The influence of predatory fish on mayfly drift: extrapolating from experiments to nature

1. A knowledge of how individual behaviour affects populations in nature is needed to understand many ecologically important processes, such as the dispersal of larval insects in streams. The influence of chemical cues from drift‐feeding fish on the drift dispersal of mayflies has been documented in small experimental channels (i.e. < 3 m), but their influence on dispersal in natural systems (e.g. 30 m stream reaches) is unclear. 2. Using surveys in 10 Rocky Mountain streams in Western Colorado we examined whether the effects of predatory brook trout (Salvelinus fontinalis) on mayfly drift, that were apparent in stream‐side channels, could also be detected in natural streams. 3. In channel experiments, the drift of Baetis bicaudatus (Baetidae) was more responsive to variation in the concentration of chemical cues from brook trout than that of another mayfly, Epeorus deceptivus (Heptageniidae). The rate of brook trout predation on drifting mayflies of both species in a 2‐m long observation tank was higher during the day (60–75%) but still measurable at night (5–10%). Epeorus individuals released into the water column were more vulnerable to trout predation by both day and night than were Baetis larvae treated similarly. 4. Drift of all mayfly taxa in five fishless streams was aperiodic, whereas their drift was nocturnal in five trout streams. The propensity of mayflies to drift was decreased during the day and increased during the night in trout streams compared with fishless streams. In contrast to the channel experiments, fish biomass and density did not alter the nocturnal nature nor magnitude of mayfly drift in natural streams. 5. In combination, these results indicate that mayflies respond to subtle differences in concentration of fish cues in experimental channels. However, temporal and spatial variation in fish cues available to mayflies in natural streams may have obscured our ability to detect responses at larger scales.     

Age,growth, and sex ratio among populations of least brook lamprey,Lampetra aepyptera,larvae: an argument for environmental sex determination

Synopsis Sex ratios of least brook lamprey,Lampetra aepyptera, larvae varied widely among 12 geographically-diverse streams of the eastern United States. The extremes were 29 and 71% male, and the proportion of males increased significantly with relative population density, which was estimated among the streams from the number of larvae collected per m2 of substrate. The skewed sex ratios were not likely due to differential mortality between the sexes or differential recruitment to the adult stock, since they were established at the time of gonadal differentiation (at ca. 2 years of age) and remained relatively constant over the subsequent 2–3 years of larval life. Furthermore, although females seemed to predominate in the oldest larval age class, thus appearing to metamorphose later than males, their numbers were small and were omitted from the overall sex ratio. Sex ratio did not vary significantly with water hardness, pH, annual thermal units, or latitude. The possible adaptive significance of density-dependent sex determination in lampreys, however, remains elusive. It has been proposed that growth-promoting conditions might yield female-biased sex ratios as a tactic for ensuring that relatively large individuals become females, thereby increasing their fecundity. As predicted, larval size at a given age was generally greater in low-density populations, but there was no relationship between sex ratio and larval size, and female larvae were not consistently larger than the males.     

Effects of structural habitat on drift distance and benthic settlement of the caddisfly,Ceratopsyche sparna

We conducted two experiments in flow-through, artificial streams to examine how habitat structure affected drift and benthic resettlement of larval hydropsychid caddisflies (Ceratopsyche sparna). In the first experiment, we quantified drift distance and the number of times larvae re-entered the drift in 9.0 × 0.51-m channels with contiguous patches (ea. 2.5-m long) of biofilm-covered gravel, biofilm-covered cobbles (– Cladophora), and Cladophora-covered cobbles (+ Cladophora). In the second experiment, we tracked nocturnal movements of larvae after benthic settlement in 2.8 × 0.1-m channels, each containing one of the three habitat types. In experiment 1, drift distance was (1) greatest in gravel and lowest in cobbles + Cladophora, (2) inversely related to hydraulic roughness of habitats, (3) independent of body size, and (4) similar for live and dead larvae. Average drift distance was relatively short (<2.5 m), regardless of habitat type. Number of drift re-entries also varied among habitats, being greatest in gravel and lowest in cobbles + Cladophora. No larvae re-entered the drift after settling in Cladophora patches. Results from experiment 2 revealed that drift propensities were higher for larvae in biofilm-covered gravel and cobbles than in cobbles + Cladophora. Larvae remaining in substrate patches (i.e. not drifting) laid fewer draglines in biofilm-covered stones than in Cladophora patches. Extent of benthic movement (i.e., crawling) by non-drifting larvae did not differ significantly among habitats. However, distance moved did differ with flow direction, being 4× greater downstream than upstream. These results highlight how local substrate and hydraulic conditions interact to affect small-scale movements of caddisfly larvae.     

