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.     

Seasonal and inter-stream variations in the population dynamics, growth and secondary production of an algivorous fish (Pseudogastromyzon myersi: Balitoridae) in monsoonal Hong Kong

1. Balitorid loaches are widespread and highly diverse in Asian streams, yet their life history and ecology have received little attention. We investigated seasonal (wet versus dry season) and spatial variation in populations of algivorous Pseudogastromyzon myersi in Hong Kong, and estimated the magnitude of secondary production by this fish in pools in four streams (two shaded and two unshaded) over a 15‐month period. 2. Mean population densities of P. myersi ranged from 6.0 to 23.2 individuals m⁻², constituting more than half (and typically >70%) of benthic fishes censused. Abundance was c. 25% greater in the wet season, when recruitment occurred. Significant density differences among streams were not related to shading conditions and were evident despite small‐scale variations in P. myersi abundance among pools. Mean biomass varied among streams from 0.85 to 3.87 g ash‐free dry weight (AFDW) m⁻². Spatial and seasonal patterns in biomass and density were similar, apart from some minor disparities attributable to differences in mean body size among populations. 3. All four P. myersi populations bred once a year in June and July, and life spans varied from 24 to 26 months. Populations consisted of three cohorts immediately after recruitment but, for most of the study period, only two cohorts were evident. Cohort‐specific growth rates did not differ significantly among streams but, in all streams, younger cohorts had higher cohort‐specific growth rates. 4. Secondary production of P. myersi estimated by the size‐frequency (SF) method was 2.7–11.5 g AFDW m⁻² year⁻¹ and almost twice that calculated by the increment‐summation (IS) method (1.2–6.6 g AFDW m⁻² year⁻¹). Annual P/B ratios were 1.17 – 2.16 year⁻¹ (IS) and 2.73 – 3.22 year⁻¹ (SF). Highest production was recorded in an unshaded stream and the lowest in a shaded stream, but site rankings by production did not otherwise match shading conditions. Wet‐season production was six times greater than dry‐season production, and daily production fell to almost zero during January and February. Cool temperatures (<17 °C) may have limited fish activity and influenced detectability during some dry‐season censuses. Estimates of abundance and annual production by P. myersi are therefore conservative. 5. Comparisons with the literature indicate that the abundance and production of P. myersi in Hong Kong was high relative to other benthic fishes in tropical Asia, or their temperate counterparts in small streams. Manipulative experiments are needed to determine the influence of P. myersi, and algivorous balitorids in general, on periphyton dynamics and energy flow in Asian streams.     

Response of grazing impacts of an algivorous fish (Pseudogastromyzon myersi: Balitoridae) to seasonal disturbance in Hong Kong streams

1. Manipulative experiments were carried out in four Hong Kong streams (two shaded, two unshaded) to investigate the impact of grazing by an algivorous fish, Pseudogastromyzon myersi, on benthic algal biomass and assemblage composition. Experiments were conducted and repeated during both the dry and wet seasons to determine whether spate‐induced disturbance modified any grazing effect. Treatments comprised fish exclusion and inclusion via closed and open cages, with a no‐cage treatment used as a control for the cage effect. Treatments were maintained for 4 weeks in each experimental run. 2. Grazing by P. myersi reduced benthic algal biomass and the organic matter content of periphyton in open cages and the no‐cage treatment relative to closed cages. The similarity between open‐cage and no‐cage treatments was evidence that the overall difference among treatments was caused by limiting fish access to closed cages and not merely an artifact of caging. Grazing effects were broadly similar in all streams, but there was a significant statistical interaction between treatments and seasons. 3. Analysis of dry‐season data matched the overall trend in inter‐treatment differences, confirming the effects of grazing by P. myersi on algal biomass and periphyton organic matter. Significant differences in algal assemblage composition between closed‐cage and no‐cage treatments during the dry season reflected reductions in the abundance of erect, stalked diatoms (Gomphonema) and filamentous cyanobacteria (Homeothrix). Removal of these vulnerable overstorey algae by P. myersi resulted in greater abundance of understorey diatoms (Achnanthes and Cocconeis) in the no‐cage treatment in all streams during the dry season. The composition of algal assemblages in open cages was intermediate between the other two treatments. 4. Although fish densities were greater in all streams during the wet season, spate‐induced disturbance obscured grazing effects and there were no significant differences among treatments attributable to fish grazing. Seasonal variation in impacts of P. myersi grazing provides support for the harsh‐benign hypothesis, and confirms that biotic factors are less important controls of stream algal biomass and assemblage structure during periods (i.e. the wet season in Hong Kong) when abiotic disturbances are frequent or intense.     

