Immigration rates of signal crayfish (Pacifastacus leniusculus) in response to manual control measures

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. A predicted outcome of trapping bias in riparian habitats is that removed individuals could be replaced by large crayfish immigrating from surrounding, untrapped, areas. We tested the hypothesis that removal by trapping of American signal crayfish from a U.K. river would result in increased rates of immigration, and increased distances moved, of crayfish from untrapped areas. 3. We studied four stretches of the River Windrush each 1 km in length and divided into three sections; a 250‐m long upstream section, a 500‐m middle section and a 250‐m downstream section. At two sites (removal sites), signal crayfish were trapped and removed from the 500‐m middle sections; at the other two (non‐removal), they were marked and returned. All crayfish captured in the upstream and downstream sections were marked and returned. 4. Probability of capture was higher for larger individuals with both chelae intact, and larger crayfish were more likely to immigrate from the upstream and downstream sections into the middle. The percentage of captured crayfish immigrating into the middle sections was the same (3.7%) in both removal and non‐removal sites. However, the mean distance that crayfish moved when immigrating was significantly greater at removal sites (239 m) than at non‐removal sites (187 m). 5. These results imply that removal of large individuals may have reduced the potential for interference competition by increasing the relative competitiveness of the immigrating individuals and permitting them to make larger movements. Consequently, the impact of manual removal strategies, both on the signal crayfish population and other biota affected by them, is likely to be reduced at the point of removal, but to extend at least 200 m beyond the trapped length of river.     

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.     

Habitat characteristics of crayfish (<Emphasis Type="Italic">Paranephrops planifrons</Emphasis>) in New Zealand streams using generalised additive models (GAMs)

Quantitative data on the habitat characteristics of stream crayfish have been generally lacking and competing demands on water resources has created a need to address this knowledge gap. We investigated day-time habitat relationships of stream crayfish (Paranephrops planifrons White) from 793 quadrats at 30 rivers and streams in the North Island, New Zealand to develop models of koura presence–absence and abundance. The model (stepwise GAM) included width, cover, median substrate size, edge location, velocity and depth, and correctly predicted presence–absence of crayfish (8–39 mm OCL) at 73.4% of quadrats and of young-of-the-year (YOY) ≤8 mm OCL at 83.4% of quadrats. Streams ranged from 1.6 to 11.5 m in width and the probability of finding both crayfish size classes reduced sharply as streams became wider than 6 m and as the substrate became large (i.e., boulder > 256 mm). Crayfish, particularly YOY, were most likely to be found in association with cover and at the stream edge. YOY were associated with shallow depths and fine substrates, whereas larger crayfish showed a preference for cobble substrate. Undercut banks, leaf litter, tree roots, and woody debris were strongly related to the presence–absence of crayfish. The model for crayfish abundance (log-linear Poisson GAM) explained 50% of the variation between quadrats with cover, velocity, edge location, depth, and the overall crayfish abundance at each particular stream being significant variables. Highest crayfish numbers were recorded in still or slow flowing water, with the majority occurring where velocities were below 0.4 m/s. Water depths up to 0.7 m were sampled, but highest numbers were found in depths of 0.2–0.3 m. Our presence–absence model determined variables that were significant over all streams, whereas our abundance model determined variables that were significant within streams. Use of the GAMs models enabled us to untangle the multiple factors contributing to habitat selection. Cover, velocity, and locations at the stream edge were important determinants of both presence–absence and abundance. Generally, substrate was important when comparing between streams, but not within streams, whereas depth was a significant determinant of abundance within streams, but not presence–absence between streams. Handling editor: K. Martens     

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.     

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.     

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.     