Assessment of the vertical distribution of larval lake sturgeon drift in the Peshtigo River, Wisconsin, USA

Drift nets have been used to document reproductive success of lake sturgeon Acipenser fulvescens. Current net designs and methods for collecting drifting larvae only sample a portion of the water column, which require assumptions of either a benthic or uniform distribution of larvae when estimating abundance or production. The objective of this study was to describe the vertical distribution of larval lake sturgeon in the Peshtigo River, Wisconsin, and to determine if drift was benthic or uniform in distribution. A net was designed to assess the vertical distribution of drifting larvae in 0.2-m increments at depths up to 1.4 m; however, during this sampling, maximum depth did not exceed 0.78 m. The distribution of larval lake sturgeon was neither benthic nor uniform. Only 5% of larvae were captured in the lowest 0.2-m increment, followed by 18% from 0.2 to 0.4 m, 41% from 0.4 to 0.6 m, and 36% from 0.6 m to the surface. Although results will likely differ among years, systems, and the precise location of sampling, our study illustrates the importance of and provides a technique for testing assumptions of the vertical distribution of larval lake sturgeon drift.     

Influence of watering and trenching ponderosa pine on a pine sawfly

Summary Neodiprion autumnalis (Smith) larvae were caged for two successive years on root-trenched, watered, and untreated ponderosa pine (Pinus ponderosa Doug. ex Laws.) to determine effects of host moisture stress on larval feeding. Levels of moisture stress (as measured by the Scholander pressure chamber) differed significantly among treatment levels during 1984 and 1985 larval feeding periods. Differences in larval feeding success were not detected in 1984. In 1985, however, larvae on trenched (stressed) trees clipped and rejected more foliage, consumed more needles, had lower pupal weights, lower survival, and a longer feeding period than larvae on watered or untreated trees. Frass production did not differ among treatment levels. The length of the feeding period was shorter for larvae on watered trees than for larvae on untreated trees, but other measures of feeding success did not significantly differ between watered and untreated trees.     

Some beneficial effects of aggregation in young larvae ofPryeria sinica moore (Lepidoptera: Zygaenidae)

The effects of group size on the survival and development of young larvae of Pryeria sinicaMoore were investigated by laboratory and field experiments. Under laboratory conditions, about 20% of isolated larvae died of unsuccessful feeding in the first instar, however, larvae survived successfully in aggregations of four or more individuals. In the field, larvae emerge in early spring and wait for new leaves to open before feeding. In this period, the larger the group size of hatchlings the survival rate became higher. The nest-web spun by hatchlings was considered to play an important role in protecting them from desiccation. In the period that larvae began to feed on leaves, more than 36 larvae are necessary to aggregate for the successful establishment of feeding groups. The nest-web played an important role also in the establishment of feeding group. However, the natural group size of the first instar larvae was larger than the minimum group size to spin a sound nest-web in the field experiment. On the other hand, in later stage, larvae in a large group did not have an excess advantage in survival or developmental rate over larvae in a small group. It was found that the experiments on survival and developmental rates could not explain the reason that this species maintain large compact groups in the most part of larval period.     

Dynamics and extent of larval lake sturgeon Acipenser fulvescens drift in the Upper Black River, Michigan

Lake sturgeon larval drift is not uniform in time or space and subsequent efforts to determine the relative abundance have suffered because of the lack of information during this early life history period. The purpose of this study was to obtain information about the early life history of lake sturgeon, determine the extent and duration of lake sturgeon larval drift, and examine this relationship to water flow and temperature in the Upper Black River, Michigan. This study also compares the results of other studies to further evaluate the dispersion of larvae. Larval production was quantified using drift nets anchored to the stream bottom from May to June in 2000–2002. Larval drift nets captured 780 larvae in 2000; 2975 larvae in 2001; and 2041 larvae in 2002. For the 2000, 2001, and 2002 spawning season, we estimated that 7107 (95% CL: ± 1470), 17 409 (95% CL: ± 5163), and 15 820 (95% CL: ± 3168) larval lake sturgeon were produced in the Upper Black River (UBR), respectively. Catch per unit effort values of drifting larvae were greatest after peak water flows, with most larvae captured in the middle of the river channel. A mean daily water temperature above 16°C was an important environmental stimulus that influenced peak larval dispersion away from spawning sites. The results of this study suggested that natural reproduction was still occurring in the Black Lake system.     

