated that the two crayfish species had similar food preferences, and that crayfish received most of their energy from feeding on invertebrates (e.g. chironomid larvae), although detritus was the most frequent food item in their guts. Signal crayfish guts were more full than those of noble crayfish, but signal crayfish in pools with fish contained significantly less food and fewer had consumed chironomids compared with predator‐free controls. Length increase of signal crayfish (35%) was significantly higher than of noble crayfish (20%), but signal crayfish in pools with fish grew less than in control pools. 6. This short‐term study indicates that fish species that do not pose a lethal threat to an organism may indirectly cause reductions in growth by affecting behaviour and feeding. This may occur even though prey are omnivorous and have access to and consume multiple food sources. These non‐lethal effects of predators are expected to be particularly important in exotic crayfish species that show a general response to fish, have high individual growth rates, and when their feeding on the most profitable food source is reduced.     

Functional significance of crayfish in stream food webs: roles of omnivory,substrate heterogeneity and sex

Heterogeneity of species interactions in food webs can result from characteristics of substrates as well as attributes of top consumers. We performed a streamside channel experiment to evaluate the impact of crayfish on lower trophic levels in detritus‐based (leaf packs) and algal‐based food webs (hard‐bottoms). After 43 days, both male and female crayfish had dramatically promoted leaf decomposition, with males processing material at a faster rate. However, the difference in leaf processing rates was not related to a greater level of male activity. Despite the sex‐related difference in residual leaf dry mass, densities of invertebrates in leaf packs were similarly low in the presence of crayfish of either sex, due to resource consumption, physical dislodgment (bioturbation) and/or predation. No trophic cascade was evident in the leaf pack assemblage. In the hard‐bottom assemblage, the results confirmed circumstantial field evidence that crayfish reduce predatory Tanypodinae and indirectly increase collector‐gatherer Chironominae following the prediction of a trophic cascade. However, no other taxa were indirectly facilitated, because of strong direct effects of crayfish on algal abundance (through direct consumption and bioturbation). Overall, impacts of crayfish on lower trophic levels were more pronounced in the structurally complex, detritus‐based assemblages than in its hard‐bottom, algal‐based counterpart. This conflicts with the expectation that net predation effects should be weaker where structural complexity is greater but is mainly a consequence of the profound engineering effects of crayfish in reducing colonisable substrate when they shred and disturb detrital material. Effects of crayfish may therefore propagate differently and with varying strength depending on substrate. Moreover, engineering activities and predation by crayfish appear to have been of overwhelming significance with subtle sex differences in leaf processing rates failing to lead to differences in invertebrate densities.     

Intensive removal of signal crayfish (Pacifastacus leniusculus) from rivers increases numbers and taxon richness of macroinvertebrate species

Invasive species are a major cause of species extinction in freshwater ecosystems, and crayfish species are particularly pervasive. The invasive American signal crayfish Pacifastacus leniusculus has impacts over a range of trophic levels, but particularly on benthic aquatic macroinvertebrates. Our study examined the effect on the macroinvertebrate community of removal trapping of signal crayfish from UK rivers. Crayfish were intensively trapped and removed from two tributaries of the River Thames to test the hypothesis that lowering signal crayfish densities would result in increases in macroinvertebrate numbers and taxon richness. We removed 6181 crayfish over four sessions, resulting in crayfish densities that decreased toward the center of the removal sections. Conversely in control sections (where crayfish were trapped and returned), crayfish density increased toward the center of the section. Macroinvertebrate numbers and taxon richness were inversely correlated with crayfish densities. Multivariate analysis of the abundance of each taxon yielded similar results and indicated that crayfish removals had positive impacts on macroinvertebrate numbers and taxon richness but did not alter the composition of the wider macroinvertebrate community. Synthesis and applications: Our results demonstrate that non‐eradication‐oriented crayfish removal programmes may lead to increases in the total number of macroinvertebrates living in the benthos. This represents the first evidence that removing signal crayfish from riparian systems, at intensities feasible during control attempts or commercial crayfishing, may be beneficial for a range of sympatric aquatic macroinvertebrates.     

