Upstream-downstream movements of aquatic invertebrates in a Rocky Mountain stream

Simultaneous collections of drift and organisms moving either upstream or downstream in association with the substrate were made using a specially designed sampler. Samples were taken in a diel series along a transect across the study riffle of a Colorado foothills stream on six dates over an annual cycle. In addition to longitudinal movements, taxonomic composition and diel periodicity were evaluated. The insect-dominated fauna showed a net downstream displacement. Only the caddisflies Helicopsyche borealis and Hesperophylax occidentalis exhibited net upstream movement, primarily a result of low drift frequencies. The taxonomic composition of moving invertebrates differed from that of the benthos. Drift resembled downstream moving substrate-associated invertebrates in composition, but differed from that of the upstream directed fauna. Taxa collectively exhibited four types of diel patterns: 1) similar downstream (drift and substrate-associated movements) patterns, which generally differed from the upstream pattern; 2) similar benthic (upstream and downstream) patterns, which differed from that of drift; 3) aperiodic patterns; and 4) independent patterns for each type of directional movement. Analysis of size classes based on head capsule width for the mayfly Baetis tricaudatus showed significantly smaller size in stationary individuals compared with moving individuals in the population and revealed that nymphs moving during the day were smaller than those moving at night.     

Asymmetric dispersal structures a riverine metapopulation of the freshwater pearl mussel Margaritifera laevis

Unidirectional water flow results in the downstream‐biased, asymmetric dispersal of many riverine organisms. However, little is known of how asymmetric dispersal influences riverine population structure and dynamics, limiting our ability to properly manage riverine organisms. A metapopulation of the freshwater pearl mussel Margaritifera laevis may be sensitive to river currents because mussels are repeatedly exposed to downstream drift during floods—a parasitic life stage is the only, limited period (~40 days) during which larvae (glochidia) can move upstream with the aid of host fish. We hypothesized that water‐mediated dispersal would overwhelm upstream dispersal via host fish, and therefore, that upstream subpopulations play a critical role as immigrant sources. To test this hypothesis, we examined the effects of both up‐ and downstream immigrant sources on the size of target subpopulations in the Shubuto River system, Hokkaido, Japan. We found that target subpopulation size was dependent on the upstream distribution range of reproductive subpopulations and the number of upstream tributaries, which are proxies for the number of potential immigrants moving downstream. In contrast, little influence was observed of downstream immigrant sources (proximity to downstream reproductive subpopulations). These results were consistent even after accounting for local environments and stream size. Our finding suggests that upstream subpopulations can be disproportionately important as immigrant sources when dispersal is strongly asymmetric.     

Gene flow and genetic structure of the aquatic macrophyte Sparganium emersum in a linear unidirectional river

1. River systems offer special environments for the dispersal of aquatic plants because of the unidirectional (downstream) flow and linear arrangement of suitable habitats.
2. To examine the effect of this flow on microevolutionary processes in the unbranched bur-reed ( Sparganium emersum ) we studied the genetic variation within and among nine (sub)populations along a 103 km stretch of the Niers River (Germany–The Netherlands), using amplified fragment length polymorphisms.
3. Genetic diversity in S. emersum populations increased significantly downstream, suggesting an effect of flow on the pattern of intrapopulation genetic diversity.
4. Gene flow in the Niers River is asymmetrically bidirectional, with gene flow being approximately 3.5 times higher in a downstream direction. The observed asymmetry is probably caused by frequent hydrochoric dispersal towards downstream locations on the one hand, and sporadic zoochoric dispersal in an upstream direction on the other. The spread of vegetative propagules (leaf and stem fragments) is probably not an important mode of dispersal for S. emersum , suggesting that gene flow is mainly via seed dispersal. Realized dispersal distances exceeded 60 km, revealing a potential for long-distance dispersal in S. emersum .
5. There was no correlation between geographical and genetic distances among the nine S. emersum populations (i.e. no isolation by distance), which may be due to the occurrence of long-distance dispersal and/or colonization and extinction dynamics in the Niers River.
6. Overall, the genetic population structure and regional dispersal patterns of S. emersum in the Niers River are best explained by a linear metapopulation model. Our study shows that flow can exert a strong influence on population genetic processes of plants inhabiting stream systems.     