Interaction of Press and Pulse Disturbance on Crayfish Populations: Flood Impacts in Pasture and Forest Streams

Interactions between press and pulse disturbances can significantly impact benthic macroinvertebrate (crayfish) populations. Press disturbances such as land-use change can make crayfish more vulnerable to the detrimental effects of pulse disturbances, by changing the habitats available to crayfish. The impact of a pulse disturbance, a major flood (1 in 28 years return period), on crayfish (Paranephrops planifrons) was significantly greater in pasture than native forest streams. Population densities of crayfish in three forest and three pasture streams had been measured seasonally for 2 years prior to the flood and many crayfish had been marked with individually identifiable tags. Low numbers of marked crayfish were recaptured after the flood in the forest streams, but no marked crayfish were identified in any pasture stream. Crayfish densities in one pasture stream declined from an average of 5 m^–2 prior to the flood to <1 m^–2 soon after the flood and it took 3 years for the population to show evidence of recovery. Macrophytes and cobbles, the dominant habitats of crayfish in pasture streams, did not appear to provide stable refugia during the flood. Habitat stability was linked to the riparian zone in forest streams where undercut banks, tree roots, and pools were important habitats for crayfish. Frequent pulse disturbance could affect population persistence of refugia-dependent species when the pressure of land-use change affects the stability of habitats, but this may only be evident over long time scales.     

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.     

Macrophyte diversity and composition in relation to substratum characteristics in regulated and unregulated Danish streams

1. The objective of the present study was to examine how the physical stream environment in regulated and unregulated lowland streams affects the diversity and distribution of macrophyte communities. We analysed the abundance, distribution and composition of macrophytes, together with physical parameters, in seven regulated and seven unregulated unshaded Danish stream reaches. 2. Total macrophyte coverage was similar in the regulated and unregulated streams, but species richness and Shannon diversity were higher in the unregulated streams. Overall, we found fifty-two different species in the regulated stream reaches and sixty-two in the unregulated stream reaches. The spatial distribution of macrophytes on the stream bottom was more heterogeneous in the unregulated streams. 3. We found positive correlations between the coverage and diversity of macrophytes and the coverage of coarse-textured substratum types on the stream bottom, as well as between macrophyte coverage and diversity and substratum heterogeneity. We also found that the macrophytes were more heterogeneously distributed where substratum heterogeneity was greater. 4. The species growing both submerged and emergent were more abundant in the regulated streams, whereas species growing only submerged were more abundant in the unregulated streams. Species growing submerged, species growing both submerged and emergent, and species only growing emergent segregated differently in a canonical correspondence analysis ordination. The submerged species were primarily associated with coarser-textured substrata, whereas species growing both submerged and emergent, and species growing only emergent were associated with finer-textured substrata. 5. The most abundant species in the regulated streams, Sparganium emersum, accounting for almost one-third of the total macrophyte coverage, was primarily associated with clay and sandy bottom substrata, whereas the most abundant species in the unregulated streams, Batrachium peltatum, was primarily associated with gravel and stony substrata.     

Variation in local abundance and species richness of stream fishes in relation to dispersal barriers: implications for management and conservation

1. Barriers to immigration, all else being equal, should in principle depress local abundance and reduce local species richness. These issues are particularly relevant to stream‐dwelling species when improperly designed road crossings act as barriers to migration with potential impacts on the viability of upstream populations. However, because abundance and richness are highly spatially and temporally heterogeneous and the relative importance of immigration on demography is uncertain, population‐ and community‐level effects can be difficult to detect. 2. In this study, we tested the effects of potential barriers to upstream movements on the local abundance and species richness of a diverse assemblage of resident stream fishes in the Monongahela National Forest, West Virginia, U.S.A. Fishes were sampled using simple standard techniques above‐ and below road crossings that were either likely or unlikely to be barriers to upstream fish movements (based on physical dimensions of the crossing). We predicted that abundance of resident fishes would be lower in the upstream sections of streams with predicted impassable barriers, that the strength of the effect would vary among species and that variable effects on abundance would translate into lower species richness. 3. Supporting these predictions, the statistical model that best accounted for variation in abundance and species richness included a significant interaction between location (upstream or downstream of crossing) and type (passable or impassable crossing). Stream sections located above predicated impassable culverts had fewer than half the number of species and less than half the total fish abundance, while stream sections above and below passable culverts had essentially equivalent richness and abundance. 4. Our results are consistent with the importance of immigration and population connectivity to local abundance and species richness of stream fishes. In turn, these results suggest that when measured at appropriate scales (multiple streams within catchments), with simple protocols amenable to use by management agencies, differences in local abundance and species richness may serve as indicators of the extent to which road crossings are barriers to fish movement and help determine whether road‐crossing improvements have restored connectivity to stream fish populations and communities.     

