Predation and drift of lotic macroinvertebrates during colonization

Summary A field experiment was carried out to determine the effect of an invertebrate predator on the colonization and drift of benthic macroinvertebrates in experimental stream channels. Lotic invertebrates colonized four replicate channels: two controls with no predators, and two channels with low densities (2.8 m^–2) of predatory stonefly nymphs, Doroneuria baumanni (Perlidae). Immigration rates were measured at the inflow of two other channels. Drift rates of invertebrates immigrating to and emigrating from channels were measured daily, and benthic samples were collected every five days. Over a 25-day colonization period, benthic densities of Baetis nymphs and larval Chironomidae were reduced by D. baumanni. Colonization curves were fit with a power function and significantly different colonization rates were indicated for both Baetis and chironomids in predation and control channels. A predator-induced drift response was exhibited by Baetis only and this response was size-dependent. In the presence of D. baumanni, large Baetis drifted more frequently than small nymphs and, correspondingly, small nymphs were more frequent in the benthos. Net predator impacts on invertebrate densities in channel substrates were partitioned into predator-induced drift and prey consumption. These estimates suggest that predator avoidance by Baetis is a prominent mechanism causing density reductions in the presence of predators. Reductions in the density of Chironomidae, however, were attributed to prey consumption only. A rainstorm during the experiment demonstrated that stream flow disruptions can override the influence of predators on benthic invertebrates, at least temporarily, and re-set benthic densities.     

Macroinvertebrate drift in a Rocky Mountain stream

An extensive series of drift collections from a Rocky Mountain stream was used to investigate quantitative patterns in the taxonomic composition of drift throughout spring, summer and fall for 1975–1978. Drift was estimated by drift rate, the number of organisms drifting past a point per 24 h; and by drift density, the numbers of organisms collected per 100 m³ of water sampled.Drift densities were up to ten times greater by night than by day, and 24 h drift densities for the total fauna approached 2000 per 100 m³ in June–July, declining to <500 by autumn. Ephemeroptera, and especially Baetis, dominated the drift. Drift rates were greatest in late spring, around 10⁶ per 24 h, which are among the highest values reported for small trout streams. Drift rates declined to <10⁵ during the summer, and shifts in the taxonomic composition are described.Multiple regression analysis of the relationship between drift rate and density, and the independent variables discharge, benthic density and temperature, showed that discharge typically was a significant predictor of 24 h drift rate, usually the best single predictor. In contrast, 24 h drift density most frequently was independent of discharge, indicating that this measure tends to correct for seasonal variation in discharge, as suggested in the literature. However, this was not invariably true. Drift density significantly correlated with benthic density in five of eight taxa inspected, thus seasonal declines in the benthos probably accounted for parallel declines in drift density.     

The relation between invertebrate drift and two primary controls,discharge and benthic densities,in a large regulated river

相似文献   

Effects of single and repeated experimental acid pulses on invertebrates in a high altitude Sierra Nevada stream

1. We examined responses of aquatic macroinvertebrates to pulsed acidification experiments in twelve streamside channels located in the Sierra Nevada, California. Experiment 1 consisted of a single 8 h acid addition, and Experiment 2 consisted of two 8 h acid additions administered 2 weeks apart. Replicated treatments (four reps/ treatment) consisted of a control (pH 6.5–6.7) and pH levels of 5.1–5.2 and 4.4–4.6. Invertebrate drift was monitored continuously and benthic densities were determined before and after acid addition. 2. Drift responses to pH reduction were: (i) increased drift during acidification in pH 5.2 and pH 4.6 treatment channels, often with depressed post-acidification drift in treatment channels relative to controls (exhibited by Baetis only). Depressed post-acidification drift in treatment channels appeared to be due to low benthic densities because a positive relationship between benthic and drift densities was noted for most common taxa; (ii) increased drift rates during acidification only at pH 4.6 (Epeorus, Drunella, Paraleptophlebia, Zapada, and Simulium); (iii) decreased drift at pH 5.2 and/or pH 4.6 relative to control channels (Rhyacaphila and chironomid larvae); (iv) no significant response to acidification (Ameletus, Amiocentrus, Dixa and Hydroporus). 3. A high proportion (45–100%) of acid-induced drift in Baetis, Epeorus, and chironomid larvae could be attributed to dead, drifting individuals. 4. Except for chironomids, most common invertebrates (i.e. Baetis and Paraleptophlebia) showed reduced benthic densities in treatment relative to control channels after acidification. 5. For sensitive taxa, drift was enhanced and benthic densities reduced by single (Experiment 1) and initial [Experiment 2(a)] acid pulses. Drift responses to a second acid pulse [Experiment 2(b)] were not as pronounced as those to the single or initial acid pulses [Experiments 1 and 2(a)], and the second acid pulse had no additional effect on benthic density.     

