Movement, dispersal and refuge use of co-occurring introduced and native crayfish

1. Dispersal and habitat use are key elements in determining impacts of introduced species. We examined if an invasive crayfish species showed a different pattern of movement, dispersal and refuge use to that of a species that it displaces. 2. Fifteen adults of the introduced American signal crayfish, Pacifastacus leniusculus and 20 adult native white‐clawed crayfish Austropotamobius pallipes were radiotagged in an area of river where in 2003 they co‐occurred and the spatial behaviour and habitat use of the two species were examined. Subsequent surveys at the study site in 2005 recorded no A. pallipes and the density of P. leniusculus had increased. 3. Clear differences in the spatial behaviour of the two species were found. The median distance moved per day by signal crayfish was over twice that of white‐clawed crayfish, and dispersal from release locations was higher in signal crayfish. A similar range of refuges with a high degree of overlap was used by both species suggesting that the potential for competitive interaction between the two species exists. 4. The greater movement and dispersal by signal crayfish recorded here supports the hypothesis that invaders are better dispersers than the species that they displace. In addition, movements by signal crayfish may allow them to make better use of patchy resources than white‐clawed crayfish and this, coupled with similar microhabitat use, may contribute to the observed replacement.     

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.     

The signal crayfish is not a single species: cryptic diversity and invasions in the Pacific Northwest range of Pacifastacus leniusculus

1. We used historical sources, morphology‐based taxonomy and mtDNA sequence data to address questions about the signal crayfish Pacifastacus leniusculus. These included evaluating unrecognised cryptic diversity and investigating the extent to which P. leniusculus may have been introduced within its presumed native range in the Pacific Northwest region of North America. Our study builds and expands on Pacific Northwest phylogeographic knowledge, particularly related to patterns of glacial refugia for freshwater species. 2. Extensive collections (824 specimens) from British Columbia (Canada), Idaho, Nevada, Oregon and Washington (United States) were used to characterise P. leniusculus at the mitochondrial 16S rRNA gene. Genetic groups within the species were elucidated by phylogenetics and amova ; evolutionary relationships within the most common and diverse group were investigated using a statistical parsimony haplotype network, a nested amova , and tests of isolation by distance. Morphological measurements were used to relate findings of molecular analyses to three historically recognised P. leniusculus subspecies and characterise cryptic diversity by morphology. 3. We found substantial cryptic diversity, with three groups highly distinct from P. leniusculus in discrete geographic regions: the Chehalis River glacial refugium, Central Oregon and the Okanagan Plateau. Disjunct distributions of P. leniusculus relative to these cryptic groups and known patterns of Pleistocene glaciation and landscape evolution cast doubt on whether P. leniusculus is native to some areas such as coastal drainages of northern Washington and southern British Columbia. Morphological traits previously used to characterise P. leniusculus subspecies still persist but may be incapable of distinguishing P. leniusculus from newly discovered cryptic groups. 4. Cryptic diversity found within P. leniusculus highlights the pressing need for a thorough investigation of the genus Pacifastacus using data based on more extensive gene and taxon sampling. It also warrants conservation attention, as introductions of P. leniusculus within the Pacific Northwest may carry risks of hybridisation and introgression for cryptic groups. Owing to high genetic diversity and limited dispersal capacity relative to more vagile organisms like freshwater fish, crayfish of the genus Pacifastacus offer powerful potential insights into the geological history and phylogeography of the Pacific Northwest region. Finally, by shedding light on the long‐neglected native range of P. leniusculus, our results should also better inform our understanding of potential source populations for, and the ecology of, this important invasive species in regions including Europe, Japan and elsewhere in North America.     

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.     

Metabolic rate in the signal crayfish (Pacifastacus leniusculus) is temporally consistent and elevated at molting

Although the temporal consistency of resting metabolic rate in individual animals is generally considered to be a universal phenomenon, studies on invertebrates are still scarce. Here, we studied the repeatability of standard metabolic rate in the signal crayfish (Pacifastacus leniusculus). We measured oxygen consumption twice on the same individuals, on average in 97-day intervals. At intermolt stage, the standard metabolic rate was a repeatable trait. However, molting increased significantly the minimum metabolic rate, thus emphasizing the role of animal physiological state in determining the rate of metabolism.     