The effect of a predatory leech,Nephelopsis obscura,on mortality,growth, and production of chironomid larvae in a small pond

Summary The effect of a predatory leech, Nephelopsis obscura, on survivorship, growth, and production of chronomid larvae was studied by enclosure experiments carried out in a small pond. The prey population was composed almost entirely of the tubiculous, microphagous chironomid larvae, Chironomus riparius and Glyptotendipes paripes. Nephelopsis significantly reduced chironomid survivorship within the enclosures, and accounted for most of the measured mortality of fourth instar larvae. The cropping by Nephelopsis was not significantly biased toward either prey species. In long-term experiments (66 d) chironomid biomass in enclosures without leeches reached much higher levels than in enclosures containing Nephelopsis. This increase in biomass was due to growth of surviving larvae, rather than recruitment, since emergence and oviposition were not going on during the course of the experiments. The enhanced survivorship of larvae within leech-free enclosures was eventually accompanied by reduced growth and specific production (daily production/biomass) for C. riparius, which made up about 90% of the larval population. Growth and specific production of G. paripes (10% of larval population) was not affected. Short-term experiments (25 d) involving manipulation of densities and species ratio (9:1 CR:GP and 1:9 CR:GP) of larvae revealed that growth of the majority species was strongly influenced by larval density, whereas growth of the minority species was not. The same pattern was observed both in the presence and in the absence of Nephelopsis and was a result of differences in resource utilization between the two species. In the shortterm experiments, growth rates estimated for larvae exposed to leeches were significantly less than those for larvae in leech-free enclosures. This could be due either to size-biased consumption of larvae by Nephelopsis, or possibly a disturbance factor leading to reduced larval food intake and/or increased metabolic costs.     

Density of an intraguild predator mediates feeding group size, intraguild egg predation, and intra- and interspecific competition

Intraguild predation (IGP) is common in most communities, but many aspects of density-dependent interactions of IG predators with IG prey are poorly resolved. Here, we examine how the density of an IG predator can affect feeding group size, IG egg predation, and the growth responses of IG prey. We used laboratory feeding trials and outdoor mesocosm experiments to study interactions between a social intraguild predator (larvae of the wood frog; Rana sylvatica) and its prey (spotted salamander; Ambystoma maculatum). Larvae of R. sylvatica could potentially affect A. maculatum by consuming shared larval food resources or by consuming eggs and hatchlings. However, successful egg predation requires group feeding by schooling tadpoles. We established from five to 1,190 hatchlings of R. sylvatica in mesocosms, then added either 20 A. maculatum hatchlings to study interspecific competition, or a single egg mass to examine IGP. Crowding strongly suppressed the growth of R. sylvatica, and IGP was restricted to the egg stage. In the larval competition experiment, growth of A. maculatum was inversely proportional to R. sylvatica density. In the predation experiment, embryonic mortality of A. maculatum was directly proportional to the initial density of R. sylvatica and the mean number of tadpoles foraging on egg masses. IGP on eggs reduced A. maculatum hatchling density, which accelerated larval growth. Surprisingly, the density of R. sylvatica had no overall effect on A. maculatum growth because release from intraspecific competition via egg predation was balanced by increased interspecific competition. Our results demonstrate that the density of a social IG predator can strongly influence the nature and intensity of interactions with a second guild member by simultaneously altering the intensity of IGP and intra- and interspecific competition.L . A. Burley and A. T. Moyer contributed equally to this work.     