Invertebrate drift in a large, braided New Zealand river

1. The spatio-temporal patterns of drifting macroinvertebrates in a large, braided New Zealand river were determined by sampling with drift nets, seasonally, for 1 year. 2. Drift densities were greatest in autumn, and at night in all seasons except winter. A greater proportion of larger animals drifted at night than during the day in all seasons. Mean annual drift densities were ninety-six animals 100m^?3 and 47 mg dry weight 100 m^?3. 3. There were relatively few taxa in the drift, and the mayfly Deleatidium spp. comprised more than 85% of the drifting aquatic invertebrates in all seasons except autumn. Chironomidae and terrestrial forms were the only other groups to occur at densities of more than one animal 100 m^?3 in all seasons. 4. Drift density was positively correlated with benthic density, which in turn was adversely affected by floods, particularly during spring and summer.     

Ontogenetic changes in the drifting of four species of elmid beetles elucidate the complexity of drift-benthos relationships in a small stream in Northwest England

1. This study aimed to quantify ontogenetic changes in the drifting of Elmis aenea, Oulimnius tuberculatus, Esolus parallelepipedus and Limnius volkmari (Coleoptera: Elmidae), and to relate their drift to benthic density. Monthly samples were taken over 39 months, using three surface nets at each of two contrasting sites in a small stream: one in a deep section with abundant macrophytes, and the other in a shallow stony section. 2. Most larvae and adults were taken in the drift at night with little variation between catches in the three nets at each site. Day catches were very low, often zero. No significant relationships could be established between mean numbers in the drift catches and benthic densities. 3. When night catches were converted to drift densities (number caught per 100 m³ of water sampled), the latter were positively related to monthly losses in the benthos, but not to benthic densities. A linear regression described the relationship, and equations for the different life‐stages within each species were not significantly different from the equation for all life‐stages combined. However, drift losses were only about 0.07% of total losses in the benthos. A severe spate in October 1967 increased the number of larvae and adults in the drift, but not drift densities, except for immature adults of E. aenea, O. tuberculatus and E. parallelepipedus. 4. Key life‐stages with the highest drift density were the earliest life‐stage soon after egg hatching for E. aenea, the start of the larval overwintering period for O. tuberculatus and L. volkmari, and mature adults during the mating season for all three species. Drift density for E. parallelepipedus was too low to identify a key life‐stage. These key life‐stages corresponded with critical periods for survival in the life cycle, as identified in an earlier study in the same stream. Mortality was high during these critical periods, hence the strong relationship between drift density and benthic losses. The latter relationship was very consistent for different life‐stages within each species, and partially supported the rarely‐tested hypothesis that drift represents surplus production in the benthos.     