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.     

Diurnal variations in physiology and behaviour of the noble crayfish Astacus astacus and the signal crayfish Pacifastacus leniusculus

Heart rate, locomotor activity, and oxygen consumption were recorded simultaneously and continuously in seven individuals of the noble crayfish Astacus astacus (Linneus 1758) and seven individuals of the signal crayfish Pacifastacus leniusculus (Dana 1852). The recordings were made in the laboratory over 7 days at 15°C under a 12 : 12 h dark : light regime. Circadian rhythms in heart rate, locomotor activity and oxygen consumption were found both in A. astacus and P. leniusculus. Increased heart rate, locomotor activity, and oxygen consumption levels during night time in both A. astacus and P. leniusculus illustrated expression of nocturnal behaviour. No differences in oxygen consumption levels were observed between A. astacus and P. leniusculus. Also, no significant difference between heart rate levels or heart rate variances was found in A. astacus and P. leniusculus at night. During day, however, heart rate levels, heart rate variances and locomotor activity were higher in P. leniusculus than in A. astacus. The higher activity level in P. leniusculus than in A. astacus during daytime indicates that P. leniusculus is less strictly nocturnal than is A. astacus.     

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.     

Testing the differential predation hypothesis for the invasion of rusty crayfish in a stream community: laboratory and field experiments

1. We tested the hypothesis that the non‐native rusty crayfish (Orconectes rusticus) is less vulnerable to predators than two native species (O. propinquus and O. obscurus) it is replacing in streams of the upper Susquehanna River catchment (New York, U.S.A.). 2. We used laboratory experiments to compare species‐specific predation rates by smallmouth bass (Micropterus dolomieu) on crayfish of equal size and field tethering experiments to compare relative predation rates between native O. propinquus and non‐native O. rusticus by the suite of crayfish predators in our system. We predicted that crayfish size would affect predation rate but that predation rates would be equal among species when size was controlled. 3. We also tested for two potential artefacts of tethering. We tethered crayfish in cages to test whether the ability to escape from tethers is size specific, and we tested whether tethering alters differential predation among crayfish species by the smallmouth bass. 4. In the laboratory, smallmouth bass predation on rusty crayfish was lower than on either of the native species. In the field, predation risk for tethered crayfish was inversely related to size but did not differ among species when size was taken into account. Because rusty crayfish in the field experiment were slightly larger than the native species, as in nature, mortality was overall lower for the rusty crayfish. 5. In cages, smaller crayfish were more probably to escape from tethers than larger ones, an artefact that may partially confound results from our tethering experiments. Unexpectedly, tethering nearly eliminated predation by smallmouth bass. This artefact prevented us from testing for an interaction of tethering with differential predation and means that the results of field tethering experiments do not include any contribution from smallmouth bass predation. 6. Our experiments highlight the importance of explicitly considering potential artefacts that could confound results. 7. Our results indicate that differential predation contributes to the rusty crayfish's invasion of a stream community. In our study system, predation rates on rusty crayfish are lower than for native species mostly because of selection by predators for smaller crayfish; species‐specific characteristics such as behaviour that further reduce predation may also contribute, especially where smallmouth bass predation is important.     

Assessing invasion risk across taxa and habitats: life stage as a determinant of invasion success

