Success and failure in a lotic crayfish invasion: the roles of hydrologic variability and habitat alteration

1. This paper examines the distribution, habitat relationships, and potential for spread of non‐native signal crayfish (Pacifastacus leniusculus) in streams of the Truckee River catchment, California, U.S.A. Crayfish associations with natural features and with impoundments and flow alteration were examined in a survey of 33 streams. Abundance changes were followed over 5 years, which included some of the highest and lowest flows on record, in three streams, two unregulated and one regulated. Movement of marked crayfish was studied in one 0.5 km stream reach just upstream from a reservoir. 2. Signal crayfish were most abundant in low‐gradient streams and were positively associated with proximity to reservoirs (both upstream and downstream). Crayfish were more likely to be found in regulated than unregulated sites, and did not occur in sites upstream of barriers, such as culverts, that separated them from reservoirs or lakes. 3. Crayfish declined in abundance in years following particularly intense and prolonged wet‐season spates, leading to a negative association between crayfish abundance and both peak discharge and duration of bankfull flows. 4. Crayfish moved distances of up to 277 m, and at rates of up to 120 m day^?1, suggesting significant dispersal ability. Larger crayfish moved greater distances and were more likely to move downstream. Female crayfish showed a pattern of upstream movement in early summer and downstream movement late in the summer, opposite the pattern found in two other studies. 5. These results suggest that natural or artificial gradient barriers and, in regulated systems, management of flow regimes to include bankfull or greater flows may help to control invasive crayfish in streams.     

Winter movements and activity of signal crayfish Pacifastacus leniusculus in an upland river,determined by radio telemetry

Radio-telemetry was used to study the late autumn and winter movements of twenty adult signal crayfish Pacifastacus leniusculus (32.9–63.8 mm carapace length) an introduced exotic crayfish species, in the upland River Wharfe, northern England. The distances moved during the study varied greatly between individuals (0–328 m). Movements were generally sporadic; crayfish would remain in one position for several weeks and make occasional movements to new locations. Total distances travelled, linear range and ranging area did not differ significantly between males and females. The distance travelled in upstream and downstream directions did not differ significantly and there was no correlation between distance travelled and crayfish size. Several high flow events occurred during the study, but these did not cause any mortality or apparent displacement of crayfish downstream, suggesting that this is not a significant factor in downstream dispersal or mortality of adults of this invasive crayfish species in winter. A marked reduction in large-scale movements occurred in mid-December which coincided with a decline in water temperature. There was a less distinct pattern in local activity which was strongly correlated with water temperature and varied before and after mid-December.     

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.     

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.

     

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.     

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.     

Self-organisation in streams: the relationship between movement behaviour and body condition in a headwater salamander

1. When movement behaviour is correlated with traits affecting fitness, it may affect population performance directly, independent of extrinsic habitat conditions. 2. In a previous 3‐year, capture–recapture study, upstream movement by the salamander Gyrinophilus porphyriticus compensated for low reproduction in the upper 500 m of Merrill Brook, a first‐order stream in northern New Hampshire (U.S.A.). This immigrant subsidy resulted from excess reproduction in the downstream section (i.e. the lower 500 m of stream length) and from consistently upstream‐biased movement by salamanders. Reproduction in the two stream sections was positively correlated with mean body condition. Using 6 years of capture–recapture data, this study examines whether the movement behaviour of G. porphyriticus is related to body condition and thereby directly influences mean body condition and reproduction in the two sections of Merrill Brook. 3. Upstream‐biased movement and greater mean body condition in the downstream section were consistent across 6 years of data collection. In Merrill Brook and four other streams, however, individuals with high body condition were more likely to move upstream and low‐condition individuals were more likely to move downstream. Movement direction was unrelated to the size, sex and initial location of individuals. Body condition was positively related to growth rate, further supporting its link to reproductive potential, and positively autocorrelated through time in individuals that moved. 4. Results of this 6‐year study suggest that the movement behaviour of G. porphyriticus partially compensated for environmental factors differentiating mean body condition and reproduction along Merrill Brook and illustrate the potential for this form of self‐organisation to occur in linear habitats such as streams and rivers.     