Macroinvertebrate movements in a large European river*

SUMMARY. 1. Quantitative variations in downstream movements of benthic macroinvertebrates were studied in a large European river, the Rhône, upstream from Lyon. Artificial substrates were suspended at three depths in the water column, both near a bunk and in mid-channel, monthly from December 1978 to March 1980. Drift nets were used to determine the diurnal rhythm in drift and to investigate the efficiency of our suspended artificial substrates in capturing the drifting macrofauna. 2. Drift densities (number and biomass) reached a maximum during summer, especially near the river bank, and at night. Mean individual weight of organisms was higher close to the bottom and at night. 3. Artificial substrates were reliable, but underestimated drift by about a quarter in number and a sixth in biomass. compared with drift nets. Two detailed analyses of the drift distribution across the width of the river revealed similar densities along both banks, and uniformity in the channel as a whole. 4. Mean annual drift densities estimated for the section of river were 100 individuals, per 100 m³ and 60 mg dry weight per 100 m³. These densities are similar to those obtained from other temperate rivers.     

Composition and seasonality of benthic invertebrates,and drift in the Naro Moru River,Kenya

A survey was carried out to establish the nature and composition of the benthos along the Naro Moru, a tropical river in central Kenya using artificial substrate baskets, from November 1986 to October 1987. A clear longitudinal zonation existed for Diptera and Ephemeroptera which were the major benthic taxa. Maximum colonization took place after ten days of exposure. Seasonal variations in abundance were also observed. All taxa collected from the bottom samples were also collected in the drift samples, but the percentage composition of the benthos showed variations with that of the drift. Simulium sp. dominated the benthos whilst Baetis spp. dominated the drift. There was a positive correlation between drift rate and benthic fauna density.     

The effect of long pools on the drift of macro-invertebrates in a mountain stream

Macro-invertebrate drift was measured entering and leaving two pools on the Middle Fork of the Cosumnes River, a third order California stream. Drift rates for Baetis spp., Chironomidae, Simulium spp., Capniidae and total drift were calculated. Significant differences in the numbers of organisms entering the two pools were found for Baetis, Chironomidae, and Capniidae. Comparisons of drift rates at the upstream and downstream ends of each pool showed that the abundance of Chironomidae, Simulium, Capniidae and total drift changed in different directions across the pools. The numbers of organisms leaving the two pools, however, were not significantly different for Baetis, Simulium, Capniidae and total drift. These findings lead us to hypothesize that long pools act as barriers, not filters, to stream macro-invertebrate drift. The composition of drift leaving the pools in this experiment appeared to be controlled by the composition of the benthic habitat at the tail of the pool and not by the composition of upstream drift entering the pools.     

Seasonal fluctuations of macroinvertebrate drift in a South Carolina Piedmont stream

The seasonal fluctuations of larval macroinvertebrate drift, exuvial drift and larval benthic density were quantitatively examined over a 1-year period in a fourth order, spring-fed stream in the Piedmont area of South Carolina. The drift was dominated by the mayfly Baetis spp. and by two species of blackfly (Prosimulium mixtum and Simulium jenningsi). Peak drift densities were noted during early spring and especially late summer. Strong correlations were noted between larval drift densities and exuvial drift, indicating a relationship between drift and seasonal growth and emergence patterns. Seasonal trends in drift and benthic densities, though less strongly correlated, were also generally similar.     

Drift behaviour and microhabitat selection in the preimaginal stages of Simulium chutteri (Diptera Simuliidae)

The aquatic stages of the cattle biting pest, Simulium chutteri Lewis utilize river flow in their dispersal and colonization behaviour. Peaks of drifting activity in S. chutteri larvae occurred in the early morning and late afternoon. It is deduced that female flies scatter eggs onto slow flowing waters upstream of rapids. Larvae hatch from drifting eggs and colonize substrates in slower flowing regions upstream of rapids, while later stage instars move into faster flowing regions within rapids where they complete their development. This microhabitat selection by S. chutteri leads to rapid attainment of large population sizes in suitable sections of river and reduces competition between different stage larvae. In their utilization of a variety of microhabitats the larvae of this species differ from co-existing simuliid species which restrict developmental stages to single habitats.Catastrophic drift was recorded for S. chutteri larvae and could be a mechanism to regulate population size.Drift of simuliid larvae off rapids was not related to benthic population densities in the rapids and was therefore not due to excessive production. It is concluded that larval drift off rapids is related to habitat disturbances associated with water flow fluctuations and the activity of aquatic predators and other animals.The distribution of S. chutteri in the Vaal River is restricted by oviposition requirements of the adult female. Knowledge of drift behaviour and water flow requirements of Simuliidae have been applied to manage the population size of S. chutteri in the Vaal River.     