Ranging behaviour of a large freshwater invertebrate, the white-clawed crayfish Austropotamobius pallipes

• 1 Radio‐telemetry and mark‐recapture methods were used to study the summer movements of adult and juvenile white‐clawed crayfish, Austropotamobius pallipes from a wild population in a small braided stream, Dalton Beck, North Yorkshire, U.K. Radio‐transmitters were attached to the chelae of 18 large (> 35 mm carapace length) crayfish and individuals were subsequently located to within 0.15 m. Additionally a total of 888 crayfish were marked with carapace brands, and 83 were recaptured.
• 2 Radio‐tracked crayfish exhibited significantly greater local activity at dusk (21.00–00.00) than at dawn (03.00–06.00), or during morning (09.00–12.00) and afternoon (15.00–18.00) monitoring periods.
• 3 The greatest movements of radio‐tracked crayfish occurred within 2 days of release. After this time, periods of residence were interspersed by movements to new locations, interpreted as establishment of ephemeral home areas. It is suggested that the initial large movements were the result of a ‘fright response’ following capture.
• 4 Movements varied widely between individuals, some moving more than 300 m in 10 days, while others showed little movement over an equivalent time period. Mean (±SE) daily movements were 4.6 ± 3.0 m for males and 1.5 ± 1.0 m for females. Although crayfish often used specific home sites for in excess of 7 days, displaced animals did not return to home sites.
• 5 The total distances travelled and the mean distance travelled per day by individual radio‐tagged crayfish did not differ significantly between upstream or downstream directions or between males and females. This was also the case for marked crayfish used in mark‐recapture studies.
• 6 Positive correlations between distance moved per day and size (carapace length) were found for downstream movements by male and female crayfish, but not for upstream movements.
• 7 Some preliminary observations of the response of crayfish to flood events suggested that these could be catastrophic with two out of five tracked crayfish found dead after a high stream‐discharge event.

     

Flow variability maintains the structure and composition of in‐channel riparian vegetation

1. Naturally variable river flows are considered to be important for structuring riparian vegetation. However, while the importance of floods for the ecology of riparian vegetation is well recognised, much less is known about the importance of small fluctuations in river flows. 2. We investigated the effect of water supply diversion weirs on the riparian vegetation of upland streams. These weirs remove within‐channel fluctuations in flow but do not prevent large floods downstream. We surveyed the in‐channel and banktop vegetation of five streams, three of which were regulated by weirs and two of which acted as controls. 3. Unexpectedly, we observed greater species richness within the channel downstream of the weirs. This was because of increased numbers of exotic and terrestrial (‘dry’) plant species. Grass cover was also greater downstream of the weirs. There were no significant differences in the banktop vegetation between the upstream and downstream sites of the regulated streams. 4. Our results highlight the role of within‐channel flow variability in maintaining the composition of vegetation within the stream channel. We suggest that greater species richness does not necessarily indicate a less‐disturbed environment. Rather, a greater number of ‘dry’ species is indicative of the impacts of flow regulation. 5. Small fluctuations in river flows are probably necessary to protect the ecosystem structure and function of regulated streams. It is recommended that variable within‐channel flows be provided in regulated streams.     

