Inverted sand dollars actively orient themselves in flow to increase likelihood of righting

The fact that sand dollars are often dislodged and inverted is an inescapable consequence of living at or slightly below the sediment–water interface. Once inverted, however, how do sand dollars effectively right themselves, given their small spines and stiff internal skeletons? Here, we examined the possibility that individuals of Mellita quinquiesperforata and Dendraster excentricus may take advantage of the interaction of their morphology and flow to increase the likelihood of righting. Based on flow tank observations, the critical velocity required to flip an inverted sand dollar varies with orientation and increases with test size. For both species, the critical velocity was lower when inverted sand dollars were oriented with the posterior margin facing directly downstream, compared with when the posterior margin was positioned in an upstream orientation. To test whether inverted sand dollars would actively rotate into a more advantageous position for flipping, we exposed inverted animals in three starting orientations – with their posterior edge directed upstream (the least favored position for flipping), perpendicular, and downstream to flow – to the minimum flow expected to induce flipping and compared their responses. Time‐lapse photography showed that regardless of initial orientation, within one hour, a majority of individuals of both species rotated into positions that were not statistically different from the downstream orientation (the most favored position for flipping). These results for D. excentricus were further confirmed in a field experiment. Taken together, these data suggest that inverted sand dollars are able to recognize flow direction and respond by modifying their orientation to maximize lift and drag for righting.     

Regulation of feeding mode by the sand dollar Dendraster excentricus in a shallow estuarine habitat

The sand dollar Dendraster excentricus is a facultative suspension feeder, which is relatively rare within the Order Clypeasteroida. Field studies of regulation of its feeding mode have been mostly conducted in exposed coastal habitats, where they are typically observed in an inclined position and oriented parallel to each other during suspension feeding. Physical (current speed and direction, reduction of drag and lift) and biological factors (increased efficiency of capture of particulates, and density) have been associated with regulation of its feeding mode in exposed coastal systems. We simultaneously measured the role of physical and biological factors in regulating the selection mode for feeding under varying tidal conditions in a shallow estuary in Baja California, Mexico. We used photographic records and direct sampling in fixed plots to determine the relationship between feeding behavior and environmental conditions. Current direction and speed, tidal level, density and content of organic matter in the water column and sediments were measured with respect to feeding mode (prone or inclined) and orientation relative to prevailing currents during spring and neap tides. Multiple regression analysis indicated that the percentage of inclined sand dollars was strongly and positively correlated only with tidal level at the densities found in the estuary (mean < 180 individuals m^− 2); there was no relationship with current velocity, density, and organic matter content of the water. The prone position, indicative of deposit feeding, was largely limited to low tidal levels. We used circular statistics to determine whether the orientation of inclined sand dollars was correlated with current direction and speed. Sand dollars were only oriented parallel to the prevailing currents during the strongest currents of spring tides (> 20 cm s^− 1). We did not observe the predominant oral:aboral configuration found in exposed coastal systems, which may be attributed to the relatively low densities of sand dollars in the exposed coastal environment (30-180 individuals m^− 2), compared to protected coastal habitats (up to 1000 individuals m^− 2). Our results suggest that regulation of the feeding mode of sand dollars in shallow and hydrodynamically complex estuarine systems differs from the feeding mode found in exposed coastal environments.     

Intraspecific density regulates positioning and feeding mode selection of the sand dollar Dendraster excentricus

Dendraster excentricus is a common sand dollar of nearshore benthic habitats along the west coast of North America, and has the ability to feed either on deposited or suspended food particles. Field surveys and manipulative experiments demonstrated that intraspecific density and sediment organic matter (SOM) content of sediments are among the factors that regulate the proportion of sand dollars that forage as deposit versus suspension feeders. High local density was associated with a lower proportion of deposit feeding animals in both field surveys and under controlled experimental conditions. Conversely, the proportion of deposit feeders was elevated in treatments in which SOM levels were subsidized, regardless of local density. These data fit Levinton's model of resource limitation in relation to deposit-and suspension-feeding communities, and expand the list of biological processes regulated through density dependence. Analyses of carbon stable isotope ratios (δ¹³C) of sand dollars and their potential sources of primary production suggest individuals rely primarily on suspended particulate organic carbon (POC) or drift macroalgae. Sediment organic matter was not a substantial source of carbon for most individuals. There was a significant inverse relationship between size and δ¹³C values; smaller individuals depended to a greater extent on macroalgae. There was no consistent relationship between isotopic ratios, feeding mode and density, which may be due to the high mobility of the species, their ability to respond rapidly to changing environmental conditions and the dynamic nature of their habitat. Our results suggest that biological interactions influence feeding mode of this species. This is a complementary mechanism to those described previously, in which physical factors such as flow and lift/washout have been shown to regulate sand dollar positioning.     