The influence of light on egg buoyancy and hatching rate of the walleye pollock, Theragra chalcogramma

Changes in buoyancy in fertilized bathypelagic eggs of the walleye pollock, Theragra chalcogramma , collected from Shelikof Strait in the Gulf of Alaska were measured under controlled laboratory conditions in density gradient columns from 90 h post–fertilization through hatching. Eggs were incubated at 6° C and exposed to either diel light or constant dark. Eggs held under diel light conditions became more dense than eggs under constant dark beginning <10 h after exposure to light and remained so until 12 h before hatching. Eggs held under constant dark then became more dense than those under diel light. Hatching of eggs under both conditions began at the same time but eggs under diel light showed a delayed hatching rate. Light–induced changes in egg density indicate the ability of walleye pollock eggs to respond to external stimuli and thereby alter their position in the water column in an ecologically meaningful way.     

Egg Deposition Behavior in the Haplodiploid Sawfly Athalia rosae ruficornis Jakovlev (Hymenoptera: Symphyta: Tenthredinidae)

The egg deposition behavior of the turnip sawfly, Athalia rosae (Hymenoptera: Symphyta), is described. Both unmated and mated females lay eggs individually inside of fresh young leaves of cruciferous plants. During an oviposition event, females exhibit a distinct pause in abdominal contractions just before the actual egg deposition act. Unmated females show a longer pause (11.31 s on average) than mated females (4.38 s on overall average). By employing an eye color mutation, the sex of the eggs laid by females was ascertained. Females mated once lay mostly fertilized (diploid female) eggs initially but begin to lay a considerable number of unfertilized (haploid male) eggs later in life. The laying of an unfertilized egg is associated with a longer pause (6.98 s on average) than the laying of a fertilized egg (3.76 s on average). These results are in contrast to previous reports on apocritan Hymenoptera, where the presence of a pause or a longer pause during oviposition was associated with the deposition of fertilized eggs rather than unfertilized eggs. The possibility that mated Athalia rosae females control fertilization and its implications for sex allocation strategies are discussed.     

Mating success of alternative maturation phenotypes in male Atlantic salmon,Salmo salar

Summary Mature male parr successfully fertilized eggs of anadromous female Atlantic salmon, Salmo salar, in competition with anadromous males under simulated natural conditions. Mating situations were created in which mature male parr and anadromous males competing for the opportunity to spawn with an anadromous female differed in allelic forms of the same enzyme. Females deposited eggs into a sequence of 3 or 4 eggs nests. The mean proportion of eggs in a redd fertilized by parr increased with increasing numbers of parr present at a redd, reaching 23% at male parr: anadromous male ratios of 20:1. Single male parr fertilized, on average, 5% of the eggs in a redd. The proportion of eggs in an egg nest fertilized by parr also depended upon the order of egg nest construction, such that parr mating success was highest at the initial nest constructed and lowest at the final nest. Parr have relatively high fertilization success for their size when compared with the smaller maturation phenotype of other salmonids.     