Nonconsumptive predator effects modify crayfish‐induced bioturbation as mediated by limb loss: Field and mesocosm experiments

1. We addressed the implications of limb loss and regeneration for multispecies interactions and their impacts on ecosystem engineering in freshwater stream environments.
2. We included regenerative and nonregenerative crayfish as well as fish predators in a 2 × 2 factorial design to assess the effects on water turbidity of interactions between crayfish ecosystem engineers differing in regenerative status and their fish predators.
3. We demonstrated that crayfish limb loss and predation risks lead to more turbidity in field and mesocosm conditions. Moreover, ongoing regeneration of crayfish increased turbidity, while fish presence seemed to hinder crayfish turbidity‐inducing behaviors (such as tail‐flipping and burrowing) in the mesocosm experiment.
4. We confirmed that greater numbers of crayfish produce a greater amount of turbidity in situ in streams.
5. Although mechanical burrowing crayfish capacities may depend on crayfish burrowing classification (primary, secondary, or tertiary), our work emphasizes the implication for turbidity levels of crayfish autotomy in freshwater streams.

     

Exotic crayfish in a brown water stream: effects on juvenile trout, invertebrates and algae

SUMMARY 1. The impact of the introduced omnivorous signal crayfish (Pacifastacus leniusculus) on trout fry, macroinvertebrates and algae was evaluated in a brown water stream in southern Sweden using in situ enclosures. We also examined the gut content of all surviving crayfish in the enclosures. Two crayfish densities in addition to a control without crayfish were used in replicate enclosures (1.26 m²) in a 1‐month experiment. Additionally, 20 trout fry (Salmo trutta) were stocked in each enclosure to assess the effects of crayfish on trout survival and growth. 2. Detritus was the most common food item in crayfish guts. Animal fragments were also frequent while algae and macrophytes were scarcer. Crayfish exuviae were found in crayfish guts, but the frequency of cannibalism was low. 3. Trout survival in enclosures was positively related to water velocity but was unaffected by crayfish. 4. Total invertebrate biomass and taxon richness were lower in crayfish treatments. The biomass of all predatory invertebrate taxa was reduced but only three of six non‐predatory taxa were reduced in the crayfish treatments. 5. Epiphytic algal biomass (measured as chlorophyll a, on plastic strips) was not related to crayfish density, whereas the biomass of epilithic algae (measured as chlorophyll a) was enhanced by high water velocity and high crayfish density. The latter was possibly mediated via improved light and nutrient conditions, as active crayfish re‐suspend and/or remove detritus and senescent algal cells during periods of low water velocity. 6. We conclude that the introduced signal crayfish may affect stream communities directly and indirectly. Invaded communities will have reduced macroinvertebrate taxon richness and the signal crayfish will replace vulnerable invertebrate predators such as leeches. In streams that transport large amounts of sediment or organic matter, a high density of crayfish is likely to enhance benthic algal production through physical activity rather than via trophic effects.     

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.     

Competition for shelter among over‐wintering signal crayfish and juvenile Atlantic salmon

Three separate effects on refuge use by signal crayfish Pacifastacus leniusculus and Atlantic salmon Salmo salar were examined: (1) the effect on Atlantic salmon of an addition of signal crayfish (doubling the total number of animals), (2) the effect on signal crayfish of an addition of Atlantic salmon and (3) intraspecific compared with interspecific competition, compared by holding total density of animals constant and varying the proportion of signal crayfish and Atlantic salmon in trials. Observations were made during winter, when both species are nocturnal. The proportion of Atlantic salmon sheltering was significantly lower in the presence than in the absence of signal crayfish when the interspecific treatment (Atlantic salmon plus signal crayfish) effected a doubling in density compared to the intraspecific treatment (Atlantic salmon alone). The proportion of signal crayfish sheltering was independent of the presence of Atlantic salmon. When total density was constant, the proportion of Atlantic salmon sheltering was significantly higher in intraspecific (52·8%) than interspecific trials (27·3%). Atlantic salmon out of shelter during the day in winter are believed to be very vulnerable to predators and the capacity for fish to share shelters with one another is known to be very low. Therefore, competition from crayfish for winter shelters may lead to detrimental effects on Atlantic salmon populations.     