Giant scallop feeding and growth responses to flow

The relationship between ambient seawater flow velocity and growth of the giant scallop Placopecten magellanicus Gmelin is shown to be a reverse ramp function with growth inhibition at flow velocities of > 10–20 cm · s⁻¹. The mechanism of inhibition involves a reduction in ration as velocities around the scallop increase. In ambient flows which are sufficient to cause overloading of the scallop gill, the feeding/filtration rate is reduced by an unknown mechanism, possibly involving the mantle edge closing or a gill bypass mechanism operating. In ambient flows where the pressure at the exhalant opening exceeds the inhalant plus the pressure head created by the gill, as when the scallop is placed dorsal edge to the flow, the tendency for flow reversal is resisted by a similar mechanism involving a reduction in feeding/filtration rate.     

Instream habitat use by blue duck (Hymenolaimus malacorhynchos) in a New Zealand river

1. The feeding habitat of a river specialist, blue duck (Hymenolaimus malacorhynchos (Gmelin 1789): Anatidae), was characterized in terms of water depth and velocity on eight occasions over a 13-month period in a river in the central North Island of New Zealand using video to record activity and relocate feeding sites. 2. Of the five feeding activities identified (‘pecking’, ‘grazing’, ‘head-dipping’, up-ending’ and ‘diving’), adult blue duck used mostly head-dipping (> 60% of feeding events on all dates), although diving or grazing from submerged surfaces of exposed boulders comprised major proportions of feeding behaviour (up to 33%) on occasions. Variations in feeding behaviour between dates partly reflected changes in antecedent flow conditions and the annual cycle of the birds. 3. Grazing and diving occurred in significantly faster water (mostly 0.3–0.45 m s^–1) and at significantly different depths (mean = 0.10 and 0.55 m, respectively) than head-dipping (0.20 m depth and 0.28 m s^–1 velocity). Adult feeding depths and velocities at four sites on different dates averaged 0.20 m and 0.31 m s^–1, respectively. Most feeding by 3–4-week-old ducklings occurred over a similar distribution of water velocities to adults but over a wider range of depths. 4. Adult birds fed in significantly shallower and lower velocity water than was available on the two dates that comparisons could be made. Ducklings also fed over a slower range of water velocities but were not selective in terms of water depth. 5. Energetically more expensive search methods were employed at times of high apparent energy demand to access flow microhabitats where larger bodied prey were more likely to be encountered. 6. These data indicate that, like other aquatic organisms, river birds can be influenced by basic hydraulic elements of river flow, but show at the same time that adult blue duck can accommodate variable lotic environments efficiently.     

Stream insects as passive suspension feeders: effects of velocity and food concentration on feeding performance

Benthic suspension feeders are important components of aquatic ecosystems, often dominating the use of space and influencing patterns of material cycling between the water column and benthos. Biomechanical theory predicts that feeding by these consumers is governed by the flux (i.e., product of food concentration and velocity) of particulate material to their feeding appendages. We performed a laboratory flume experiment to test how feeding by larval black flies (Simulium vittatum Zett.) responds to independent manipulations of flow and food concentration. We quantified larval body posture, flick rate of the labral fans, and ingestion rate as a function of two concentrations of a baker's yeast/chalk suspension (0.96 and 4.44 mg l^-1) and five water velocities (20, 30, 45, 60, and 90 cm s^-1). Using analysis of covariance, we found that both flick rate and ingestion rate increased in a decelerating manner with increasing velocity, while fan height decreased linearly with increasing velocity. In contrast, food concentration had no effect on any aspect of feeding behavior. Thus, although both velocity and food concentration contribute to particle flux, our results indicate that the two were not substitutable under the range of conditions tested here.     