Diel interactions between prey behaviour and feeding in an invasive fish, the round goby, in a North American river

1. We studied the diet of the invasive round goby (Neogobius melanostomus) on a diel basis in the Flint River, a warmwater stream in Michigan, U.S.A. Diet and available prey samples were collected seven times over a 24 h period in four consecutive months. The section of river studied lacked zebra mussels (Dreissena polymorpha), the primary prey of adult round gobies elsewhere in the Great Lakes region. 2. Diet changed on a diel basis with hydropsychid caddisfly and chironomid larvae predominating during the day, chironomid pupae dominating in the evening and heptageniid mayflies dominating at night. Simultaneous study of macroinvertebrate drift suggested that caddisfly and chironomid larvae were most likely picked from submerged rocks, chironomid pupae were most likely taken during their emergent ascent and mayflies were either captured from the drift or picked from rocks. 3. The Flint River lacks a diverse darter (Family: Percidae) and sculpin (Family: Cottidae) fauna and it appears that the round goby has occupied a generalised darter/sculpin niche. Our results indicate that round gobies have the potential to invade successfully riverine systems, particularly those lacking a diverse benthic fish assemblage.     

Ontogenetic shifts in drift periodicity and benthic dispersal in elmid beetles

1. There is a paucity of information on ontogenetic changes in the dispersal of benthic invertebrates, which is an important aspect of their ecology. This study quantifies ontogenetic changes in diel periodicity in drift, and in upstream–downstream dispersal on the substratum for Elmis aenea, Oulimnius tuberculatus, Esolus parallelepipedus and Limnius volkmari (Coleoptera: Elmidae). Three drift nets were emptied every 3 h over 24 h in each month (October 1965–December 1968) at two contrasting sites: one in a deep section with abundant macrophytes, the other in a shallow stony riffle. Comparisons of periodicity between life‐stages of the same species were limited to months when numbers in the drift were highest. Dispersal was evaluated in six experimental stream channels, placed above the stream, with initial numbers of each life‐stage varying from 20 to 80. 2. Drift numbers were always highest at night with few or no animals in the day samples. Ontogenetic shifts in diel periodicity were similar for all four species. Drift catches were similar throughout the night for the early and intermediate larval instars and for mature adults, but were highest in the early hours of the night with a gradual decline thereafter for later larval instars and immature adults. These patterns were unaffected by a severe spate, even though drift numbers increased considerably. 3. Dispersal was density‐independent; the number of dispersing animals was a constant proportion of the initial number for each life‐stage. The relationship between dispersal distance and the number of animals travelling that distance was well described by an inverse power function. Median and maximum distances (m day^?1) were estimated for each life‐stage. 4. Ontogenetic shifts in dispersal in the stream channels matched those shown in diel drift periodicity. For all four species, the later larval instars and immature adults showed little movement in either direction, whereas early and intermediate larval instars and mature adults dispersed predominantly upstream, adults travelling further than any other life‐stage. 5. Ontogenetic shifts in diel drift periodicity and dispersal were related to seasonal changes in drift density and critical periods in the life cycle. Such shifts have not been quantified in other stream invertebrates, but should be considered when evaluating the role of dispersal in their population dynamics and their colonization ability.     

Size-dependent drift responses of mayflies to experimental hydrologic variation: active predator avoidance or passive hydrodynamic displacement?