Environmental DNA detects a rare large river crayfish but with little relation to local abundance

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Predicting the distribution of invasive crayfish (Pacifastacus leniusculus) in a Kusiro Moor marsh (Japan) using classification and regression trees

Summer distributions of the invasive signal crayfish (Pacifastacus leniusculus) were investigated in relation to physicochemistry in a Kusiro Moor marsh and its inflows and outflows in northern Japan. Maximum crayfish abundance and biomass were 1.04 individuals/m² and 3.56 g dry mass (DM)/m² in littoral marsh habitats, and 5.84 individuals/m² and 13.48 g DM/m² in stream habitats. Classification tree analysis was used to predict crayfish occurrence at 102 sites from all habitats (i.e. littoral marsh, pelagic marsh and stream) while regression tree analyses were used to predict crayfish abundance at littoral marsh and stream sites separately. The classification tree showed that crayfish occurrence was primarily determined by undercut bank volume regardless of habitat identity. When undercut bank volume was <0.0054 m³, crayfish were predicted to be absent at marsh sites, but expected to occur at stream sites where pH and water temperature exceeded 6.5 and 14.3°C, respectively. The regression tree using only littoral marsh sites showed that undercut bank volume, followed by dissolved oxygen level, determined the splits of the tree. Crayfish abundance was highest when undercut bank volume was >0.61 m³, and moderately high when dissolved oxygen was >9.09 mg/l and undercut bank volume was <0.61 m³. On the other hand, the regression tree using only stream sites showed that water temperature was the major predictor that determined the splits. We discuss the roles of physicochemical factors as limiting factors of the distribution pattern of the invasive crayfish.     

Summer use of a small stream by fish and crayfish and exchanges with adjacent lentic macrohabitats

1. Movements between a stream reach and two adjacent lentic macrohabitats, a beaver pond and a lake, were followed for the Appalachian crayfish and two fish species, brook charr and brown bullhead, over an 85‐d period from early June to late August, and were analysed in relation to water level, maximum water temperature, photoperiod length, lunar luminosity, and age, by use of time‐series regressions. 2. Brook charr showed strong net immigration to the stream reach for underyearling (age class 0+) fish but net emigration for 1+ fish. Both immigration and emigration were positively related to water level and temperature; migratory responses to temperature were age‐specific. 3. Brown bullhead used the stream primarily as a corridor for downstream migration from the beaver pond to the lake. As with brook charr, water level and temperature had a positive effect on movement and responses were stronger in younger individuals. 4. Crayfish emigrated from the stream during the summer. Movements were positively related to increases in water level and temperature, with responses of 1+ crayfish much stronger than those of older individuals. 5. The results indicate that the stream tributary played different roles for brook charr (nursery), brown bullhead (dispersal corridor) and Appalachian crayfish (early summer refuge), and suggest that variation in water level or temperature resulting from climatic change or local anthropogenic activities might modify connectivity between macrohabitats, particularly for younger individuals.     