Aim Many aquatic invertebrates produce dormant life‐history stages as a means to endure inhospitable environments and to facilitate natural long‐distance dispersal, yet we have little understanding of the role of dormant stages as a mechanism for human‐mediated introductions of non‐indigenous species. We explore the survival of invertebrate dormant eggs in collected ships’ ballast sediment over a 1‐year period to determine relative invasion potential across taxa (i.e. rotifers, copepods, cladocerans and bryozoans) and different habitats (freshwater, marine). Location Canadian Atlantic and Pacific coasts and Laurentian Great Lakes. Methods During 2007 and 2008, 19 ballast samples were collected as a part of a larger study. The degradation rate of dormant eggs was assessed by enumerating dormant eggs and by conducting viability hatching experiments. Results Taxa examined included rotifers, copepods, anomopods, onychopods and bryozoans. Dormant eggs of rotifers degraded at the highest rate of all taxa examined, with no viable eggs remaining within 10 months. Copepods showed a less rapid degradation rate than rotifers. The degradation rate of anomopod dormant eggs was significantly slower than that of both rotifers and copepods. Onychopods and bryozoans did not visibly degrade at all over 12 months. Viability hatching experiments were successful for rotifers, copepods, and anomopods. Onychopods and bryozoans did not hatch during any of the three hatching trials. Main conclusions Dormancy is not equally beneficial to all invertebrate taxa. Our results indicate that dormant eggs of rotifers and copepods degrade at a rapid rate and may not pose high invasion risk. In contrast, the slow degradation rate of anomopod dormant eggs and the lack of degradation of onychopod and bryozoan dormant eggs could result in high invasion risk because of their accumulation in ballast tanks. Species having resistant dormant eggs mostly belong to freshwater taxa making freshwater habitats at higher invasion risk by dormant invertebrates than marine habitats.     

Influence of suburban edges on invasion of Pittosporum undulatum into the bushland of northern Sydney,Australia

Abstract Suburban edges are shown to exert a major influence on invasion of the native Pittosporum undulatum Vent, in dry sclerophyll bushland of northern Sydney. Transect data from fifteen urban bushland sites spanning approximately 90 years of development indicate significant increases of P. undulatum with time. Basal area, density and frequencies of plants in all size-classes increased significantly with age of adjoining development. Mean basal area at edges of old sites was 5700 times greater than in comparable bushland in larger reserve interiors. The effect of age was compounded by the greater impact of edge effects in narrower reserves of older suburbs. Basal areas and proportion of reproductive plants decreased significantly with distance into the reserve. Older sites contained larger but fewer individuals at the edge, compared with high densities of smaller plants further into the reserve, suggesting an advanced successional stage. The overall relationship of density with distance was not significant. The observed clumping of seedlings beneath canopies is consistent with previous work on seed dispersal by vectors, and enhanced seedling recruitment in safe sites. Pittosporum basal area and density reached no maximum levels with age, implying ongoing invasion. The study indicated some factors which appeared to promote the establishment of P. undulatum in the study area. Signs of fire were negatively correlated across all sites with basal area and density of P. undulatum. Human disturbance was positively correlated with the same variables. On the basis of findings in this study, some broad priorities for management of Pittosporum in urban bushland are suggested.     

Temporal shifts in physical habitat of the crayfish,Orconectes neglectus (Faxon)

A total of 300 samples was collected from February 1985 to August 1986 in a medium order Ozark Mountain stream. Physical habitat measurements of temperature, mean water column velocity, depth, and substrate character were recorded for each of the 25 monthly samples along with length and sex of all individuals of Orconectes neglectus (Faxon). Analysis of habitat utilization and suitability (or preference) was conducted using exponential polynomial models of hydraulic stress models. There appeared to be equal preference for depth over the range measured. Both substrate and velocity preference curves were bimodal with each mode designating certain crayfish size classes. Young-of-the-year were found primarily in cobbled, high velocity areas while adults were found in low velocity, macrophyte beds. Utilization curves for laminar sublayer thichness also reflected size-dependent phenomena where young-of-the-year were found in thin sublayer areas and adults were found primarily in thick sublayers. When separated by time and size, adults were found to occupy higher velocity, cobbled habitats during at least two months. This time period corresponded with the time of egg-bearing and further analysis yielded a time-dependent habitat suitability surface which accounted for this movement pattern. We suggest that the application of these suitability surfaces, which reflect habitat changes during the annual life cycle, will produce more accurate predictions of density and will allow better habitat management decisions under various regulated flow scenarios.     