Length–weight relationships for 19 headwater fish species from streams in the Sinos River basin,southern Brazil

This study reports the length–weight relationship estimates (LWRs) for 19 fish species inhabiting low order headwater streams in the Sinos River basin of southern Brazil. Between 2010 and 2014, 19 species at 47 sites were sampled in 10 sub‐basins by electrofishing in winter and summer. For two species, Bryconamericus iheringii and Phalloceros caudimaculatus, the estimates in this study offer new maximum size ranges compared to previous studies.     

The ecology of pre-imaginal simuliidae (Diptera) in South-East Queensland,Australia

A study of the pre-imaginal simuliid fauna of South-East Queensland was conducted to elucidate some of the aspects of their ecology. A total of 17 species of three genera (Cnephia, Austrosimulium and Simulium) bred in the area. The highest diversity of species was found to occur in mountainous areas near the coast which had the following conditions: high rainfall, steep relief, cool temperatures, on the edge of forests with streams of the order 1–3 (using a 1 : 250,000 map).More detailed studies of A. bancrofti, S. nicholsoni and S. ornatipes were made to investigate microdistribution, colonization, drift, pupation timing and population changes. Current velocity was the most important factor determining the distribution of A. bancrofti, while the distribution of S. nicholsoni was influenced by a preference for vegetation substrate and current velocity. Late instar larvae of A. bancrofti and S. ornatipes occurred in faster currents than early instar larvae. Pupae of these two species primarily occurred on the downstream side of submerged substrates. Colonization of a particular stream was dependent on the oviposition preference of gravid females while drift, looping, or descent on silk threads was used for larval dispersal from the oviposition site. Early instars of A. bancrofti drifted from quiet reaches of mature rivers to rapids, while S. ornatipes larvae appeared to be more sessile with limited dispersal from site of egg masses. Drift occurred throughout 24 hours with little change in total numbers. Early instars of A. bancrofti however showed a definite diurnal tendency while the proportion of late instars increased at night. Pupation of S. ornatipes was diurnal under undisturbed conditions. The larvae of A. bancrofti and S. nicholsoni both developed in the Brisbane River system, however the populations of A. bancrofti peaked in late winter, a dry season with stable water conditions while S. nicholsoni larvae were most abundant from late summer, at the end of the rains to the early winter. The larvae of S. ornatipes, which occur in small streams down to temporary trickles, were most numerous during the warm rainy period.Department of Parasitology, University of Queensland     

The Chaoborus (Diptera: Chaoboridae) component of the invertebrate drift of the Mwenda river,Lake Kariba,Central Africa

Chaoborus larvae were collected from the riverine drift of the Mwenda River during the 1972/3 rainy season. Large numbers of larvae were found in the early stages of flow following the 1972 dry season, fewer were found ten days later.Larvae which reached the lake in this way appeared to show no ill effects. Possible mortality arising prior to reaching the sampling point was doubtful since no dead larvae or parts were found in the material collected.     

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.     