The importance of crayfish in the breakdown of rhododendron leaf litter

1. Rhododendron (Rhododendron maximum) is a common evergreen shrub in riparian areas of the southern Appalachians, where its leaves can comprise a large proportion of leaf litter in streams. However, they are relatively refractory and generally considered a low quality food resource for detritivores. 2. Our objective was to assess whether macroconsumers [primarily crayfish (Cambarus bartonii)] influence rhododendron leaf breakdown in a forested southern Appalachian stream in both summer (when leaves other than rhododendron are relatively scarce) and autumn (when other leaves are relatively abundant). We conducted two leaf decay experiments, one in summer and one in autumn, using pre‐conditioned leaves. Macroconsumers were excluded from the benthos of a fourth‐order stream using electric ‘fences’; we predicted that excluding macroconsumers would reduce the decay rate of rhododendron leaves in both summer and autumn. 3. In both experiments, breakdown rate was lower in exclusion treatments. Macroconsumers accounted for approximately 33 and 54% of rhododendron decay in summer and autumn, respectively. We attribute this effect to direct shredding of rhododendron by crayfish. Biomass of insect shredders, insect predators and fungi did not differ between control and exclusion treatments, indicating that insectivorous sculpins (Cottus bairdi) had no effect on rhododendron decay and that omnivorous crayfish did not exert an indirect effect via alteration of insect or fungal biomass. 4. The influence of shredding insects varied between summer and autumn. In summer, when other, more palatable leaf types were not available, rhododendron leaf packs appeared to provide ‘resource islands’ for insect shredders. There was a significant inverse relationship between insect shredders and leaf pack mass in the summer exclusion treatment: insects were the only organisms eating leaves in this treatment and, as shredder biomass increased, remaining leaf pack mass decreased. In the control treatment, however, we did not see this relationship; here, the effect of insect shredders was presumably swamped by the impact of crayfish. In autumn, when other leaves were abundant, insect shredder biomass in rhododendron leaf packs was less than one‐third of summer values. 5. Even at low density (approximately 2 m^–2) crayfish were able to influence an ecosystem process such as leaf decay in both summer and autumn. Given the threatened status of many crayfish species in the United States, this finding is especially relevant. Even small alterations in crayfish assemblages, whether via loss of native species and/or introduction of exotic species, may have significant repercussions for ecosystem function.     

Stream channel geomorphology influences mussel abundance in southern Appalachian streams, U.S.A.

1. Studies of North American streams have shown that hydraulic parameters and stream geomorphology can explain unionid mussel abundance at both the reach and catchment scale. However, few studies have examined applicability of hydrogeomorphic variables across broader spatial scales, such as across whole catchments, or have elucidated conditions under which spates can affect mussel populations in streams. 2. We quantified freshwater mussel abundance and species richness and their physical habitat at 24 sites in eight streams in southern Appalachian catchments in 2000 and 2001. In addition, we modelled site‐specific hydraulic parameters during summer baseflow and bankfull stages to estimate high‐ and low‐discharge conditions, respectively. 3. Mussel abundance was related to stream geomorphology, whereas richness was related to stream size. Baseflow habitat parameters explained only minor variation in abundance or richness, and both measures were highly correlated with mean current velocity or stream size. Bankfull shear stress composed a relatively low proportion of overall mussel habitat variability, but it accounted for significant variation in abundance and richness. 4. Mussel abundance was highly variable at sites subject to low‐shear stress during spates, whereas abundance always was low at sites subject to high‐shear stress. These data suggest that habitat conditions during floods, rather than those at summer baseflow, limit the abundance of mussels in Appalachian streams. These data also suggest that mussel abundance and assemblage structure may be sensitive to any changes in channel geomorphology and hydraulic conditions that might result from land use in the catchment.     

Grazer species effects on epilithon nutrient composition

1. Field and laboratory experiments were conducted to investigate the excretion stoichiometry of nitrogen (N) and phosphorus (P) of two benthic macroinvertebrate grazers, the crayfish Orconectes propinquus and the snail Elimia livescens, that differ in body stoichiometry (mean body molar N : P 18 and 28, respectively). Crayfish excretion had a significantly higher ammonium : soluble reactive phosphorus (SRP) ratio in the laboratory and in three natural streams than did snails, as predicted by ecological stoichiometry theory. 2. In greenhouse recirculating artificial streams, treatments consisting of crayfish, snails, or no grazers were used to examine responses in dissolved nutrient concentrations and epilithon nutrient composition and limitation. SRP concentrations depended upon the grazer species, with the snail treatment having a higher SRP concentration than other treatments (P < 0.05). Dissolved inorganic N was not affected by grazers, but appeared to be rapidly incorporated in epilithon. 3. Epilithon N content was dependent upon the grazer species present, with the crayfish treatment having a significantly higher N content than other treatments (P = 0.001). No grazer species effects on epilithon P content were found. However, both grazer treatments had significantly lower epilithon P content than the no‐grazer treatment. 4. Traditionally, studies have focused on how grazer‐induced structural changes to epilithon can alter epilithon nutrient dynamics, but this structural mechanism could not solely explain differences in epilithon nutrient contents and ratios in the present study. Our results rather suggest that benthic grazers can alter epilithon nutrient composition and limitation via nutrient excretion. Consequently, macroinvertebrate grazers may serve as ‘nutrient pumps’ that partly regulate the availability of nutrients to algae in stream ecosystems.     