A model for accurate drift estimation in streams

1. This paper explores the experimental difficulties involved with the use of drift nets in small streams, and outlines a method whereby the estimation of drift density (number of specimens m⁻³ of water) can be improved.
2. Changes in the filtering efficiency of the net caused by trapping of organic debris ('clogging') has the effect of reducing net entrance velocities, causing errors in the calculation of sampled water volume, and thus drift density. A model of the reductions in net entrance velocity based on empirical measurements of trapped debris is developed.
3. Cross-sectional velocity calculations suggest that errors can also be introduced into drift density calculations by positioning sampling nets only on the bed. A method to allow this effect is demonstrated.
4. As adjustments to the calculation of sampled volume are required when sampling in rivers that undergo marked changes in discharge during the sampling period, a method whereby these effects can be accommodated to improve drift density estimations is also outlined.
5. The results of this study imply that theoretical links between flow hydraulics and short-term drift behaviour are poorly understood.     

A biomechanical hypothesis explaining upstream movements by the freshwater snail Elimia

1. Many taxa of freshwater invertebrates show active upstream movements, particularly the snails. Hypotheses explaining this behaviour invoke the search for food or space, compensation for drift, avoidance of predation and hydrodynamic effects. The pervasiveness of upstream movements among remote lineages of snails (two subclasses, three orders, 10 families), however, suggests that snails may move upstream for mechanical rather than adaptive reasons.
2. It is proposed that upstream movements by snails are a function of torque on the snail's foot generated by hydrodynamic drag on the shell. When subject to high broadside drag-forces on their shells, snails are able to reduce torque and stabilize orientation only by directing their anterior aspect upstream.
3. Movements of the freshwater pleurocerid snail Elimia were studied by following marked free-ranging individuals in six streams in Alabama, USA (four species, eight populations).
4. Populations showed either no net movement (two streams) or significant upstream movements ranging to a mean of ≈40 m over a 3-month period (four streams). Movement patterns were stream specific rather than species or population specific. Within populations showing significant upstream movements, snails with shell lengths ≤10 mm showed little net movement. Larger snails showed movements from 0 to >200 m upstream.
5. A torque-constrained random walk model was used to perform a post hoc test of the hypothesis that upstream movements were a function of torque on the snail's foot generated by hydrodynamic drag on the shell. The model predicted upstream and size-dependent movement patterns that approximated those observed for snails in the field.     

Differences in endozoochorous dispersal between aquatic plant species, with reference to plant population persistence in rivers

Summary 1. In river ecosystems, populations are continuously subjected to unidirectional downstream currents resulting in a downstream movement of populations. To ensure long-term population persistence in rivers, organisms must have a mechanism for upstream dispersal, which allows them to re-colonise upstream areas.
2. In this study we assessed differences in the potential for endozoochorous seed dispersal of Sparganium emersum and Sagittaria sagittifolia , two aquatic plant species with different seed morphologies, by mallard ( Anas platyrhynchos ) and teal ( Anas crecca ), two duck species with different body weights.
3. We found no significant differences in seed retrieval (the proportion of ingested seeds retrieved after gut passage) and seed retention time (time between seed ingestion and retrieval), between mallard and teal, despite the difference in body weights. We did find a significantly higher germination (%) over retention time of S. emersum seeds retrieved from teal compared with mallard, most likely related to a more efficient removal of the seed coat during passage through the gut of teal.
4. There were large differences between S. emersum versus S. sagittifolia in: (i) seed retrieval (22.65 ± 20.8% versus 1.60 ± 2.4%, respectively); (ii) seed retention time in duck gut, with a maximum of 60 h versus 12 h; (iii) the effect of gut passage on seed germination, with an increase of approximately 35% versus a decrease of 25%; and (iv) the effect of gut passage on seed germination rate, with an acceleration of 10 days versus a delay of 3 days on average. The results show that S. emersum has a higher potential for endozoochorous dispersal by ducks and postdispersal establishment than S. sagittifolia .
5. We propose that, in rivers, bird-mediated seed dispersal may promote re-colonisation of upstream areas, enabling long-term plant population persistence.     

A test of the hypothesis of compensatory upstream dispersal using a stream-dwelling waterstrider,Gerris remigis Say

Summary This paper reports the results of 2 experiments designed to determine if adult Gerris remigis (Hemiptera, heteroptera), a stream-dwelling waterstrider, tend to disperse preferentially upstream, as predicted by the colonization cycle hypothesis summarized by Muller (1982). Markrecapture observations and experimental removals were used to assess the distance and direction of movement of adult G. remigis along a small mountain stream, over a full year. These experiments indicated that adult G. remigis show a significant upstream bias in movement distance, but not in numbers of animals moving. This upstream bias is characteristic of pre-reproductive, sexually immature adults of both sexes, and occurs primarily in association with movements to and from diapause sites. Although the existence of a significant upstream bias in movement distances tends to support the colonization cycle hypothesis, the data from the removal experiments clearly show that upstream dispersal is not sufficient to compensate entirely for downstream drift.     