Effects of trout on the diel periodicity of drifting in baetid mayflies

Some benthic invertebrates in streams make frequent, short journeys downstream in the water column (=drifting). In most streams there are larger numbers of invertebrates in the drift at night than during the day. We tested the hypothesis that nocturnal drifting is a response to avoid predation from fish that feed in the water column during the day. We surveyed diel patterns of drifting by nymphs of the mayfly Baetis coelestis in several streams containing (n=5) and lacking (n=7) populations of rainbow trout, Oncorhynchus mykiss. Drifting was more nocturnal in the presence of trout (85% of daily drift occurred at night) than in their absence (50% of daily drift occurred at night). This shift in periodicity is due to reduced daytime drifting in streams with trout, because at a given nighttime drift density, the daytime drift density of B. coelestis was lower in streams occupied by trout than in troutless streams. Large size classes of B. coelestis were underrepresented in the daytime drift in trout streams compared to nighttime drift in trout streams, and to both day and night drift in troutless streams. Differences in daytime drift density between streams with and without trout were the result of differences in mayfly drift behaviour among streams because predation rates by trout were too low to significantly reduce densities of drifting B. coelestis. We tested for rapid (over 3 days) phenotypic responses to trout presence by adding trout in cages to three of the troutless streams. Nighttime drifting was unaffected by the addition of trout, but daytime drift densities were reduced by 28% below cages containing trout relative to control cages (lacking trout) placed upstream. Drift responses were measured 15 m downstream of the cages suggesting that mayflies detected trout using chemical cues. Overall, these data support the hypothesis that infrequent daytime drifting is an avoidance response to fish that feed in the water column during the day. Avoidance is more pronounced in large individuals and is, at least partially, a phenotypic response mediated by chemical cues.     

Drift and settlement of stream insects in a complex hydraulic environment

1. The hydraulic and geomorphic characteristics of stream patches are often associated with distinctive assemblages or densities of stream invertebrates, and it is routinely presumed that these patterns reflect primarily species‐specific habitat requirements. An alternative hypothesis is that such patterns may be influenced by constraints on movement, such as the results of departure and settlement processes. We describe a manipulative experiment that examined how the hydraulic environments created by topographic bedforms influenced the drift behaviour and potential settlement sites for two species of mayfly (Baetis rhodani and Ecdyonurus torrentis). These species are common in the drift and often co‐occur in streams, but differ in their small‐scale distribution patterns, body shape and movement behaviour. 2. Flume experiments were carried out to determine how the hydraulic environments conditioned by a step bedform influence the behaviour of mayflies in the drift (swimming, posturing, tumbling), and the consequences of those behaviours (drift distance and time), compared to drift over a plane bed. The ramped step in the flume mimicked step bedforms that are common in coarse‐grained, high‐gradient streams. In contrast to the plane bed, a zone of recirculating flow was created downstream of the step, above which flow was faster and more turbulent. Uniform flows are used in most flume studies of drift; our approach is novel in recreating a complex hydraulic environment characteristic of stream channels. 3. Both species had some behavioural control over drift, and drift distances and times were shorter for live larvae than for dead larvae over the plane bed. The step had no impact on drift time or distance for live Baetis, but dead larvae were trapped in the flow separation eddy and drift time increased accordingly. Some Ecdyonurus also became trapped in the eddy, but live larvae drifted farther than dead larvae, and farther over the step than the plane bed. 4. Whilst in the drift, larvae altered their behaviour according to the ambient hydraulic environment, but in a species‐specific manner. Over the plane bed, Baetis had occasional swimming bursts, but primarily postured (maintained a stable body orientation), whereas Ecdyonurus spent roughly equal time posturing and swimming. In the more turbulent flows generated by the step, Baetis spent proportionately more time swimming, whereas Ecdyonurus spent more time posturing and often tumbling as body orientation became unstable. 5. In a high‐gradient stream, Baetis was more abundant close to steps than in plane bed patches with less complex flow, whereas the opposite pattern held for Ecdyonurus. Thus, the small‐scale distribution patterns of these species within streams correspond to their drift behaviours and ability to access various hydraulic patch types in our flume. These results are consistent with the hypothesis that constraints on movement and settlement may be important driver of distribution patterns within streams.     

Seasonal variation in diel activity and microhabitat use of an endemic New Zealand stream-dwelling galaxiid fish