The effect of water temperature and velocity on barnacle growth: Quantifying the impact of multiple environmental stressors

Organisms employ a wide array of physiological and behavioral responses in an effort to endure stressful environmental conditions. For many marine invertebrates, physiological and/or behavioral performance is dependent on physical conditions in the fluid environment. Although factors such as water temperature and velocity can elicit changes in respiration and feeding, the manner in which these processes integrate to shape growth remains unclear. In a growth experiment, juvenile barnacles (Balanus glandula) were raised in dockside, once-through flow chambers at water velocities of 2 versus 19 cm s⁻¹ and temperatures of 11.5 versus 14 °C. Over 37 days, growth rates (i.e., shell basal area) increased with faster water velocities and higher temperatures. Barnacles at high flows had shorter feeding appendages (i.e., cirri), suggesting that growth patterns are unlikely related to plastic responses in cirral length. A separate experiment in the field confirmed patterns of temperature- and flow-dependent growth over 41 days. Outplanted juvenile barnacles exposed to the faster water velocities (32±1 and 34±1 cm s⁻¹; mean±SE) and warm temperatures (16.81±0.05 °C) experienced higher growth compared to individuals at low velocities (1±1 cm s⁻¹) and temperatures (13.67±0.02 °C). Growth data were consistent with estimates from a simple energy budget model based on previously measured feeding and respiration response curves that predicted peak growth at moderate temperatures (15 °C) and velocities (20–30 cm s⁻¹). Low growth is expected at both low and high velocities due to lower encounter rates with suspended food particles and lower capture efficiencies respectively. At high temperatures, growth is likely limited by high metabolic costs, whereas slow growth at low temperatures may be a consequence of low oxygen availability and/or slow cirral beating and low feeding rates. Moreover, these results advocate for approaches that consider the combined effects of multiple stressors and suggest that both increases and decreases in temperature or flow impact barnacle growth, but through different physiological and behavioral mechanisms.     

Estimation of preferred water flow parameters for four species of Simulium (Diptera: Simuliidae) in small clear streams in South Africa

Blackfly larvae typically occur in fast-flowing riffle sections of rivers, with different blackfly species showing preferences for different hydraulic conditions. Very little quantitative data exist on hydraulic conditions linked to the blackfly species occurring in South African streams. Stones-in-current biotopes (i.e. fast riffle flows over cobbles) were sampled from four sites in three small clear streams in the Eastern and Western Cape provinces of South Africa. Mean water column velocities at each sampled stone were measured using a mini current meter, while flow velocities closer to the boundary layer where blackfly larvae occurred were estimated using indirect techniques (standard hemispheres and aerating tablets). Standard hemispheres were also used to calculate more complex hydraulic parameters such as Froude and Reynolds numbers. Four species of Simuliid were sampled in sufficient numbers to show trends in flow velocity preferences. Simulium impukane and S. rutherfoordi both occurred at their highest densities at velocities of 0.3m s^?1, while S. merops preferred velocities of 0.7m s^?1. Simulium nigritarse SL attained the highest densities of all the blackfly species sampled, and its relative abundances were greatest at velocities of 0.8–0.9m s^?1. Within the streams surveyed, all blackfly species occurred in subcritical-turbulent flows — based on a classification using Froude and Reynolds numbers — although two of the species were also found in high densities in supercritical flows where these existed at the sites. Local hydraulics within the stones-in-current biotope are complex, but in the absence of fine-scale equipment for measuring micro-velocities, standard hemispheres are a useful, cost-effective technique for the initial quantification of hydraulic parameters in small, clear streams. Such an approach facilitates further understanding of links between hydraulics and aquatic invertebrates in South African streams.     