Towards a mechanistic understanding of individual-level functional responses: Invasive crayfish as model organisms

1. In novel communities, a rising number of new and emerging invasive species interact with resident species, some of which are non-native themselves. We implemented an innovative trophic interaction framework for novel communities and quantified the interaction strength and impact potential of a truly novel species (marbled crayfish Procambarus virginalis) with a resident non-native counterpart (spiny-cheek crayfish Faxonius limosus). As prey, we used Dreissena mussels, which are non-native as well and now hold a key position in many European and North American aquatic ecosystems.
2. For both crayfish species, we predicted functional responses based on a mechanistic model that we parameterised with a set of experimental observations of foraging behaviour and satiation. We compared these predicted functional responses to empirically observed responses. In addition, we incorporated behavioural traits such as aggression, activity, and boldness in the comparisons between the species and individuals to determine their influence on functional responses. We tested individuals from aquarium stocks as well as naturalised individuals from invaded water bodies.
3. Altogether, we performed 1,095 experiments with 26 individual crayfish. We found that per capita predation of spiny-cheek crayfish exceeded that of marbled crayfish from aquaria and naturalised individuals. Functional responses differed between species and were mostly higher for spiny-cheek crayfish males. Marbled crayfish, however, were more voracious and reached satiation more slowly. Consumption rates correlated with aggression for marbled crayfish and with an aggressive threat response for spiny-cheek crayfish.
4. We conclude that spiny-cheek crayfish can reach higher short-term consumption rates than marbled crayfish, but both species probably do not substantially affect Dreissena mussel populations in the field. For marbled crayfish, high long-term consumption, interspecific aggression, and reproduction rates can promote their establishment and spread. Risk assessments of these invaders should be improved by considering numerical responses, and different prey organisms and predators.

     

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.     

Disentangling relationships between habitat conditions, disturbance history, plant diversity, and American black cherry (Prunus serotina Ehrh.) invasion in a European temperate forest

Whether non-native plant invasions are causes, consequences, or independent of the low species diversity in recipient ecosystems remains a debated question. We tried to test these three hypotheses in the special case of the American black cherry ( Prunus serotina Ehrh.), a gap-dependent tree species, which is invading European temperate forests. We compared plant communities, soil properties, and disturbance history between P. serotina -invaded and uninvaded paired-stands in a managed mixed forest. Relationships between invasion, disturbances, plant communities, and environmental conditions were investigated using redundancy analyses with variation partitioning. Several soil characteristics differed between paired stands, but were rather components of stand invasibility than invasion effects, except for topsoil available phosphorus. The disturbance history was similar among paired stands except for the amount of storm-induced tree falls, which correlated with the invader's density. Wild boar-disturbed soil areas were more important beneath P. serotina canopies, suggesting a positive feedback on its own establishment. Overall, species assemblages in invaded and uninvaded stands were similar; their ecological inconsistency suggested a management-sustained non-equilibrium. Habitat conditions and disturbances explained most of the variation in both plant diversity and P. serotina density, the last two factors exhibiting a weak direct association. We conclude that in managed forest ecosystems where plant communities are mainly driven by non-interactive factors and immigration processes, non-native plant species can naturalize without being directly influenced by measured features of the plant community in the receiving environment on the short term.     

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.     

Behavioral effects of invaders: alien crayfish and native sculpin in a California stream

Invasive crayfish have been shown to have negative impacts on a range of taxa, though the mechanisms for those effects have not always been evaluated. Signal crayfish (Pacifastacus leniusculus) in Sagehen Creek were associated with reduced growth rates and gut fullness of Paiute sculpin (Cottus beldingi) in earlier experiments. This paper assesses potential behavioral mechanisms of competition between the two species. I conducted experiments to determine crayfish effects on sculpin behavior and habitat use in a stream observation facility at the Sagehen Research Station, California, USA. Sculpin reduced their use of refuges and pools, shifted into higher-velocity microhabitats, and spent more time fleeing in the presence of crayfish. Crayfish used refuges, pools, and low-velocity habitats more than sculpin in either treatment. Both species were notably nocturnal, with most activity at dusk and night observations, although crayfish were more strongly so than sculpin. Detailed field surveys of lower Sagehen Creek found that potential refuges (unembedded rocks) were closely associated with total crayfish and sculpin numbers, suggesting that cover is at least sometimes limiting under natural conditions. By displacing sculpin from refuges and pools and increasing their activity rate, crayfish may increase the likelihood of predation on sculpin. Behavioral shifts in sculpin appear to be at least partly responsible for the reduced growth rates of sculpin in the presence of crayfish.