1. Seasonal variation in microhabitat use and activity of 14 giant kokopu Galaxias argenteus, a drift‐feeding galaxiid fish, was compared using radiotelemetry. 2. During winter giant kokopu predominantly used low velocities and intermediate depths by night and day. Activity recorded during 24 and 72 h periods indicated that fish were consistently active at night and inactive during the day. Activity data corresponded with point‐in‐time habitat use data, both of which indicated that fish were concealed amongst cover during the day and used open water habitats at night. 3. During summer, giant kokopu used higher water velocities, shallower depths and coarser substrata, particularly at night but also occasionally during the day relative to winter. Giant kokopu were active by both day and night in summer, although periods of activity were less defined and less predictable than during winter. 4. Adults used predictable home reaches at base‐flow, with most individuals repeatedly using of one or two cover locations within their ‘home’ reach. Reaches used by fish were relatively short (rarely exceeding 26 m) irrespective of season and always included a single pool‐riffle sequence. 5. Diel and seasonal behaviour of giant kokopu was generally comparable with that exhibited by other drift feeding fish species in small temperate streams. However, the nocturnal activity of giant kokopu contrasts with activity patterns in various salmonids, indicating that the impact of predation by different drift feeding fish may vary considerably.     

The effect of rainfall on tick challenge at Kyle Recreational Park,Zimbabwe

The effect of rainfall pattern on tick challenge was investigated at Kyle Recreational Park, Zimbabwe, from 1991 to 1992 using drag and removal plot methods to sample environmental tick density. The abundance of adults and nymphs of the brown ear-tick Rhipicephalus appendiculatus and larvae of the bont tick Amblyomma hebraeum was positively correlated with monthly rainfall, whereas no relationship with rainfall was revealed for larval R. appendiculatus, adults of the red-legged tick R. evertsi, or larvae of the blue tick Boophilus decoloratus. A comparison between 1991 (490 mm rainfall) and the drought year of 1992 (161 mm) revealed significant differences in the abundance of R. appendiculatus, A. hebraeum, and B. decoloratus. During the wet season, R. appendiculatus adults were 2–3 time more numerous in the environment during the higher rainfall year of 1991. A. hebraeum larval abundance exhibited a similar pattern to that of R. appendiculatus adults, but B. decoloratus larvae were more abundant in the drought year of 1992 during both the wet and dry seasons. Comparable tick abundance data collected at Kyle during the above-average rainfall years of 1975–1977 (mean = 1029 mm) were compared with tick challenge during the below-average rainfall years of 1991–1992 (mean = 326 mm). In grassland sand habitat and all habitats combined R. appendiculatus adults, nymphs, and larvae were much more abundant during the high rainfall years. In contrast, larvae of B. decoloratus were more numerous during the drier years. A. hebraeum larvae were also more abundant during the drier years. The strong positive correlation of adult R. appendiculatus abundance with rainfall and the coincidence of increased adult tick challenge with increased rainfall indicates that adult R. appendiculatus tick burden on hosts would be heaviest during the wet season and high rainfall years.     

Drift and upstream movement in Yuccabine Creek,an Australian tropical stream

Drift and upstream movement were monitored over 14 months in a seasonal upland tropical stream in northeastern Australia. There were distinct seasonal pulses in the drift with variable peak levels in the summer wet season and low more stable levels during the dry season. Drift density ranged from 0.36 to 3.98 animals per m³ (monthly mean = 1.26). There was no correlation between drift density and either benthic density or stream discharge. In the absence of catastrophic drift, drift was dispersive, not depletive in the wet season. A total of 121 taxa were caught in the 14 drift samples. Most taxa had nocturnal maximum drift levels with a peak immediately after sunset, a pattern apparently related to level of light and not temperature. Compensation for drift by upstream-moving nymphs and larvae was least during the wet season and increased during the dry season to a peak of 27% by numbers. Mean compensation was 8.2%. It is suggested that apart from in the wet season when an animal may drift substantial distances, most riffle animals will spend their larval lives in one small stretch of stream.     

A manipulative study of macroinvertebrate grazers in Hong Kong streams: do snails compete with insects?

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Seasonal variability of ichthyoneuston assemblage on the continental shelf and slope of the Southwest Atlantic Ocean,Brazil (20°–23°S)