Applying flow tank measurements to the surf zone: Predicting dislodgment of the Gigartinaceae

Our understanding of how flow affects the survival of members of the Gigartinaceae has advanced considerably in recent years. In particular, methods have been developed that give phycologists a powerful tool with which to predict mechanistically the survival of wave-swept macroalgae. One limitation of such methods, however, is the lack of hydrodynamic data for macroalgae in surf-zone flows. The method instead relies upon the extrapolation of low-velocity flow tank measurements (< 3.5 m s^–1) to velocities well beyond their measured range (10–20 m s^–1). Such extrapolation is particularly tenuous for flexible organisms, such as macroalgae, that can reconfigure in flow. This study uses three members of the Gigartinaceae to evaluate two commonly used methods for extrapolation of flow tank data to high water velocities: one that allows for continued thallus reconfiguration at extreme water velocities and one that does not. Limitations of both models are evident and underscore the need for improved measures of force generation on macroalgae exposed to extreme flows. Additionally, the extent to which the mechanical design might influence life history traits of the Gigartinaceae is discussed.     

A versatile laboratory stream with examples of its use in the investigation of invertebrate behaviour

A versatile and simple laboratory stream was designed and used to investigate the burrowing activity of two insects in response to changes in water velocity and substrate type.Aphelocheirus aestivalis adults were unable to burrow into sand, however, a small proportion of juveniles did burrow in this substrate. The presence of sand in gravel reduced the burrowing success of adults. Steady increases in flow stimulated the burrowing response of both adults and juveniles on gravel and sandy gravel.Ephemera danica was unable to burrow in sand alone at the velocities used in the experiment. The presence of particles greater than 2 mm in diameter in the substrate appeared to be essential for successful burrowing under the test conditions. An increase in flow from 3 to 8 cm s^–1 resulted in an increase in burrowing. The time taken for each specimen to burrow varied widely within replicate tests but most specimens had penetrated the substrate within 150 seconds of introduction. The implications of these observations in influencing the microdistribution of these species are discussed.     

The ectosymbiont crab Dissodactylus mellitae-sand dollar Mellita isometra relationship

The presence of the ectosymbiont Dissodactylus mellitae on Mellita isometra was studied at Tybee Island in Georgia. The number of crabs found on sand dollars, stage of maturity, sex, numbers of eggs, and size of eggs produced by crabs were noted. The test diameter of sand dollars, and the number and diameter of eggs produced were also noted. Infestation rates of crabs on sand dollars varied over time. The maximum number of crabs found on a sand dollar was 10. The three types of population dispersion, random, uniform, and clumped, were observed for crabs on sand dollars throughout the sampling period. Clumped or gregarious settling was observed when juvenile crabs were abundant, uniform and random distribution when mature crabs were abundant. Female crabs were significantly larger than male crabs, with carapace width of the largest mature female crab being 4.6 mm and the largest male 3.1 mm. Crabs produced between 80 and 300 eggs from 0.188 to 0.291 mm in diameter. Bigger crabs produced significantly more eggs than smaller crabs. Sand dollar sizes were 50-110 mm, with a mode between 60 and 70 mm. The effect of crab burden on egg production in the sand dollar was time-dependent. The presence of crabs on sand dollars correlated with total egg production of sand dollars in May, the peak of the spawning season, with sand dollars carrying one or two crabs having a lower egg production than those without crabs. Overall, variation in egg size was attributed to variation among females followed by variation between seasons and crab burden.     

Investigation of the effects of current velocity on mussel feeding and mussel bed stability using an annular flume

An annular flume was used to measure the effect of increasing current velocity on mussel (Mytilus edulis) feeding rate and the stability of mussel beds sampled from the mouth of the Exe estuary (SW England). It was found that, in contrast to earlier flume studies, the feeding rates of mussels from open coast sites were unaffected by current velocities up to 0.8 m s^–1. Algal cell depletion in the water column above mussels was a function of current velocity, increasing with declining currents below 0.05 m s^–1. The erodability/stability of the mussel bed, measured in terms of critical erosion velocity, sediment mass eroded and mean erosion rate, was found to be a function of the nature of the substrate and the density of the mussels. Erosion of mussel beds on sandy substrate showed a non-linear relationship with mussel bed density. In comparison with the sand (0% mussel cover), sediment resuspension was about five and four times higher for 25% and 50% cover, respectively. This was due to the increased turbulence and scouring around the clumps of mussels in low-density parts of the bed, and this resulted in some mussels detaching from the bed. At ~100% mussel cover, the sandy bed was more protected by the dense surface layer of mussels, and none became detached during erosion due to the high number of byssal attachments between individuals. The sediment resuspension from the 100% mussel cover was about three times lower than the 0% cover. Erosion of the bed with 50% cover resulted in burial of a large proportion of the mussels, with a 6 cm increase in sediment level. However, the mussels returned to the surface and recovered in 1–2 days, due to a combination of migration upwards and substrate settlement. Channels on the edge of the main Exmouth mussel bed were characterised by a more stable substrate comprising pebbles and sand with varying mussel densities. At these sites, where mussels experience high current velocities on spring tides (up to 0.9 m s^–1), there was no difference between the erodability of pebble/sand substrate with 0% and 100% mussel cover. The sediment erosion was also lower than the 100% mussel cover on the sandy substrate, particularly at currents >0.4 m s^–1. Sampling of different parts of the mussel bed at Exmouth showed mussels at low densities were made up of smaller clumps with a lower mass ratio of mussels to attached substrate (pebbles/sand), thus providing a greater degree of anchorage. Electronic Publication     