The effect of manual removal on movement distances in populations of signal crayfish (Pacifastacus leniusculus)

1. Crayfish are amongst the most frequently introduced non‐native aquatic organisms, with well‐documented negative effects on a large number of freshwater taxa. Many crayfish‐control strategies make use of manual removal by trapping, a method known preferentially to remove the largest individuals, leaving the juvenile population almost entirely untrapped. 2. Removal by trapping may be used in an attempt to delay colonisation of new stretches by invasive crayfish. It is, however, unclear what effects trapping may have on movement distances of crayfish in wild populations. We examine the impacts of removal by trapping on the movements of American signal crayfish in two UK rivers. 3. We studied four 100 m stretches of two rivers, the Evenlode and Thame, comprising two removal and two non‐removal stretches. Each river supported both treatments. Half of the crayfish captured from the removal sections were removed and humanely destroyed by freezing, and half were marked with their trap location and released there. All crayfish captured from the non‐removal sections were marked and returned at the point of capture. 4. Mean movement distances were smaller in the removal stretches than the non‐removal stretches, both within capture sessions (10.8 and 16.0 m, respectively) and between sessions (14.5 and 24.6 m, respectively), suggesting that removal trapping resulted in the remaining crayfish making smaller movements. Larger crayfish under both treatments made substantially larger movements than those with smaller carapace lengths, both within capture sessions (range 7.6–19.6 m) and between range capture sessions (range 8.9–32.6 m). 5. The results of this study are consistent with expectations if removal by trapping lowered population densities, which we speculate may have affected movement distances directly or indirectly through increasing the availability of food and shelter. 6. This study suggests that trapping at the margins of a population may be sufficient to delay colonisation of new stretches by: (i) maintaining low densities and therefore reducing movements, and (ii) preferentially reducing the population of large individuals, which make the largest movements. However, it remains unlikely that any trapping programme can entirely prevent emigration /dispersal, and therefore colonisation, by signal crayfish.     

Threat of non-native crayfish introductions into Turkey: global lessons

Introductions of crayfish species from their home range to new environments have been carried out in many parts of the world. The most important introduced crayfish species are Procambarus clarkii, Pacifastacus leniusculus, Cherax destructor, C. quadricarinatus, Orconectes limosus, O. rusticus and Astacus leptodactylus. The environmental impact of crayfish introductions can be positive, negative or neutral. However, native crayfish populations in Europe have been negatively affected by introductions of non-indigenous crayfish species from America. Negative effects of non-native crayfish introductions included displacement of native crayfish species, transfer of disease (crayfish plague), consumption of fish eggs, reduction of fish stocks, consumption of large amounts of macrophytes, indirect and direct effects on other invertebrates and upsetting production in rice fields. As a result of non-native crayfish introductions, the natural harvest and crayfish industry in Europe have been severely affected. Large quantities of Turkish A. leptodactylus were harvested (approximately 7,000 tonnes annually) and exported to Europe before the crayfish plague was observed in these populations. The total harvest of A. leptodactylus in Turkey reduced dramatically to 320 in 1991 after the plague. Therefore, although Turkey currently has no known non-native crayfish species, there is a threat of non-native crayfish introduction in order to increase crayfish productions and subsequent harvest. The North American spiny-cheek crayfish, O. limosus, has been spreading quickly down the River Danube and could soon reach neighboring countries including Turkey. The North American signal crayfish, P. leniusculus is known from Greece and could be a threat to native stocks if it is introduced into Turkey for aquaculture. Additional threats may come from the release of other North American species, which are widely available through the aquarium trade. We conclude that the spread of non-native crayfish introductions throughout Turkey will increase local problems, because introductions of non-native crayfish in many parts of the world have been known to have caused important reductions in population density and numbers of native crayfish species. Furthermore, freshwater ecosystems may be altered by such introductions and the economic viability of native crayfish species fisheries could be severely reduced in Turkey.     

Tuning in to multiple predators: conflicting demands for shell morphology in a freshwater snail

1. We examined the response to chemical cues from fish and crayfish, two predators with contrasting feeding modes, and their single and combined effect on shell morphology in the freshwater snail Radix balthica. 2. Snails were subjected to four treatments: tench (Tinca tinca), signal crayfish (Pacifastacus leniusculus), a combination of tench and signal crayfish and no predators (control). Shell shape, crushing resistance and shell thickness were quantified. We also analysed whether shape or shell thickness contributes most to crushing resistance. 3. Chemical cues from the fish induced a rounder shell shape in R. balthica, a thicker shell and a higher crushing resistance, whereas crayfish chemical cues had no effect on shell morphology, shell thickness or crushing resistance. Shell shape contributed more to crushing resistance than shell thickness. 4. The combined predator treatment showed an intermediate response between the fish and crayfish treatments. Shell roundness was reduced compared with the fish treatment, but the reduced crushing resistance that comes with a less rounded shell was compensated by an increased investment in extra shell material, exceeding that of the fish treatment. 5. Our study extends previous studies of multipredator effects on phenotypically plastic freshwater snails by showing that the snails are able to fine‐tune different elements of morphology to counter predator‐specific foraging modes.     