Flight activity of insects along a mountain stream: is directional flight adaptive?

1. Flight activity of Trichoptera, Plecoptera and Ephemeroptera was studied by sticky trapping for 12 months at five sites along a New Zealand mountain stream. Over 19 000 insects were captured by the traps, which were located in forest and grassland reaches, including a reach with intermittent flow.
2. Most species occurred predominantly in forest or grassland, although some were trapped throughout the stream. Longitudinal distributions of adults and their larvae were strongly correlated.
3. Flight periods of 24 caddisflies, three mayflies and four stoneflies ranged from 2 to 12 months. Six species were trapped in all months and 17 (55%) in more than 5 months.
4. The most abundant forest-dwelling caddisfly species were over-represented on the downstream sides of sticky traps located in, and immediately below, forest indicating a majority was flying upstream. Upstream flight compensates for downstream drift of larvae and should maximize the likelihood that forest-dwelling species will locate preferred habitat for egg, larval and/or adult development. Unlike the caddisflies, the stonefly Spaniocerca zelandica was over-represented on the upstream sides of traps, suggesting that some adults may float or fly downstream following emergence.
5. In contrast to forest-dwelling species, only one common caddisfly ( Oxyethira albiceps ) was over-represented on the downstream sides of traps at grassland sites. Unlike the forest-dwelling species, most species taken at the downstream sites probably came from a variety of sources, including a nearby stream.     

Tributary and mainstem benthic macroinvertebrate communities linked by direct dispersal and indirect habitat alteration

Benthic communities in tributary–mainstem networks might interact via downstream drift of invertebrates or material from tributaries and adult dispersal from the mainstem. Depending on the strength of these interactions, mainstem downstream communities are expected to be more similar to tributary communities due to drift or habitat alteration. Communities not connected by flow are expected to be similar due to adult dispersal but decreasing in similarity with distance from the mainstem. We investigated interactions between invertebrate communities of a 7th order river and 5th order tributary by comparing benthic community structure in the river upstream and downstream of the tributary confluence and upstream in the tributary. Non-metric multidimensional scaling showed invertebrate communities and habitat traits from river locations directly downstream of the tributary clustered tightly, intermediate between tributary and mid-channel river locations. In addition, Bray–Curtis dissimilarity increased between the mainstem and tributary with distance upstream in the tributary. Our results indicate that similarities between mainstem and tributary communities are potentially caused by direct mass effects from tributary to downstream mainstem communities by invertebrate drift and indirect mass effects by habitat restructuring via material delivery from the tributary, as well as potential effects of adult dispersal from the river on proximal tributary communities.     

Using passive sonic telemetry methods to evaluate dispersal and subsequent movements of hatchery-reared white sturgeon in the Kootenay River

A total of 35, age 1 juvenile Kootenay River white sturgeon ( Acipenser transmontanus ), were fitted with sonic tags in 2005 and released as part of larger hatchery release groups at five sites to evaluate dispersal and subsequent movements (seven tags per site). Juvenile sturgeon released at three locations within the deep, low gradient reach (typical gradient of 0.02 m km⁻¹ and velocities of <0.4 m s⁻¹) of the Kootenay River below Bonners Ferry, ID showed substantial dispersal both up and downstream; however, downstream redistribution was more common. White sturgeon from all three release locations overlapped during dispersal, with 9% of tagged fish moving from river release sites into Kootenay Lake. The three hatchery release locations in this low gradient reach produced good dispersal of hatchery progeny into available habitats. Tagged fish released above Bonners Ferry in the shallow, higher gradient reach (typical gradient of 0.6 m km⁻¹, and velocities >0.8 m s⁻¹) at two additional sites all moved downstream of the gradient break at Bonners Ferry, ID into the lower gradient reach within 2 months of release. In total, 93% of these tagged fish relocated to the low gradient section within 25 days of release, with some fish undertaking this movement within 1 day. In general, age 1 hatchery release juveniles were mobile and capable of substantial movements.     