Swimming ability and burrowing time of two cirolanid isopods from different levels of exposed sandy beaches

Most of the macroinfauna from sandy beaches is highly mobile, emerging out of the sediment when the tide rises, and using the swash to migrate up and down the beach face or feed (searching for prey or carrion). After swash excursions, they usually burrow back into the sediment, maintaining zonation at low tide. Therefore, the different species abilities to emerge, move around and burrow under different swash climates and sediment conditions are expected to influence observed distribution patterns. Nonetheless, few attempts have been made to understand behavioral mechanisms of these organisms in moving fluids.In this study, we used a flume tunnel to investigate the orientation, swimming ability and burrowing time of two similar species of cirolanid isopods, Excirolana armata Dana and Excirolana braziliensis Richardson, under current velocities ranging from 5 to 30 cm·s⁻¹. E. armata inhabits middle levels of dissipative to intermediate beaches, while E. braziliensis is found towards the upper level of a wider range of beach states. Both species oriented countercurrent above a threshold velocity, which turned out to be significantly lower for E. armata than for E. braziliensis. E. armata proved to be a stronger swimmer as shown by the higher velocities surmounted, and the less drags experienced at the highest current velocity. E. armata also burrowed faster than E. braziliensis. Burrowing time was affected by sediment grain size and water content, but not by water flow. Once organisms managed to begin burrowing under different flow conditions, they were not affected by current velocity. Nonsaturated sand precluded burial, while coarse sand retarded it. Differences in the observed patterns of across-beach distribution may thus be the result of species-specific behavioral responses to swash climate, manifested in swimming ability, burying and orientation in directional flows.     

Water velocities around water plants in chalk streams

Water velocities were measured around water plants in a chalk stream. Low velocities (<100 mm.s^?1) were associated with stands of plants while faster flowing water (>500 mm.s^?1) was found in open water and above weed beds. An indirect method of estimating mean water velocity through weed beds, using discharge measurements, revealed velocities between 40 and 140 mm.s^?1, while open water velocities ranged between 200 and 370 mm.s^?1. The significance of the velocities is discussed.     

Evaluation of swimming performance for fish passage of longnose dace Rhinichthys cataractae using an experimental flume

The swimming performance of longnose dace Rhinichthys cataractae, the most widely distributed minnow (Cyprinidae) in North America, was assessed in relation to potential passage barriers. The study estimated passage success, maximum ascent distances and maximum sprint speed in an open‐channel flume over a range of water velocities and temperatures (10·7, 15·3 and 19·3° C). Rhinichthys cataractae had high passage success (95%) in a 9·2 m flume section at mean test velocities of 39 and 64 cm s^–1, but success rate dropped to 66% at 78 cm s^–1. Only 20% of fish were able to ascend a 2·7 m section with a mean velocity of 122 cm s^–1. Rhinichthys cataractae actively selected low‐velocity pathways located along the bottom and corners of the flume at all test velocities and adopted position‐holding behaviour at higher water velocities. Mean volitional sprint speed was 174 cm s^–1 when fish volitionally sprinted in areas of high water velocities. Swimming performance generally increased with water temperature and fish length. Based on these results, fishways with mean velocities <64 cm s^–1 should allow passage of most R. cataractae. Water velocities >100 cm s^–1 within structures should be limited to short distance (<1 m) and structures with velocities ≥158 cm s^–1 would probably represent movement barriers. Study results highlighted the advantages of evaluating a multitude of swimming performance metrics in an open‐channel flume, which can simulate the hydraulic features of fishways and allow for behavioural observations that can facilitate the design of effective passage structures.     