Effects of Predation Risk on Habitat Selection by Water Column Fish, Benthic Fish and Crayfish in Stream Pools

Predation risk can affect habitat selection by water column stream fish and crayfish, but little is known regarding effects of predation risk on habitat selection by benthic fish or assemblages of fish and crayfish. I used comparative studies and manipulative field experiments to determine whether, (1) habitat selection by stream fish and crayfish is affected by predation risk, and (2) benthic fish, water column fish, and crayfish differ in their habitat selection and response to predation risk. Snorkeling was used to observe fish and crayfish in, (1) unmanipulated stream pools with and without large smallmouth bass predators (Micropterus dolomieui >200 mm total length, TL) and (2) manipulated stream pools before and after addition of a single large smallmouth bass, to determine if prey size and presence of large fish predators affected habitat selection. Observations of microhabitat use were compared with microhabitat availability to determine microhabitat selection. Small fish (60–100 mm TL, except darters that were 30–100 mm TL) and crayfish (40–100 mm rostrum to telson length; TL) had significantly reduced densities in pools with large bass, whereas densities of large fish and crayfish (> 100 mm TL) did not differ significantly between pools with and without large bass. Small orangethroat darters (Etheostoma spectabile), northern crayfish (Orconectes virilis), and creek chubs (Semotilus atromaculatus) showed significantly greater densities in pools without large bass. The presence of large smallmouth bass did not significantly affect depths selected by fish and crayfish, except minnows, which were found significantly more often at medium depths when bass were present. Small minnows and large and small crayfish showed the greatest response to additions of bass to stream pools by moving away from bass locations and into shallow water. Small darters and sunfish showed an intermediate response, whereas large minnows showed no significant response to bass additions. Response to predation risk was dependent on prey size and species, with preferred prey, crayfish and small minnows, showing the greatest response. Small benthic fish, such as darters, are intermediate between small water column fish and crayfish and large water column fish in their risk of predation from large smallmouth bass.     

Effects of an invasive crayfish on trophic relationships in north‐temperate lake food webs

1. The introduction of invasive species is one of the main threats to global biodiversity, ecosystem structure and ecosystem processes. In freshwaters, invasive crayfish alter macroinvertebrate community structure and destroy macrophyte beds. There is limited knowledge on how such invasive species‐driven changes affect consumers at higher trophic levels. 2. In this study, we explore how the invasive rusty crayfish Orconectes rusticus, a benthic omnivore, affects benthic macroinvertebrates, as well as the broader consequences for ecosystem‐level trophic flows in terms of fish benthivory and trophic position (TP). We expected crayfish to decrease abundance of benthic macroinvertebrates, making most fish species less reliant on benthic resources. We expected crayfish specialists (e.g. Lepomis sp. and Micropterus sp.) to increase their benthic dependence. 3. In 10 northern Wisconsin lakes, we measured rusty crayfish relative abundance (catch per unit effort, CPUE), macroinvertebrate abundance, and C and N stable isotope ratios of 11 littoral fish species. We used stable isotope data and mixing models to characterise the trophic pathways supporting each fish species, and related trophic structure to crayfish relative abundance, fish body size and abiotic predictors using hierarchical Bayesian models. 4. Benthic invertebrate abundance was negatively correlated with rusty crayfish relative abundance. Fish benthivory increased with crayfish CPUE for all 11 fish species; posterior probabilities of a positive effect were >95%. TP also increased slightly with crayfish CPUE for some species, particularly smallmouth bass, largemouth bass, rock bass and Johnny darter. Moreover, both fish body size and lake abiotic variables explained variation in TP, while their effects on benthivory were small. 5. Rusty crayfish abundance explained relatively little of the overall variation in fish benthivory and TP. Although rusty crayfish appear to have strong effects on abundances of benthic macroinvertebrates, energy flow pathways and trophic niches of lentic fishes were not strongly influenced by invasive rusty crayfish.     