Assessment of lake sturgeon spawning stocks using fixed-location, split-beam sonar technology

Fixed‐location, split‐beam sonar technology was used successfully to identify adult lake sturgeon Acipenser fulvescens as they moved upstream and downstream for spawning in the Sturgeon River, Michigan, May–June 2004. A Hydroacoustic Technology Inc. Model 241 Split‐Beam Echo Sounder operating at 200 kHz and a single 4 × 10° elliptical‐beam transducer with a near field range of 1.7 m set perpendicular to the river flow was used. Data collected from migrating lake sturgeon included direction of movement, swimming speed, range from transducer, time and date of passage, and target strength. The spawning population of lake sturgeon was estimated to be at 350–400 fish, with almost equal numbers of fish seen moving upstream as downstream. Most fish were recorded moving within the mid‐section of the river, 1.5–1.65 m deep, and swimming speeds upstream were slower than those for downstream moving fish. These results show that spilt‐beam sonar can be applied to lake sturgeon assessments, without the stress of actually handling these large, pre‐spawning fish.     

Adult philopatry and dispersal in the field vol Microtus agrestis

Summary Using mark-recapture data, we related the movements of adult field voles to population density, sex ratio and population growth. Dispersal movements (defined as distances larger than 1 home range diameter) were few in both sexes; 4 out of 197 (2.0%) in males and 8 of 316 (2.5%) in females. The distance moved between sequential trapping periods was similar for males and females; the mean being 10.2 m and 9.0 m respectively. Both males and females moved larger distances during the breeding season than during the nonbreeding period. The distance moved between sequential trapping periods showed a strong negative relation to density, i.e. both sexes moved shorter distances at higher densities, but there were no differences between periods of increasing and declining population densities. These results contradict the dispersal predictions of all major hypotheses proposed to explain population fluctuations in small mammals. The dispersal patterns fit a geometric distribution, suggesting that competition is the primary factor determining the dispersal characteristics of this population.     

Pattern of movements of adult Barbus haasi in a small Mediterranean stream

Most adult Barbus haasi , at a 1950 m-long site in Vallvidrera creek, were highly sedentary and resided within a home range <20 m (32 m²), while a small group were more mobile. On successive sampling occasions, between 52·3 and 64·9% of fish were recaptured in the same 10-m-long section in which they had previously been captured. Movements over long distances were infrequent, and only 5·6% of the fish moved >100 m. The movement pattern of the population was seasonally stable, although the mean distances were slightly greater in summer because of the medium-range movements of a few individuals. Overall, upstream and downstream movements were equally common but a significant downstream movement occurred in spring. The size of the fish did not influence the movement rate. Fish inhabiting those sections of the stream with greater depth, slower current and more cover had a lower movement rate than fish occupying shallower, exposed sections. The restricted movement of B. haasi could increase the survival of fish by increasing the probability of staying in the remaining pools during summer dry-out, and may therefore be of adaptive significance.     

Population genetic analysis of microsatellite variation of guppies (Poecilia reticulata) in Trinidad and Tobago: evidence for a dynamic source–sink metapopulation structure,founder events and population bottlenecks

Riverine fish populations are traditionally considered to be highly structured and subject to strong genetic drift. Here, we use microsatellites to analyse the population structure of the guppy (Poecilia reticulata), focussing on the headwater floodplain area of the Caroni drainage in Trinidad. We also analyse the population genetics of guppies in the Northern Drainage in Trinidad, a habitat characterized by rivers flowing directly into the sea, and a small isolated population in Tobago. Upland Caroni populations are highly differentiated and display low levels of genetic diversity. However, we found no evidence to suggest that these upland populations experienced recent population crashes and the populations appear to approach mutation–drift equilibrium. Dominant downstream migration over both short‐ and long‐time frames has a strong impact on the population genetics of lowland Caroni populations. This drainage system could be considered a source–sink metapopulation, with the tributary furthest downstream representing a ‘super sink’, receiving immigrants from rivers upstream in the drainage. Moreover, the effective population size in the lowlands is surprisingly low in comparison with the apparently large census population sizes.     