Secondary settlement of cockles Cerastoderma edule as a function of current velocity and substratum: a flume study with benthic juveniles

Some newly-settled bivalve molluscs can experience a second dispersal stage in the water column and colonize areas distant from initial settlement zone ('secondary settlement'). To document mechanisms involved in such a process, experiments were conducted in a recirculating 13-m flume, using juvenile cockles Cerastoderma edule (shell length between 0.8 and 5.7 mm). After 4 h and under current surface velocities of 10, 20 and 24 cm s^?1, all juveniles left the plexiglass substratum (site of initial introduction) and 42.4, 58.6 and 76.2% of juveniles, respectively, were retrieved from a downstream sand area (which only represented 7.7% of the total flume surface). Naked-eye observations showed that smaller individuals were borne within the water column, whereas larger individuals tended to roll or slip on the substratum. Byssus threads produced by the juveniles were often seen, sometimes covered in sand grains. These observations were confirmed by finding larger juveniles in the upstream part of the sand area. When sand was replaced by mud, a reduced proportion of cockles was retrieved in the experimental substratum (18.4, 20.0 and 16.4%, respectively). Observations showed that juveniles rarely succeeded in anchoring themselves in mud. When initially introduced on a favourable substratum (medium sand), more than 87% of juveniles were retrieved from that sand array at all flow velocities. This study shows that secondary settlement occurs for juvenile cockles up to 5.7 mm in shell length and depends not only on flow velocity but on substratum type.     

The flow velocity as driving force for decomposition of leaves and twigs

The water velocity and associated physical forces affect the distribution of organisms in rivers and streams; changes in the flows can also affect the organic matter decomposition and transport of detritus in lotic systems. The main objective of this study was to evaluate (120 days) the effects from the water flow velocity on the mass loss of leaves and stems in artificial channels (at 24 ± 1.0°C; current velocities: 0, 5, and 10 cm s^?1). The kinetic model used suggested that in addition to abrasion, the velocity of flow stressed the detritus fibers altering the decomposition pathways. Overall, the changes in the kinetic parameters varied linearly with the increase in flow velocity. Weight losses of the leaves were more affected by the flow velocity than those of the twigs. However, abrasion was more effective in the twigs than that in the leaves detritus. In the lotic systems, the leaves have greater importance in the detritus chains in the short term and in the conditions of backwater. The boost of flow velocity tends to change the refractory fractions (RPOM) into more labile/soluble compounds and increase the values of RPOM rate constant, owing to abrasion.     

The response of invertebrates to a gradient of flow reduction – an instream channel study in a New Zealand lowland river