Food partitioning between coexisting Atlantic salmon and brook trout in the Sainte‐Marguerite River ecosystem, Quebec

Food resource partitioning between similar‐sized, sympatric Atlantic salmon Salmo salar and brook trout Salvelinus fontinalis was examined as a possible mechanism enabling their coexistence in a stream (Allaire) of the Sainte‐Marguerite River ecosystem, Quebec, Canada. Fish stomach contents and invertebrate drift were collected concurrently during three diel cycles in August to September 1996. The food and feeding habits of an allopatric brook trout population in a nearby stream (Epinette) were studied for comparison. The diel feeding rhythms of the two coexisting fish species were similar. The composition of their diet, however, showed significant differences. Atlantic salmon predominantly (60–90%) fed on aquatic insects, mainly Ephemeroptera (35–60% of the diet). The brook trout mostly (50–80%) fed upon the allochthonous terrestrial insects (mainly adults of Coleoptera, Hymenoptera and Diptera) which comprised 5–40% of the stream drift. The allopatric brook trout fed opportunistically on the more abundant aquatic insects and terrestrial insects rarely formed 25% of its diet. The allopatric trout fed nearly twice as much as the sympatric brook trout during a day. The results suggest that the differences in feeding by brook trout in the two streams (with and without Atlantic salmon) are the result of inter‐specific interaction with Atlantic salmon and are not related to the differences in food availability between the two streams. Food resource partitioning between Atlantic salmon and brook trout may be viewed as an adaptive response resulting in a greater exploitation of available resources and coexistence.     

Night — day drift patterns and the size of larvae of two aquatic insects

A test of Allan's (1978) hypothesis about differential drift abundance of mayflies of the genus Baetis between night and day, and the size of larvae was performed at a mountain stream in Idaho. Palisades Creek, Idaho, contains a different species of mayfly, B. tricaudatus, and vertebrate predator, Salmo clarki, than Cement Creek, Colorado (B. bicaudatus, and brook trout, Salvelinus fontinalis). Consequently it was not known if B. tricaudatus would exhibit a similar pattern as its congener in Cement Creek, with large instars tending to avoid daylight drift, as found by Allan (1978). However, similar results were observed in the present study. It appears that the earlier hypothesis may have generality for geographically distinct streams with a different vertebrate predator and mayfly prey. The existence of a similar pattern for chironomid larvae was also tested, however, no such pattern existed. This discrepency between taxa may be due to differential predation, or to inherent differences in drift abilities.     

Field experiments on the relationship between drift and henthic densities of aquatic insects in tropical streams (Ivory Coast). III Trichoptera

SUMMARY. 1. Based on in situ gutter trials we related the drift of caddis flies to their benthic densities and to various abiotic factors in streams in the Ivory Coast (West Africa). Members of the families Hydropsychidae, Philopotamidae. Hydroptilidae and Leptoceridae were considered in detail.
2. The drift of larvae peaked at night in both early and late larval instars.
3. Drift of a larval group (a certain instar, species or higher taxon) was more often related to the benthic density of other larval groups than to its own benthic density.
4. Self-regulation of an upper benthic density of a larval group by emigration through drift was not statistically evident.
5. There was no straightforward relationship between drift and abiotic factors.
6. Drift rates differed between taxa as well as between larval instars (size groups) within a taxon. Newly hatched larvae had very high drift rates, whereas the last larval instar usually had the lowest drift rate.
7. We related these results to the violently fluctuating discharge of the streams in the study area and the consequent variability of space for lotic insects.
8. Drift estimates, made at the same time as a monitoring programme on possible side-effects of insecticides (Onchocerciasis Control Programme), failed to reflect benthic densities except in the night drift of Hydropsychidae.     

Development of a meridic diet for Hylobius transversovittatus (Coleoptera: Curculionidae) and the role of carbohydrates in feeding, growth, and survival of larvae