An invasive crayfish affects egg survival and the potential recovery of an endangered population of Arctic charr

1. Many fish stocks have declined, because of overharvesting, habitat destruction and introduced species. Despite efforts to rehabilitate some of these stocks, not all are responding or are recovering only slowly. 2. In freshwater systems, introduced crayfish are often problematic, and it has been suggested that their egg predation could reduce recruitment in depleted stocks of native fish. 3. Here, we report the results of a field experiment, using experimental cages, on the extent of predation on eggs of great Arctic charr (Salvelinus umbla) in Lake Vättern, Europe’s fifth largest lake. Here, the great Arctic charr has declined dramatically and is listed as critically endangered. 4. We were able to partition the total loss rate of eggs into background mortality, predation by introduced signal crayfish (Pacifastacus leniusculus) and predation by native fish. The mortality rate of charr eggs because of crayfish was estimated at more than five times that because of native fish. Of the total loss of eggs, 80% is believed to be caused by crayfish and 14% by fish, with 6% being natural background mortality. 5. In a worst case scenario, our data infer that only 25% of the original number of eggs would survive, compared with 75% in the absence of crayfish. This could impair recovery of the stock of the endangered great Arctic charr in Lake Vättern. 6. Contrary to earlier claims that crayfish predation on eggs of great Arctic charr is insignificant, our results indicate that crayfish predation may exceed fish predation and suggest that the abundance of signal crayfish on the spawning sites of great Arctic charr should be managed.     

An invasive species may be better than none: invasive signal and native noble crayfish have similar community effects

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Growth and population size of crayfish in headwater streams: individual- and higher-level consequences of acidification

1. Environmental stress may have indirect positive effects on population size through modification of food‐web interactions, despite having negative effects on individuals. Here we evaluate the individual‐ and population‐level effects of acidification on crayfish (Cambarus bartonii) in headwater streams of the Allegheny Plateau (PA, U.S.A.) with field experiments and survey data. Median baseflow pH of 24 study reaches in nine streams varied from 4.4 to 7.4, with substantial variation found both among and within streams. 2. Two bioassays were conducted to evaluate the relationship between stream pH and crayfish growth rates. Growth rates were always higher in circumneutral reaches than in acidic reaches. Crayfish originating in acidic water grew less when transplanted into neutral water than did crayfish originating in neutral water, providing some evidence for a cost of acclimation to acidity. 3. Stream surveys showed that fish were less abundant and crayfish more abundant in acidified streams than in circumneutral streams. Crayfish density was sixfold higher in reaches with the lowest pH relative to circumneutral reaches. Large crayfish made up a higher proportion of crayfish populations at sites with high fish biomass, consistent with the hypothesis that fish predation on small individuals may be limiting crayfish population size at these sites. 4. Although individual crayfish suffered lower growth in acidified streams, increased acidity appeared to cause an increase in crayfish population size and shifts in size structure, possibly by relieving predation pressure by fish.     

Coupling long-term studies with meta-analysis to investigate impacts of non-native crayfish on zoobenthic communities

1. Biological invasions are widely recognised as a significant component of human‐caused environmental change and a primary threat to native biodiversity. The negative impacts of species invasions are particularly evident for freshwater crayfish faunas. 2. This study provides novel insight into the ecological effects of native and non‐native crayfish on zoobenthic communities (with emphasis on the non‐native rusty crayfish, Orconectes rusticus) across broad scales by combining a meta‐analysis of small‐scale experimental studies with a long‐term observational study conducted over a 24 year period in Sparkling Lake, Wisconsin, U.S.A. (46°00′N, 89°42′W). 3. The meta‐analysis summarised quantitatively the results of cage experiments for seven species of crayfish spanning four continents. We found that total zoobenthos densities (primarily Gastropoda and Diptera) were significantly lower in treatments containing crayfish relative to controls; a result that was significant for non‐native crayfish but not for crayfish in their native range, perhaps owing to a small sample size. In contrast to other species, rusty crayfish were also negatively associated with Ephemeroptera. 4. Results from the time series analysis comparing temporal trends in rusty crayfish and invertebrate abundances from Sparkling Lake were consistent with the findings from the meta‐analysis. Rusty crayfish were negatively correlated with the abundance of total zoobenthos, Diptera, Ephemeroptera and Odonata, as well as families of Trichoptera. 5. By coupling the results from short and long‐term research, our study offers greater insight into the nature of crayfish‐invertebrate interactions in aquatic systems, revealing consistent effects of invasive crayfish on native fauna. The control and management of invasive species is facilitated by the knowledge that well executed small‐scale studies may be extrapolated to understand larger‐scale ecological interactions.     