Short-term movements of chalk-stream invertebrates

SUMMARY 1. Trays of gravel were implanted for 24 h periods in several chalk streams. The trays were variously modified to monitor colonization from different sources, particularly drift (D), other downstream movements (DSM), upstream movements (USM), and all sources combined (ALL).
2. The abundance of invertebrates in the streams was high (maximum of 717 per 0.04 m²) and colonization of the trays was extensive, especially by Gammarus pulex , the predominant species in all but one of the streams (maximum number of individuals of G. pulex per tray: for D, 473; for DSM, 462; tor USM, 286; and tor ALL, 639). In some cases benthic densities were achieved in the trays in the 24 h exposure period, and in most cases any one source of colonization could provide benthic densities in a few days.
3. Patterns of movement differed between species, between size classes of G. pulex , and between sites for particular species. No one source of colonization could be regarded as generally predominant although downstream movements in total (D + DSM) usually far exceeded USM. DSM was usually an important source of colonization.
4. The extent and direction of movement appeared to be determined by local factors and were explained in terms of the high densities of invertebrates and the current and substratum characteristics of each site.
5. Estimates of the relative importance of different sources of colonization varied greatly according to the criteria used in determining proportions.     

Genetic diversity and migration patterns of the aquatic macrophyte Potamogeton malaianus in a potamo-lacustrine system

1. Previously, the Yangtze River connected thousands of shallow lakes which together formed a potamo-lacustrine system capable of sustaining a rich variety of submerged macrophytes.
2.  Potamogeton malaianus is one of the dominant submerged macrophytes in many lakes of this area. Genetic variation and population structure of P. malaianus populations from ten lakes in the potamo-lacustrine system were assessed using inter-simple sequence repeat markers.
3. Twelve primer combinations produced a total of 166 unambiguous bands of which 117 (70.5%) were polymorphic. Potamogeton malaianus exhibited a moderate level of population genetic diversity ( P _P = 70.5%, H _E = 0.163 and I =  0.255), as compared with that of plants in the same habitat and range. The main factors responsible for this moderate value were the plant's mixed breeding system (both sexual and asexual) and the hydrological connectivity among habitats.
4.  F statistics, calculated using different approaches, consistently revealed a moderate genetic differentiation among populations, contributing about 20% of total genetic diversity. An estimate of gene flow (using F _ST) suggested that gene flow played a more important role than genetic drift in the current population genetic structure of P. malaianus ( Nm  = 1.131).
5. The genetic diversity of P. malaianus did not increase downstream. A high level of linkage–disequilibrium at the whole population level suggested that metapopulation processes may affect genetic structure. The migration pattern of P. malaianus was best explained by a two-dimensional stepping stone model, indicating that bird-mediated dispersal could greatly influence gene movements among lakes.     

Dispersal of organisms in a patchy stream environment under different settlement scenarios

1.  Previous work in streams, where many organisms disperse downstream by drifting in the water column, has suggested dispersal distances could be related to the overall proportion of the stream within dead water zones (DWZ), places such as backwaters, pools or behind large obstacles, where the water is out of the main flow and is stored temporarily. However, dispersal distances might also be influenced by the spatial organization of DWZ within a reach; this would require us to distinguish between reaches containing many small DWZ compared with those having a few large ones in order to understand fully differences in dispersal patterns among reaches differing in this regard.
2.  We constructed a spatial model of flow in stream reaches, in which we varied the density and size distribution of DWZ distributed through the reach. We defined flow conditions under which stream organisms, modelled as passive particles, would settle or continue to drift downstream. We then examined the horizontal dispersal profiles of such particles through reaches differing in arrays of DWZ.
3.  Our results support those of previous studies, but suggest that the spatial arrangement of DWZ may be just as important as the overall proportion of DWZ within a reach. On a more general note, the importance of patch configuration observed in this study reflects the growing view that a thorough understanding of population and ecosystem processes will require the explicit consideration of spatial pattern in the environment.     

Mortality of zebra mussel, Dreissena polymorpha, veligers during downstream transport

1. Streams flowing from lakes which contain zebra mussels, Dreissena polymorpha , provide apparently suitable habitats for mussel colonization and downstream range expansion, yet most such streams contain few adult mussels. We postulated that mussel veligers experience high mortality during dispersal via downstream transport. They tested this hypothesis in Christiana Creek, a lake-outlet stream in south-western Michigan, U.S.A., in which adult mussel density declined exponentially with distance downstream.
2. A staining technique using neutral red was developed and tested to distinguish quickly live and dead veligers. Live and dead veligers were distinguishable after an exposure of fresh samples to 13.3 mg L⁻⁻¹ of neutral red for 3 h.
3. Neutral red was used to determine the proportion of live veligers in samples taken longitudinally along Christiana Creek. The proportion of live veligers (mean ± SE) declined from 90 ± 3% at the lake outlet to 40 ± 8% 18 km downstream.
4. Veligers appear to be highly susceptible to damage by physical forces (e.g. shear), and therefore, mortality in turbulent streams could be an important mechanism limiting zebra mussel dispersal to downstream reaches. Predictions of zebra mussel spread and population growth should consider lake-stream linkages and high mortality in running waters.     