Natal dispersal in relation to population density and sex ratio in the field vole,Microtus agrestis

Summary In a sample of 240 juvenile field voles 8% of the males and 22% of the females reached sexual maturity within their natal home range. Among individuals retrapped as adults, 58% of males and 23% of females had dispersed, i.e. had moved more than one home range diameter. The mean distance moved for males (58.5 m) exceeded that for females (28.6 m). Male movement distances were negatively associated with total density, and with density of adult females, but not with male density. Female movements were not related to population density. There were no relation between sex ratio and distance moved. The distribution of distances moved for both males and females fit a geometrical distribution, suggesting the importance of competitive processes.     

Migratory movements of pacu, Piaractus mesopotamicus, in the highly impounded Paraná River

A mark‐recapture study was conducted in 1997–2005 to investigate movements of stocked pacu, Piaractus mesopotamicus, in the Paraná River Basin of Brazil, Paraguay, and Argentina. Fish raised in cages within the Itaipu Reservoir and in ponds were tagged externally (n = 2976) and released in the Itaipu Reservoir (53.2%) and bays of its major tributaries (46.8%). In total, 367 fish (12.3%) were recaptured. In all, 91% of the pacu moved away from the release site; upstream movements were more extensive than downstream movements. Pacu traveled upstream a maximum of 422 km (average of 41.3 km) at a maximum rate of 26.4 km day⁻¹ (av. 0.8). Downstream movements were limited in terms of number of individuals and distance moved. Fish released during the wet season moved farther than those released during the dry season, and feeding rather than spawning might have been the compelling reason for movement. Although fish passed downstream through dams, none of the marked fish were detected to have moved upstream through the passage facilities. Pacu showed movement patterns not radically different from those of other neotropical migratory species, but their migratory movements may not be as extensive as those of other large migratory species in the basin.     

The influence of tidal cycles and freshwater inflow on the distribution and movement of an estuarine resident fish Acanthopagrus butcheri

Movement patterns and habitat utilization by black bream Acanthopagrus butcheri (Sparidae), an estuarine resident species, were investigated using acoustic telemetry in a small estuary on the east coast of Tasmania, Australia. Thirty‐four adult A. butcheri were tracked for periods of up to 187 days between August 2005 and January 2006. Although able to tolerate a wide range of salinities, the fish spent most of the time within the upper and middle regions of the estuary, where brackish conditions dominated. The species exhibited extensive movements linked to tidal cycles, with small‐scale upstream movements during incoming tides and downstream movements during out going tides. The extent of these movements was positively correlated with the tidal height difference between consecutive tidal peaks and troughs. Freshwater inflows and resultant changes in salinity also significantly influenced distribution and movement patterns. Fish moved downstream during the periods of heavy inflows, returning upstream as salinities increased to c. >10. During the peak of spawning period (November to December) fish moved into the upper region of the estuary, where they aggregated to spawn. Periodic increases in freshwater discharge, however, resulted in fish leaving the spawning grounds and moving downstream. Towards the end of the spawning season (January), the fish became more dispersed throughout the entire estuarine system.     

Indicators of movement and space use for two co-occurring invasive crayfish species