Many marine organisms spend the early life history stages in neuston domain. Although the importance of ichthyoneuston, few studies were developed in the Southwest Atlantic Ocean. Therefore, this study aimed to improve the knowledge on ichthyoneuston of the Brazilian coast analyzing the vertical stratification and horizontal distribution of these organisms. Neuston samples were collected in daylight between 20°S and 23°S from February to April 2009 (late summer/early autumn; rainy season) and from August to September 2009 (late winter/early spring; dry season). Eggs of six taxa were identified: Anguilliformes, Engraulidae, Clupeidae, Synodontidae, Trichiuridae, and Maurolicus stehmanni (Sternoptychidae). The neustonic larval fish assemblage was composed by 40 families and 63 species. Mullidae and Myctophidae larvae were the most abundant in the rainy period while Mullidae and Mugilidae dominated in the dry season. Seasonal and spatial variation of larval fish assemblages in the neuston were structured by oceanographic features. The larval fish abundance on the outer slope stations may have been favored by the advection of an anticyclone that encompassed most of the study region during the cruise periods. In the rainy season, salinity and local depth structured the larval fish assemblage in a cross‐shelf gradient, while in the dry season the larval assemblage was structured around temperature and north‐south gradient. In the dry season, the cross‐shelf gradient was less pronounced mainly because of the low abundance and frequency of mesopelagic larvae. The low frequency and abundance of some species are probably related to the net avoidance of fish larvae during the day or dial vertical migration, as many species migrate to deeper layers during the day and ascend to neuston only at night. Nevertheless, the present study presented baseline information about the seasonal and spatial variation of the neustonic larval fish assemblage influenced by the oceanographic conditions in the Campos Basin. We recommend to additionally collect night samples to decrease larval escapement rates and to compare night versus day catches to further investigate the influence of daily migration in the neustonic larval fish in the area.     

Seasonal variation in the composition of fish assemblages in small Amazonian forest streams: evidence for predictable changes

1. The effects of seasonal inundation on the biology of fishes on floodplains of large Amazonian rivers are well studied. However, the small seasonal changes in headwater streams are generally considered to have little effect on fish assemblages. 2. In this study, we analysed seasonal changes in the species composition and abundance of fish in small Amazonian forest streams. We sampled fish with hand and seine nets in headwater streams in a 10 000 ha terra‐firme forest reserve near Manaus, Brazil. Each stream was surveyed at the end of the 2005 dry season, at the beginning of the 2006 rainy season and at the beginning of the 2006 dry season, by means of a standardized sampling effort. 3. The numbers of individuals and species caught were higher in the dry season, but rarefaction analyses indicated that greater species numbers could have been due simply to the larger number of individuals caught. 4. Between the dry and rainy season, the direction of changes in species composition in multivariate space varied among sites, especially for quantitative (abundance) data. However, the observed variation among sites was the less than expected if the directions of change were random. 5. Fish assemblages in the second dry season were more similar to those in the previous dry season than expected if changes in species composition among seasons were random. This indicates that a general seasonal pattern in fish assemblages can be detected, despite the existence of some erratic site‐specific changes. 6. Most of the species that showed large seasonal variations in density occupy temporary ponds during the rainy season, when much of the valley is inundated and pond networks form adjacent to streams. Short‐duration lateral migrations to these ponds may play an important role in the seasonal fish‐assemblage dynamics in Amazonian headwater streams. 7. Our results contrast with previous studies on small Amazonian streams, which have found little seasonal change in fish assemblages, and highlight the importance of the abundance of common species as an indicator of general fish assemblage structure in biological monitoring programmes.     

Nested species subsets, gaps, and discrepancy

Chemical cues from fish can alter the behaviour of stream invertebrates in experimental tanks but their effect in natural streams has received little attention. By adding brook trout (Salvelinus fontinalis) odour to a trout stream in the Rocky Mountains of Colorado, USA, we tested whether changes in the concentration of chemical cues from visually feeding predatory fish would alter the drift of mayfly nymphs (Ephemeroptera). Stream water was piped from stream-side tanks with (odour) and without (control) three brook trout to two locations in the stream 3.5 m upstream of drift nets at six replicate sites. Five-minute drift samples were collected downstream from odour and control pipes before, during and after the release of water from the tanks into the stream during both the day and night. Almost all drift occurred at night and consisted predominantly of Baetis bicaudatus nymphs. The odour manipulation had no measurable effect on Baetis drift during the day but statistical power was low. During the night, however, the drift of large (>0.65 mm head capsule width, HCW) Baetis nymphs decreased significantly during the odour addition compared to control drift. In contrast, the drift of small nymphs (≤0.65 mm HCW) increased both during and after the odour addition in comparison to control drift. Since the stream contains brook trout (0.04–0.18 m⁻²), and water from the stream (presumably containing fish odour) altered the behaviour of fishless-stream Baetis nymphs in another experiment, we conclude that the changes in Baetis drift density were a response to an increase in the concentration of fish odour in the stream. Furthermore, we were able to detect the effect within 5 min. of odour addition, indicating that mayfly behavioural response to trout odour was rapid. These results suggest that mayflies can distinguish different concentrations of trout odour in natural streams and that the response is size-specific, according to the relative risk of predation of large and small Baetis. Received: 12 May 1998 / Accepted: 23 October 1998     