Do pools impede drift dispersal by stream insects?

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Three ways of assessing metapopulation structure in the butterfly Plebejus argus

1. Three independent methods were used to investigate population structure in the butterfly Plebejus argus . First, migration and dispersal ability were measured by mark–release–recapture in seven adjacent habitat patches, and by release of butterflies in unoccupied habitat. Secondly, colonization of newly created habitat was observed over 7 years. Finally, genetic differentiation of local populations within a metapopulation was investigated. Sampled local populations included parts of the mark–release–recapture study area.
2. Plebejus argus is relatively sedentary: the maximum movement detected was 395 m, and only 2% of individuals moved further than 100 m between recaptures on different days. None the less, adjacent local populations in the mark–release–recapture study area were linked by occasional migration, with ≈ 1.4% of individuals moving between patches separated by 13–200 m.
3. Despite low mobility, observed colonizations occurred rapidly over distances of 1 km. Because P. argus occurs at high population densities, 1.4% migration can generate enough migrants to colonize newly suitable habitat quickly at this spatial scale.
4. Mark–release–recapture data were used to predict that there would be limited genetic differentiation through drift between local populations at this spatial scale. The prediction was supported by allele frequency data for the same local populations.
5. Genetic differentiation often indicates higher levels of migration than are revealed by the movements of marked individuals. This study shows that when experimental releases and extensive marking are undertaken in areas that are large relative to most movements, indirect measures of gene flow and direct measures of dispersal can concur.
6. Evidence from the three different approaches was complementary, indicating that P. argus occurs as metapopulations within the study area.     

Movement patterns of Rhyssomatus lineaticollis Say (Coleoptera: Curculionidae) within and among Asclepias syriaca (Asclepiadaceae) patches in a fragmented landscape

Abstract.  1. Dispersal capabilities of organisms are critical in determining the landscape population structure of species as well as their likelihood of survival in fragmented landscapes. Using mark–recapture techniques on the monophagous weevil Rhyssomatus lineaticollis Say (Curculionidae), within- and between-patch dispersal capabilities, landscape level population structure, and the role of beetle density and host patch characteristics in setting distances, amounts, and timing of dispersal were studied.
2. The data indicate that R. lineaticollis is sedentary, with 50% of recaptured beetles moving < 1 m and the maximum distance moved < 1 km. Within- and between-patch movement of beetles was unrelated to host plant patch characteristics and beetle densities.
3. Despite limited dispersal, R. lineaticollis probably functions as a patchy population in east-central Iowa, U.S.A. because dispersals between patches are common and because all host patches surveyed contained this herbivore, indicating a lack of suitable vacant patches, a prerequisite for metapopulation structure.
4. Between-patch distances are well within the dispersal capabilities of R. lineaticollis , although this may be the result of an increase in the density of patches of its host, Asclepias syriaca , in the landscape over the last 150 years as a result of human disturbance and this species' weedy habit.
5. Metapopulation structure in monophagous prairie herbivores may be most likely in species whose non-weedy host plants form highly predictable resources in space and time, but which are now widely scattered in habitat fragments.     

Delivery of hatching larvae to estuaries by an amphidromous river shrimp: tests of hypotheses based on larval moulting and distribution