1. The impacts of low flows on invertebrates were assessed in six experimental channels placed in a lowland stream. Each channel consisted of replicate stacked gravel‐filled baskets (buried 0–7, 8–14 and 15–21 cm deep) from which invertebrates were sampled. Samples were collected before, and 1 and 2 months after flows were reduced by varying amounts. We predicted that invertebrate communities would change most with the lowest flows that persisted longest. The effects of reduced flow on channel and intragravel‐velocities, temperature, dissolved oxygen and algal biomass were also monitored. 2. Mean velocity (17 cm s⁻¹) and depth (20 cm) were similar in all channels prior to flow reduction, but were reduced in five channels by 25% to 98%, a sixth channel serving as a control. Mean intragavel velocity prior to flow reduction (1.7 mm s⁻¹) was also reduced from 17% to 51%. Flow reduction had no effect on temperature, which ranged from 11.2 °C (night) to 19.6 °C (day). Dissolved oxygen was generally high (mean = 94% saturation) but variable (range from 50% to 145%), and did not differ between channels after flow reduction. Periphyton biomass (chlorophyll a and AFDM_algae) increased over time and was positively related to velocity after 1 month of flow reduction. After 2 months, however, this relationship reversed, with higher chlorophyll and AFDM_algae in lower velocity channels. Organic matter biomass (AFDM_total) was significantly higher in channels with lower velocities, and did not change with time. There was no relationship between AFDM_total and velocity in baskets from different depths. 3. Greater than 85% of animals were found in the upper baskets (0–7 cm deep), and flow reduction had no influence on the vertical distribution of invertebrates. After 1 month, invertebrate density had declined roughly in proportion to the magnitude of flow reduction in all channels where flows were reduced more than 25%. Densities recovered to pre‐reduction levels within 2 months. The amphipod, Paracalliope fluviatilis dominated all channels numerically prior to flow reduction, but its density declined markedly during the study. Densities of Ostracoda, Oxyethira albiceps and Cladocera increased dramatically after 2 months of flow reduction, especially in the lowest flow channels. Of the physical variables measured, chlorophyll a biomass and discharge best explained the temporal changes in the invertebrate community. 4. The effects of increased duration and magnitude of flow reduction on invertebrate communities were restricted to changes in the relative abundances of just a few taxa. Our results suggest that invertebrates in this lowland stream were resistant to the experimental flow reduction, presumably because of their broad ecological tolerances. We also found that more prolonged low flows did not result in predictable changes. This finding may have implications in terms of using hydraulic‐habitat models to set minimum flows in lowland streams if invertebrates can persist at flows much lower than their supposed optima.     

Effect of relative water motion on photosynthetic rate of red alga Gracitaria conferta

Bulk water velocities and local relative velocities generated in experimental tanks around and within thalli of free moving Gracilaria conferta were estimated according to the dissolution rate of benzoic acid sticks. Boundary-layer thickness and HCO ₃ ^– -mass-transfer coefficient were derived from the water velocities. Average relative velocities varied between 12 cm s ^–1 to less than 0.1 cm s ^–1 as a function of the absolute water flow in the tank, alga shape and location within the thallus. The lower range of velocities was observed at 20% of maximum aeration in the inner part of the plant. In laboratory experiments, photosynthetic rates, as determined in a closed Clark-type O₂-electrode system, increased by 30%–50% when water velocity was increased from zero to about 1.5 cm s ^–1. Another minor increase was obtained between 1.5 cm s ^–1 and 8 cm s ^–1 water velocity. This response to water motion was affected by bulk inorganic carbon concentration and by plant condition, as was reflected from the differences in the response in the winter and spring. It might be suggested that under carbon saturation, water velocity above 2 cm s^–1 provided almost sufficient flow to saturate carbon uptake.     

EFFECTS OF SUBSTRATE RELIEF ON THE DISTRIBUTION OF PERIPHYTON IN LABORATORY STREAMS,I. HYDROLOGY1

We examined the hypothesis that the heterogeneity of epilithic algal assemblages in streams may be partly a result of hydrologic differences created when water flows over a rough substrate. A 32-day experiment was conducted in laboratory streams that contained either 22.5 × 22.5 × 4 cm or 7.5 × 22.5 × 4 cm tile blocks. Free water velocities in the streams overaged 28 cm·s^?1. Hydrologic parameters and algal assemblages associated with surfaces on top of blocks and with recessed surfaces between blocks were compared to corresponding surfaces in streams with of relief. In streams with blocks, shear velocities averaged 1.7 cm·s^?1 on the top of blocks and 0.8 cm·s^?1 in the recessed areas. Shear velocity at corresponding surfaces in the control (no relief) streams averaged 1.9 cm·s^?1 and exhibited little variation. The hydrologic differences created by the larger blocks significantly affected the distribution of algal biomass, with recessed areas having an average of 2.6 g·m^?2 AFDW more biomass than surfaces on the top of blocks. Differences in shear velocities and biomass accumulation between top and recessed areas for the smaller blocks were less than for large blocks. Successional changes on all substrates were similar with the exception that recessed surfaces had a significantly greater abundance of the filamentous chlorophyte Stigeoclonium tenue (Ag.) Kütz after day 16. The results suggest that in cobble riffle areas of natural streams, the interaction between current flow and substrate relief has the potential to create patches of algae which are different in biomass and taxonomic composition.