Microhabitat use by foraging white-clawed crayfish (<Emphasis Type="Italic">Austropotamobius pallipes</Emphasis>) in stream pools in the NE Iberian Peninsula

The white-clawed crayfish (Austropotamobius pallipes) is an endangered species across most of its distribution range, and information on its ecological requirements is needed to implement effective conservation measures. Its habitat use has been studied in different areas and at various spatial scales. However, being a nocturnal species, there is scarce information on its habitat selection during foraging periods. In this work we analyse nocturnal habitat use of white-clawed crayfish in pools of a small stream in the northeastern Iberian Peninsula at two different scales: (1) microhabitat selection and (2) pool characteristics. Large crayfish showed a clear positive selection for deeper microhabitats, a selection pattern that was weaker for medium-sized crayfish and absent for small ones. On the other hand, crayfish of all sizes avoided cobble and boulder microhabitats and positively selected fine substrate and more exposed microhabitats. Crayfish abundance in pools was positively influenced by pool area, pool depth and the availability of fine substrates, especially silt. While studies on white-clawed crayfish habitat use have often stressed the importance of rough substrates as crayfish refuge, our results show that fine substrates are positively selected by foraging crayfish of all size classes and promoted active crayfish abundance in pools. These apparently contradictory results may be due to the differences in microhabitat preferences exhibited by active and inactive crayfish. Thus, our results help to better complete the picture of white-clawed crayfish habitat requirements.     

Contrasting effects of an invasive crayfish (Procambarus clarkii) on two temperate stream communities

1. The effects of omnivorous exotic species on native communities are often difficult to predict because of the broad diets and behavioural flexibility of the omnivore, and the diverse abiotic and biotic characteristics of invaded systems. We investigated experimentally the effects of a gradient of density of the introduced, omnivorous red swamp crayfish Procambarus clarkii (Decapoda: Cambaridae) on two stream communities in southern California, U.S.A. 2. The Ventura River is a clear, flowing stream with a cobble substratum, with abundant algae but low densities of large invertebrates, small herbivores and snails. The Santa Ynez River at the time of the study consisted of a series of drying pools underlain by sand, with abundant charophytes, large predatory invertebrates and herbivores, including snails. 3. In the Ventura River, periphyton biomass and inorganic sediment decreased with increasing crayfish abundance, but in the Santa Ynez River, periphyton and sediment were unrelated to crayfish densities. 4. In the Ventura River, the biomass and density of all benthic invertebrates combined, chironomids, micropredators, the meiofauna (chydorid cladocerans, copepods and ostracods), and specific predatory and herbivorous taxa, as well as taxon richness, were negatively related to crayfish density. In the Santa Ynez River, the biomass and average body size of benthic invertebrates, predatory invertebrates, herbivores and chironomids, but not total invertebrate density or taxon richness, were negatively related to crayfish density. 5. Fewer large predatory invertebrates and snails (Physella gyrina) in both streams, and baetid mayflies in the Ventura River, were visible at night in channels where crayfish were abundant. Snails responded to crayfish by moving above the water line in the Santa Ynez River, but not in the Ventura River. 6. We suggest that the same omnivore had different effects on these neighbouring streams because of crayfish predation on large invertebrates in the Santa Ynez River and the scarcity of such prey in the Ventura River, leading to increased crayfish grazing on periphyton, and reductions in periphyton‐associated invertebrates, in the Ventura River.