1.  Amphidromous shrimps live and breed in freshwater rivers and streams, but their larvae require development in sea water. Larvae may hatch upstream and then drift to the sea, although in some species females have been reported to migrate to the coast before larvae are released. Here, we tested the relative importance of larval drift and female migration in Macrobrachium ohione (Decapoda: Palaemonidae) in a distributary of the Mississippi River in Louisiana, U.S.A.
2.  Newly hatched (stage-1) larvae are nonfeeding and will not moult to stage 2 (first feeding stage) without encountering salt water. A factorial experiment was conducted in the laboratory to test the effects on moulting to stage 2 of (i) time spent by stage-1 larvae in fresh water before (ii) exposure to and maintenance in water of different salinity. Larvae kept in fresh water for 1 or 3 days before a change to saline water at 6 or 10 ppt showed a greater frequency of moulting than those kept for longer (5 days) in fresh water or changed to less saline water (2 ppt). Non-moulting larvae died or were moribund within 11 days of hatching.
3.  The relative abundance of stage-1 larvae was measured with plankton tows at two locations in the river c. 150 km apart, one near the sea and one upstream. Larval abundances near the sea were significantly greater than those upstream.
4.  The results indicate that hatched larvae of M. ohione have a limited period in which to drift in fresh water before reaching water sufficiently saline to stimulate moulting to the first feeding stage. Female migrations may play an important role in delivering larvae of amphidromous species from large continental river systems in which distances to the sea are great, while larval drift alone may be sufficient in species living in short streams, like those found in many small mountainous tropical islands.     

Detecting dispersal of Nuphar lutea in river corridors using microsatellite markers

1. River corridors are well-known for their role in plant dispersal. The buoyancy of seeds, the possibility of dispersal by vegetative fragments, and the frequency and efficiency of dispersal among different river catchments determine linear distribution patterns. Little is known about the relative importance of these factors to observed patterns of genetic variation.
2. One hundred and fifty-six Nuphar lutea individuals from forty-four sampling sites in the river catchments of the Cidlina River, the Mrlina River and the Labe River (Czech Republic) were studied using ten microsatellite markers. Interpretation of patterns in genetic variation allowed several conclusions about dispersal mechanisms.
3. Vegetative long-distance dispersal is probably very limited in this species. Only one multilocus genotype was found in more than one sampling site. The distance between the sites was about 75 km.
4. To explain the distribution of Bayesian based clusters of related multilocus genotypes, both along-river and inter-river long-distance dispersals have to be invoked.
5. A marginally significant tendency for higher genetic diversity in the lower part of the river Cidlina was detected. Continuous downstream dispersal of seeds by water currents could be a valid explanation.
6. Significant positive autocorrelation was found among individuals at within-river distances of up to 25 km. Repeated dispersal of seeds over distances in the range of tens of kilometers is common.     

The upstream/downstream movement paradox of lotic invertebrates: quantitative evidence from a Welsh mountain stream

• 1 This study attempts to determine the mechanisms by which the downstream displacement of lotic invertebrates is compensated by the upstream movements of immatures and adults. To this end, submerged and aerial traps were set up at three sites, each 100 m apart on a small mountain stream (Yr Ogof) in North Wales and operated for 12 months.
• 2 At Site 1 (the source), 19.3% of the benthic invertebrates lost as drift were replaced by the upstream movement of aquatic stages. At Site 2, the average upstream compensation was 51.1% (although there was a marked difference between the two traps at the site). At downstream Site 3 the average compensation was 55.4%. The net loss of benthos downstream over 1 year was estimated to be 261920 individuals from Site 1, 41891 from Site 2, and 40470 from Site 3.
• 3 At Sites 1 and 2, the numbers of invertebrates drifting were positively correlated with both current speed and water depth. At Sites 2 and 3, the numbers moving upstream were positively correlated with water temperature. At all three sites, the numbers of animals drifting were positively correlated with the numbers moving upstream, under water.
• 4 In none of the eight common species of stoneffy did females show a strong overall upstream flight preference—however, two species, Leuctra nigra and Nemoura erratica, showed a significant downstream preference.
• 5 As with the stonefly nymphs, most of the caddisfly larvae showed a longitudinal distribution. Agapetus fuscipes larvae were most abundant at Site 1, yet females showed no preferred flight direction—this would seem to be inappropriate for maintaining this larval distribution pattern. However, females caught moving upstream at Site 2 were bigger and contained twice as many eggs, which were also larger, than females moving upstream at Site 1. The flight patterns of the other Trichoptera also seemed inappropriate for maintaining their respective larval distributions. Immature insects moved upstream at certain sites and times, but trends were not consistent. There was no correlation between female flight direction and local wind speed and direction.
• 6 Downstream displacement of the most abundant non-insect species, Gammarus pulex, was quite considerable at all three sites. However, at several times during the year the numbers moving upstream approached or exceeded those drifting. Maintenance of benthic population densities in this species is thought to be through a combination of upstream movement (which was highly correlated with benthic density at Sites 1 and 3) and production of young over a long (10 month) reproductive period. Maintenance of the benthic populations of the immature stages of the majority of insect species in Yr Ogof is thought to be through oviposition by females in the vicinity of their emergence sites.