The root-feeding weevil Hylobius transversovittatus Goeze (Coleoptera: Curculionidae) is used for biological control of the invasive plant purple loosestrife, Luthrum salicaria L. (Lythraceae). A simple rearing system for this weevil was developed with the goals of improving production techniques and increasing the availability of insects for field introduction. Additionally, the dietary effects of digestible and indigestible carbohydrates were explored. A meridic diet for rearing H. transversovittatus was formulated through nutritional alterations of a boll weevil, Anthonomus grandis grandis Boheman, diet. Diet attractiveness was evaluated on two levels: first, by recording the incidence of initial tunneling, and second, by estimating the larval establishment rate. The performance of test diet formulations was further assessed by measuring developmental and survival rates of H. transversovittatus. Sucrose, starch, and three types of indigestible carbohydrates were tested as components to improve diet performance. Physical properties of the diet, modified by fillers in test formulations, produced major effects on the initial tunneling of hatchlings. The establishment of hatchlings was affected by chemical properties of the diet. Increases in sucrose concentration decreased larval establishment, decreased the rate of larval development, and decreased larval survival. However, omitting sucrose from the diet, or replacing it with starch, increased mortality of first instars. In advanced stages of larval development, omitting sucrose from the diet did not significantly affect larval survival. The developmental rate of larvae was increased when the amount of digestible carbohydrate was reduced. To date, seven generations of the univoltine H. transversovittatus have been successfully produced on this new meridic diet.     

Micro-habitat use within a guild of newt larvae (Trituridae) in an Alpine lake

Population and ecological parameters such as numbers of larvae, microhabitat use, niche breadth and niche overlap of three species of syntopic larval newts (Alpine newt Triturus alpestris, Italian crested newt T. carnifex, and common newt T. vulgaris) were studied for two years in a small pond at 1160 m a.s.l. in NE Slovenia. Differences in microhabitat partitioning among larval newts were small. The largest niche breadth was estimated for larval T. alpestris, and the narrowest estimate was for larval T. carnifex in both years. Ecological differences seem to be very small and quite variable among sites and years. It appears that the developmental stage and size of newt larvae are more important in explaining resource partitioning than the characteristics of each species. Because of the absence of potential invertebrate predators and adult newts in the second half of the breeding season, the injuries could only be caused by intra-and interspecific predation attempts.     

Gregariousness,field distribution and defence in the sawfly larvae Croesus varus and C. septentrionalis (Hymenoptera,Tenthredinidae)

Summary The larvae of Croesus varus are cryptic and those of C. septentrionalis aposematic. In both species, females lay eggs in groups, but the groups formed by young larvae are smaller in C. varus than in C. septentrionalis. In both species, group size decreases during successive larval instars, and the cryptic species becomes solitary, while the aposematic species remains gregarious during all larval instars. On Alnus shrubs in the field, the distribution of the groups ofC. septentrionalis larvae is more aggregative. They are found mainly on mediumsized shrubs (1.5–2.5 m high), whereas groups of C. varus larvae are also found on larger shrubs. Moreover, during the 2 years of observation, C. septentrionalis was present for a shorter time than C. varus, even when the former was much more abundant than the latter. Thus, conspicuousness of the aposematic species should be amplified by this spatial and temporal aggregation. The efficiency of both defensive strategies is compared and discussed, taking into account these particular field distributions, predation and some parasitism factors, and the chemical defences of both species.     

Effects of larval age and prolonged simulated acid rain on the susceptibility of European pine sawfly to virus infection

We studied the effect of simulated acid rain treatment of host trees on the susceptibility of the European pine sawfly larvae to virus, and possible differences when larvae of two different ages were infected. Older larvae were less susceptible to virus. Most larvae treated with virus 2 days after they started feeding on experimental foliage (group A) died rapidly within 10 days after the virus treatment, and survival to the end of the larval period was only 8–25%. Larvae treated with virus 1 week later (group B) were less affected by the virus and 36–49% survived. In group A the larval survival in the pH 3 treatment was higher than in other treatments; at the end of the larval period the difference was twofold. In group B there were no clear effects of acid rain on the susceptibility of larvae to virus. The study yielded the following new information: (1) the effect of prolonged acid rain treatment on reducing the efficacy of virus on young larvae was more distinct than in a previous study with shorter exposure to acid rain, and the difference was maintained to the end of larval period; (2) the susceptibility of older larvae to virus was not affected by acid rain treatments; (3) pH inside the needles did not explain the larval mortality caused by virus.     

Downstream drift of the larvae of Chironomidae (Diptera) in the River Chew,S.W. England

The drift of larval Chironomidae (Diptera) was investigated at two sites on the River Chew, using a pump filtration system with 50 µm mesh-aperture aerial nets, situated on the river bank.Chironomid larvae were found to drift in far greater numbers than previously reported, even in slow-flowing water. First and second instar larvae dominated the drift.Variation in drifting behaviour between taxa was observed, where Orthocladiinae drifted in all instars and Chironominae predominantly as first and second instars. Proportionally different rates of drift were observed between these taxa from the benthos at the two sites.It is suggested that the majority of chironomid drift represents an active dispersal and colonisation mechanism by which population redistribution and habitat selection occurs.     