Summary Larger nymphs within aquatic insect taxa have been frequently observed to be transported down-stream in the stream drift only at night. Others have hypothesized this pattern results primarily from large nymphs' behavioural avoidance of entering drift during daylight, when size-selective, visually-feeding fish predators are most active. This hypothesis assumes that animals can actively control their entry into the drift, which may not be the case under all flow conditions. We experimentally induced streamflow increases and decreases in adjacent riffles in a hydrologically-stable stream during the daytime to examine whether changes in diel patterns of drift abundance and size-distribution of mayflies were consistent with the hypothesis of active avoidance of diurnal drift. We assessed the likelihood of active vs. passive mechanisms of diurnal drift entry and transport for four taxa that differ with respect to body size, morpho-behavioural attributes, microhabitat use, and general propensity to drift. In each of three seasons, diurnal and nocturnal drift samples were collected in three riffles over two diel cycles. Background drift patterns were established on the first day (no flow manipulation). Six h before sunset on the second day, flow was experimentally increased in one riffle, decreased in the second, and not altered in the third (control). Between-day differences in diurnal and nocturnal drift rate and size composition were then compared among the treatment and reference riffles. Responses of two taxa were consistent with active control over drift entry, transport, or both. For Baetis spp., drift-prone mayflies typically preyed upon by fish, diurnal drift rates immediately increased following both flow reduction and flow elevation in all seasons, but only small individuals comprised the drift. Drift by large individuals was delayed until nighttime. Epeorus longimanus also exhibited significant increases in drift rates following flow reduction and elevation, but responses of this large-bodied species were restricted to nighttime. Drift responses for these two taxa were largely independent of direction of hydrologic change, thus indicating a strong behavioural control over drift. By contrast, numbers and sizes of drifting Paraleptophlebia heteronea and Ephemerella infrequens depended strongly on direction of flow change. Drift rates for both species generally declined after flow reduction and increased after flow elevation. Moreover, after flow elevation, larger individuals often drifted diurnally, a finding consistent with expectations under a passive hydrodynamic model. These experiments indicate that size-dependent mayfly drift reflects not only presumed risk from visual fish predators, but also functional attributes of species such as morphology, behaviour, and microhabitat affiliation, which influence aspects of drift entry and transport under variable hydrologic conditions.     

Black fly (Diptera: Simuliidae) response to phosphorus enrichment of an arctic tundra stream

This study reports on the density, growth, and production response of the dominant black fly, Prosimulium martini, to whole river fertilization of the Kuparuk River in arctic Alaska during the summer of 1984. Beginning in 1983, a long term study of fertilization effects was initiated in the Kuparuk River. Increased nutrient supply stimulated algal and microbial biomass and microbial activity, which in turn affected the larval growth and abundance of Prosimulium. This experiment allowed us to isolate the effects of nutrient supply from other factors in determining black fly growth and abundance. Phosphorus addition had the following indirect effects on Prosimulium: growth was higher, but abundance decreased in the enriched section, leading to a net decrease in secondary production from 2.62 g m⁻² yr⁻¹ to 0.77 g m⁻² yr⁻¹. Prosimulium emergence rates were not measureably affected. The decrease in abundance and production appears to be a result of competitive displacement by the caddisfly Brachycentrus americanus which increased in abundance in the fertilized section of the river.     

Diel drift of Chironomidae in a large lowland river (Central Poland)

The diel drift patterns of Chironomidae larvae were investigated in a seventh order section of the Warta River (Central Poland) over two diel cycles during May 1989. Three nets (mesh size 400 m) were installed in a cross section of the Warta River.The estimated drift density was low, but was comparable to that calculated for other large rivers. Spatio-temporal fluctuations in abundance and composition of macroinvertebrate drift, including Chironomidae, were observed with the highest density of drifting macrobenthos recorded near the depositional bank of this river. The ratio benthosdrift indicated differing propensities for of the older instars of a given chironomid taxon to drift. Orthocladiinae larvae were the most abundant subfamily of Chironomidae in drift but not in benthos, reaching up to 73% of the total drifting chironomid larvae. More taxa but fewer individuals (about 20% of the chironomid larvae collected) belonged to the tribe Chironomini, the dominant group in benthos.A major part of chironomid drift collection may represent behavioural drift because the net mesh size used in the Warta River was insufficient to catch the earliest instars (distributional drift). Both at the family and subfamily level chironomid larvae exhibited a distinct nocturnal drift periodicity. Nocturnal periodicity was documented for the dominant species, but due to the low density of many chironomid species, it was impossible to determine their diel drift pattern. Some Chironomidae appeared to be aperiodic.     

Zur Fortpflanzungsbiologie vonLineus viridis (Nemertini)

The fertilization of the heteronemerteanLineus viridis Müller takes place in a cocoon built up by the female surrounding itself and the male. After the male has left the cocoon, the female builds a mucous layer within the cocoon, in which the fertilized eggs enclosed in egg capsules are embedded. Observations on mating behavior and spawning are described and discussed.