Predator effects on prey population dynamics in open systems

Animal population dynamics in open systems are affected not only by agents of mortality and the influence of species interactions on behavior and life histories, but also by dispersal and recruitment. We used an extensive data set to compare natural loss rates of two mayfly species that co-occur in high-elevation streams varying in predation risk, and experience different abiotic conditions during larval development. Our goals were to generate hypotheses relating predation to variation in prey population dynamics and to evaluate alternative mechanisms to explain such variation. While neither loss rates nor abundance of the species that develops during snowmelt (Baetis bicaudatus) varied systematically with fish, loss rates of the species that develops during baseflow (Baetis B) were higher in streams containing brook trout than streams without fish; and surprisingly, larvae of this species were most abundant in trout streams. This counter-intuitive pattern could not be explained by a trophic cascade, because densities of intermediate predators (stoneflies) did not differ between fish and fishless streams and predation by trout on stoneflies was negligible. A statistical model estimated that higher recruitment and accelerated development enables Baetis B to maintain larger populations in trout streams despite higher mortality from predation. Experimental estimates suggested that predation by trout potentially accounts for natural losses of Baetis B, but not Baetis bicaudatus. Predation by stoneflies on Baetis is negligible in fish streams, but could make an important contribution to observed losses of both species in fishless streams. Non-predatory sources of loss were higher for B. bicaudatus in trout streams, and for Baetis B in fishless streams. We conclude that predation alone cannot explain variation in population dynamics of either species; and the relative importance of predation is species- and environment-specific compared to non-predatory losses, such as other agents of mortality and non-consumptive effects of predators. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Diversity of parasites in Cichlasoma amazonarum Kullander, 1983 during rainy and dry seasons in eastern Amazon (Brazil)

This study investigated the influence of the rainy and dry seasons on the parasite communities of Cichlasoma amazonarum (Kullander, 1983) in a tributary of the Amazon River system, northern Brazil. Of 112 fish examined, 95.5% were parasitized by Ichthyophthirius multifiliis, Piscinoodinium pillulare, Gussevia disparoides, Posthodiplostomum sp., Procamallanus (Spirocamallanus) inopinatus, and Echinorhynchus paranensis, as well as by the Glossiphoniidae leech. Ichthyophthirius multifiliis and P. pillulare were the dominant parasite species, with I. multifiliis the most prevalent and abundant; the leech (Glossiphoniidae) was the least prevalent and least abundant. Parasites presented an aggregate dispersion pattern and seasonal variations in infestation levels, influenced by the environmental conditions in the rainy season. Variation in the parasite dynamics created overall changes in the parasite communities, characterized by greater diversity, species richness and evenness during the rainy season. However, there was a high similarity (99.8%) in the parasite community structure between the rainy and dry seasons. During the dry season the hosts had predominantly 1–3 parasite species compared to 3–4 parasites in the rainy season. Some parasites in the eastern Amazon undergo population changes relating to seasonality. These results thus indicate the correct season to apply adequate prophylactic measures to reduce negative impacts of parasites in this wild ornamental fish when captured.     