Crayfish in lakes and streams: individual and population responses to predation, productivity and substratum availability

1. In a correlative study, we investigated the relative importance of fish predation, refuge availability and resource supply in determining the abundance and size distributions of the introduced and omnivorous signal crayfish (Pacifastacus leniusculus) in lakes and streams. Moreover, the biomass and food selection of predatory fish was estimated in each habitat type and stable isotopes of carbon and nitrogen were measured in perch (Perca fluviatilis), the dominant predator in the lakes, and in its potential food sources (crayfish, juvenile roach and isopods). 2. In lakes, crayfish were the most frequent prey in large perch (46%), followed by other macroinvertebrates (26%, including the isopod Asellus aquaticus) and small fish (25%). Crayfish and fish dominated the gut contents of large perch with respect to biomass. Nitrogen signatures showed that perch were one trophic level above crayfish (approx. 3.4‰) and a two‐source mixing model using nitrogen isotope values indicated that crayfish (81%) contributed significantly more to perch isotope values than did juvenile roach (19%). A positive correlation was found between the abundance of crayfish and the biomass of large perch. Crayfish abundance in lakes was also positively correlated with the proportion of cobbles in the littoral zone. Lake productivity (chlorophyll a) was positively correlated with crayfish size, but not with crayfish abundance. 3. In streams, brown trout (Salmo trutta) were the most abundant predatory fish. Gut contents of large trout in a forested stream showed that terrestrial insects were the most frequently found prey (60%), followed by small crayfish (27%) and isopods (27%). In contrast to lakes, the relative abundance of crayfish was negatively correlated with the total biomass of predatory fish and with total biomass of trout. However, abundance of crayfish at sites with a low biomass of predatory fish varied considerably and was related to substratum grain size, with fewer crayfish being caught when the substratum was sandy or dominated by large boulders. The mean size of crayfish was greater at stream sites with a high standing stock of periphyton, but neither predator biomass nor substratum grain size was correlated with crayfish size. 4. Our results suggest that bottom‐up processes influence crayfish size in lakes and streams independent of predator biomass and substratum availability. However, bottom‐up processes do not influence crayfish abundance. Instead, substratum availability (lakes) and interactions between predation and substratum grain size (streams) need to be considered in order to predict crayfish abundance.     

An invaded invader: high prevalence of entocytherid ostracods on the red swamp crayfish <Emphasis Type="Italic">Procambarus clarkii</Emphasis> (Girard, 1852) in the Eastern Iberian Peninsula

The American red swamp crayfish Procambarus clarkii (Girard, 1852) was introduced in 1973 into the Iberian Peninsula for commercial purposes. As a result of both the expansion from the Iberian Peninsula and, probably, further introductions in other European countries, now it is widely distributed throughout much of Europe. The ecological impacts of this invading crayfish have received increasing attention, but nothing is known about its symbiotic entocytherid ostracods outside the American continent. The present survey has examined more than 200 crayfishes from 12 localities distributed over a wide area of Eastern Spain. Entocytherid ostracods were extracted from individual crayfishes and they were identified, counted, assigned to developmental instars and sexed. In all the study locations but one, we found at least one crayfish individual infected by entocytherid ostracods and the species determined was the same in all cases: Ankylocythere sinuosa (Rioja, 1942). The number of ostracods on individual P. clarkii varied notably in relation to crayfish size and also differed significantly among sampling sites. The crayfish size effects on ostracod densities might be related to the amount of resources and to the crayfish age and moulting frequency affecting ostracod distribution and population structure. In addition, the spatial variation in ostracod densities could also be related to site-specific habitat traits and the variability of crayfish population dynamics. Our study represents the first citation of an alien entocytherid species in Europe and demonstrates its wide distribution in the Iberian Peninsula. Further research is needed to know the potential effects of this ostracod species on the ecology of P. clarkii and of native species, with implications on the management of this aquatic invader.