Red swamp crayfish (Procambarus clarkii) and signal crayfish (Pacifastacus leniusculus) are two invasive freshwater species with a worldwide distribution. The objective of this work was to investigate how the two species move and use space in an area of recent coexistence. Simultaneously, we test the use of new tools and indices to describe their movement patterns. To accomplish this we performed a radio-tracking program within a river-type habitat during two different periods (September/October 2010 and June/July 2013). We used spatial analysis tools to map crayfish radio-location data with and without accounting for the curvature of the river. To assess the consistency of the direction of movement and of the distances traveled by crayfish, two indices were developed. To assess the habitat preferences of each species we applied Ivlev's Electivity Index and the Standardized Forage Ratio. Movement of P. clarkii and P. leniusculus differed. The average detected movement was 8.8 m day⁻¹ for P. clarkii and 17.5 m day⁻¹ for P. leniusculus. However, crayfish behavior ranged from almost complete immobility – sometimes during several days – to large movements, in half a day, up to a maximum of 255 m for P. clarkii and 461 m for P. leniusculus. The proportion of upstream or downstream movements was independent of the species and both species displayed no preference for either direction. The indices of consistency of movement showed a large interindividual variation. Species and period (2010 or 2013) affected the mean daily distance traveled, maximum observed distance from location of release and percentage of observations under vegetation cover. The Ivlev's Electivity Index and the Standardized Forage Ratio presented similar results. P. clarkii showed a preference for pool areas with riparian vegetation cover while P. leniusculus preferred riffle and pool areas with riparian vegetation cover. Our work provided new and valuable data for modeling the active dispersal of these two problematic invaders in a context of coexistence.     

Distribution, movements, and mortality of anadromous arctic char, Salvelinus alpinus L., in the Cumberland Sound area of Baffin Island

During 1972 the downstream movement of anadromous artic char, Salvelinus alpinus L., from two rivers flowing into Cumberland Sound, Baffin Island, began during the middle of May and was completed within 2 weeks. This movement took place during both the day and night. Upon reaching saltwater, many char older than 9 years (longer than 20 cm) began to migrate along the shore of the Sound at a rate of 0–6–0-9 km/day with the result that the average maximum distance travelled from the natal river was 40–50 km. Char 6–9 years old, 10–20 cm in length, remained much closer to the rivers while those younger than 5 years never entered saltwater. During the period of saltwater residence, the fish frequently moved into the intertidal zone and freshwater. Maturing fish of both sexes were occasionally found in saltwater. The upstream migration to the lakes began during the second week of August and was completed within 5–6 weeks. Females tended to ascend the rivers before males, although fish of all different size and age migrated upstream simultaneously. Migration occurred during both the day and night. The maximum distance anadromous char moved from saltwater was 40 km. The char were found only in freshwater between May and September and were normally located near the bottom of lakes at a depth of 15–40 m. During this period, large scale movements were probably very limited. Average annual mortality was estimated at 16.0% per year with the highest rates (25–30%) occurring at ages 10 and 15–17 years. Many fish probably died of old age but physical deterioration during and after spawning was also an important cause of death.     

Dispersal and upstream migration of an amphidromous neritid snail: implications for restoring migratory pathways in tropical streams

1. The amphidromous life cycle of several species of neritid snails, shrimp and gobies throughout the tropics includes juveniles that migrate from the ocean to breed in fresh water. In many Hawaiian streams, the decline of Neritina granosa, an endemic gastropod, has been associated with habitat degradation and water withdrawal, which are common factors affecting tropical rivers around the world. 2. We investigated the effects of water withdrawal and density on dispersal and upstream migration of N. granosa using three experimental treatments: (i) reduced flow (RF) owing to a stream diversion, (ii) natural flow (NF) and (iii) natural flow with artificially increased snail density. For each treatment, snails were differentially tagged and released in a stream without a natural, extant population of N. granosa. 3. Capture rates ranged from 17 to 65% over a 63‐day period following release. Captures on 2–6 days after release measured initial dispersal and migration, whereas longer‐term migration rates were calculated from snails captured 16–63 days after release. Snails under NF displayed positive rheotactic behaviour, with only 3–12% demonstrating initial downstream movement. Under RF, 22–77% of snails moved downstream or showed no bias either way. 4. Initial mean upstream migration rate (UMR) was 0.25, 0.66 and 1.16 m day^?1 under RF, NF and natural flow with increased snail density, respectively. Longer‐term migration rates did not differ significantly between treatments, and the overall mean was 0.62 m day^?1. 5. Principal component analysis and generalised linear models were used to identify habitat characteristics important to UMR, with habitat and reach‐scale hydraulics as the most important factors. The relationship between discharge and UMR suggested it would take 11–35 years for snails to migrate past the most upstream water diversion. However, rates from published studies of neritid snail species migrating en masse or in long lines under natural situations suggested that N. granosa could migrate above stream diversions within 72 days–2.5 years (when in an aggregation) and 29 days–1.1 years (when following in long lines of individuals). 6. An understanding of upstream neritid snail migration can be used for the management and conservation of this and other migratory species in tropical streams.     