Life histories, production dynamics and resource utilisation of mayflies (Ephemeroptera) in two tropical Asian forest streams

SUMMARY 1. A 2‐year study of the life histories, production dynamics and resource utilisation of five mayfly species was undertaken in two forest streams in Hong Kong [Tai Po Kau Forest Stream (TPKFS) and Shing Mun River (SMR)]. Afronurus sp. and Cinygmina sp. (Heptageniidae), Procloeon sp. and Baetiella pseudofrequenta (Baetidae), and Choroterpes sp. (Leptophlebiidae) were abundant in both streams and contributed more than 50% of the total mayfly populations. 2. All species had asynchronous larval development with recruitment occurring throughout the year. Mean annual production (all mayflies combined) was 3.1 and 2.0 g dry weight m^?2 year^?1 in SMR and TPKFS, respectively – the higher value at SMR reflecting greater mayfly densities – with more than 70% of production occurring during the wet season. Mayfly production varied between years, decreasing by 5% in TPKFS and 43% in SMR during 1996–97, reflecting lower densities of heptageniids relative to 1995–96. Annual biomass turnover rates (P/B) were high in both sites ranging from 27.2 to 94.6 in TPKFS (Cinygmina sp. and Procloeon sp.) and from 31.8 to 109.8 in SMR (Cinygmina sp. and B. pseudofrequenta). 3. Patterns of daily production in both streams showed that Afronurus sp., Cinygmina sp. and Choroterpes sp. were most productive during the wet season, while Procloeon sp. maintained high production levels throughout the year. The highest daily production of B. pseudofrequenta occurred during the wet season in TPKFS, but in the dry season at SMR. Temporal overlap in production and hence resource utilisation in both streams, calculated using the proportional similarity index (PS), ranged from 0.39 to 0.81. It was highest (0.63–0.81) between pairs of species of Heptageniidae and Baetidae, and lowest between Choroterpes sp. and other mayflies (0.39–0.61). No clear temporal segregation was observed among any species. However, when using the fraction of production attributable to each food, lower PS values were obtained for all species in both sites. In SMR, trophic segregation may have occurred between the two species pairs Procloeon sp.–Cinygmina sp. and Procloeon sp.–Choroterpes sp. (PS=0.17 and 0.03, respectively). 4. A combination of production data and information on the stable isotope signature of mayflies revealed that, during both the wet and dry seasons, more than 50% of total mayfly production in TPKFS was derived from autochthonous foods. In SMR, 68% of production was supported by allochthonous foods during the wet season, and 72% by autochthonous sources in the dry season. Considering that more than 70% of the total production occurred in the wet season, the trophic basis of mayfly production in SMR is mostly allochthonous (58%) while in TPKFS it is mainly of autochthonous origin (66%). The year‐round importance of autochthonous foods in shaded streams such as TPKFS is surprising, but the wet season contribution of allochthonous foods (especially in SMR) may have resulted from depletion of algal biomass during spates.     

Benthivorous fish reduce stream invertebrate drift in a large-scale field experiment

Drift as a low-energy cost means of migration may enable stream invertebrates to leave risky habitats or to escape after encountering a predator. While the control of the diurnal patterns of invertebrate drift activity by fish predators has received considerable interest, it remains unclear whether benthivorous fish reduce or increase drift activity. We performed a large-scale field experiment in a second-order stream to test if invertebrate drift was controlled by two benthivorous fish species (gudgeon Gobio gobio and stone loach Barbatula barbatula). An almost fishless reference reach was compared with a reach stocked with gudgeon and loach, and density and structure of the invertebrate communities in the benthos and in the drift were quantified in both reaches. The presence of gudgeon and stone loach reduced the nocturnal drift of larvae of the mayfly Baetis rhodani significantly, in contrast to the findings of most previous studies that fish predators induced higher night-time drift. Both drift density and relative drift activity of B. rhodani were lower at the fish reach during the study period that spanned 3 years. Total invertebrate drift was not reduced, by contrast, possibly due to differences in vulnerability to predation or mobility between the common invertebrate taxa. For instance, Chironomidae only showed a slight reduction in drift activity at the fish reach, and Oligochaeta showed no reduction at all. Although benthic community composition was similar at both reaches, drift composition differed significantly between reaches, implying that these differences were caused by behavioural changes of the invertebrates rather than by preferential fish consumption. The direction and intensity of changes in the drift activity of stream invertebrates in response to the presence of benthivorous fish may depend on the extent to which invertebrate taxa can control their drifting behaviour (i.e. active versus passive drift). We conclude that invertebrate drift is not always a mechanism of active escape from fish predators in natural streams, especially when benthos-feeding fish are present.