Invertebrate infestation on eggs and hatchlings of the loggerhead turtle, Caretta caretta, in Dalaman, Turkey

The damage caused by some invertebrates to the eggs and hatchlings of loggerhead turtles, Caretta caretta, was investigated during the summer of 2002 on Dalaman beach, Turkey. The specimens, identified to family or genus levels, from nine families representing seven orders were recorded as infesting nests of loggerhead turtles. The heaviest impacts on loggerhead turtle nests was made by Pimelia sp. (Tenebrionidae, Coleoptera). Twenty-four (36.3%) out of 66 intact loggerhead hatched nests were affected by these larvae. Larval damage by Pimelia sp. was recorded in 188 (10.6%) out of 1773 eggs, but only in two (0.28%) hatchlings. The results show that fewer insects were in the nest the further from vegetation and therefore the relocation of nests from the water's edge to further inland close to vegetation may increase the infestation rate of the eggs.     

Hydraulic microdistribution patterns of larval fire salamanders (Salamandra salamandra salamandra) in the Weidlingbach near Vienna, Austria

1. This study examined the longitudinal distribution of larval fire salamanders ( Salamandra salamandra salamandra L.) in the Weidlingbach system, a first- to fourth-order tributary of the Danube near Vienna, Austria. On the microhabitat scale, actual current velocities, Reynolds numbers and Froude numbers at larval locations were measured and larval positions mapped.
2. Larval densities were highest in shallow first- and second-order tributaries where mean current speeds were less than 20 cm s⁻¹, mean Reynolds numbers were less than 12 000 and mean Froude numbers were less than 0.30. Young larvae appeared on 26 April, fully grown larvae were observed from 11 May and larvae started metamorphosis from 27 June. Young adults left the brook between 26 August and 8 September. A severe flood on 25 May significantly reduced larval density, especially at sites near the source.
3. At a given sampling station young larvae favoured microhabitats exposed to only minor hydraulic stress. As larvae grew, they also colonized microhabitats with moderate current velocity (4–15 cm s⁻¹). At sites with generally low discharge, larvae were distributed over a wider range of current velocities than at sites with higher discharge; at the latter, larvae were constricted to sheltered microhabitats.     

Using spatial, seasonal, and diel drift patterns of larval Lost River suckers Deltistes luxatus (Cypriniformes: Catostomidae) and shortnose suckers Chasmistes brevirostris (Cypriniformes: Catostomidae) to help identify a site for a water withdrawal structure on the Williamson River, Oregon

A small irrigation diversion dam near Chiloquin, Oregon, was removed and replaced with a pump station to improve fish passage for Lost River suckers (Deltistes luxatus) and shortnose suckers (Chasmistes brevirostris) entering the Sprague River on their spawning migrations. During the developmental phase of the pump station, a need was identified to better understand the larval drift characteristics of these endangered catostomids in order to reduce entrainment into the irrigation system. The spatial, seasonal, and diel distribution of drifting larvae was measured during the 2004 spawning season at two proposed sites on the Williamson River where the pump station could be located. Larval drift for both species coincided with the irrigation season making them subject to entrainment into the irrigation system. Drift occurred almost exclusively at night with larvae entering the drift at sunset and exiting the drift at sunrise. Nighttime larval densities were concentrated near the surface and at midchannel at both sites. Densities were generally greater on the side of mid-channel with greater flow. During early morning sampling we detected a general shift in larval drift from surface to subsurface drift. We also observed an increase in larval densities towards the shore opposite from the proposed pump station at the upper site whereas larval densities remained high at midchannel at the lower site. During daytime sampling, the few larvae that were collected were distributed throughout the water column at both pump sites. This study found that larvae drifting during all time periods were generally distributed further across the cross section, deeper in the water column, and closer to where the proposed water withdrawal structure would be built at the downstream site when compared to the upstream site. Recommendations were provided to locate the withdrawal facility at the upstream site and operate it in a manner such that larval entrainment would likely be minimized.