Fine scale gene flow and individual movements among subpopulations of Centrolene prosoblepon (Anura: Centrolenidae)

Dispersal capabilities determine and maintain local gene flow, and this has implications for population persistence and/or recolonization following environmental perturbations (natural or anthropogenic), disease outbreaks, or other demographic collapses. To predict recolonization and understand dispersal capacity in a stream-breeding frog, we examined individual movement patterns and gene flow among four subpopulations of the Neotropical glassfrog, Centrolene prosoblepon, at a mid-elevation cloud forest site at El Copé, Panama. We measured male movement directly during a two year mark-recapture study, and indirectly with gene flow estimates from mitochondrial DNA sequences (mtDNA). Individuals of this species showed strong site fidelity: over two years, male frogs in all four headwater streams moved very little (mean = 2.33 m; mode = 0 m). Nine individuals changed streams within one or two years, moving 675-1,108 m. For those males moving more than 10 m, movement was biased upstream (p < 0.001). Using mtDNA ND1 gene sequences, we quantified gene flow within and among headwater streams at two spatial scales: among headwater streams within two adjacent watersheds (2.5 km2) and among streams within a longitudinal gradient covering 5.0 km2. We found high gene flow among headwater streams (phi(ST) = 0.007, p = 0.325) but gene flow was more limited across greater distances (phi(CT) = 0.322, p = 0.065), even within the same drainage network. Lowland populations of C. prosoblepon potentially act as an important source of colonists for upland populations in this watershed.     

Migration and Movement Patterns of Green Sturgeon (Acipenser medirostris) in the Klamath and Trinity rivers, California, USA

Green sturgeon, Acipenser medirostris, movement and migration within the Klamath and Trinity rivers were assessed using radio and sonic telemetry. Sexually mature green sturgeon were captured with gillnets in the spring, as adults migrated upstream to spawn. In total, 49 green sturgeon were tagged with radio and/or sonic telemetry tags and tracked manually or with receiver arrays from 2002 to 2004. Tagged individuals exhibited four movement patterns: upstream spawning migration, spring outmigration to the ocean, or summer holding, and outmigration after summer holding. Spawning migrations occurred from April to June, as adults moved from the ocean upstream to spawning sites. Approximately 18% of adults, those not out mignation in the spring, made spring post-spawning outmigrations. The majority of adults, those not outmigrating in the spring, remained in discrete locations characterized as deep, low velocity pools for extended periods during the summer and early fall. Fall outmigration occurred when fish left summer holding locations, traveled rapidly downstream, and exited the river system. High river discharge due to the onset of winter rainstorms and freshets appear to be the key environmental cue instigating the fall outmigration.     

Homing behaviour of stream-dwelling brook charr following experimental displacement

Precise homing of 24 brook charr Salvelinus fontinalis (157–215 mm L_F), displaced from their site of capture in a natural stream, was rare both in fish displaced upstream and those displaced downstream. Most fish settled in preferred habitats (pool or glide) away from their home site and showed restricted movement up to the pre-spawning period. Partial return responses, however, were stronger in fish displaced downstream than in fish displaced upstream. Most fish displaced downstream showed directional upstream movement soon after displacement, whereas fish displaced upstream did not show directional upstream movement, suggesting that it is primarily olfactory cues from upstream locations, and not positive rheotaxis, which mediate the return response. Homing success was unrelated to distance of displacement, body size, or population density in the home site, but daily mobility increased with body size for fish displaced downstream. There was no evidence of switching between specific areas between night and day, nor of differences between daytime and night-time mobility, suggesting that diel movement cycles did not bias the evaluation of homing success. Site attachment in stream-dwelling brook charr may be characterized as persistent (undisturbed fish remain stationary over long periods), but readily disrupted by involuntary displacement.