Nocturnal drift of mayfly nymphs as a post-contact antipredator mechanism

1. The predominantly nocturnal constrained drift of stream invertebrates is commonly regarded as a behaviour that avoids encounters with visually foraging fish in the water column. The alternative explanation, that drift peaks are caused by bottom-feeding, nocturnal predators, has rarely been tested.
2. We examined these hypotheses by collecting invertebrate drift in five streams in northern Finland: one with brown trout ( Salmo trutta , a drift-feeding fish), one with alpine bullhead ( Cottus poecilopus , a benthic fish), one with both species, and two fishless streams.
3. Drift by Baetis mayflies was aperiodic or slightly diurnal in both fishless streams on all sampling occasions. In contrast, drift was nocturnal in streams with trout and, to a lesser extent, in the stream with bullhead. Non-dipteran prey drifted mainly nocturnally in all streams with fish, whereas Diptera larvae were less responsive to the presence of fish.
4. In laboratory experiments, bullheads were night-active, causing a much higher frequency of drift by touching Baetis at night than during the day. Thus, increased nocturnal drift may serve to avoid both visual predators (a pre-contact response) and benthic fish (a post-contact response). In streams with bottom-feeding fish, nocturnal drift should be caused by increased drift by night rather than by reduced drift by day.     

Diel vertical movements by lotic mayfly nymphs under variable predation risk

1. The diel foraging periodicities of two grazing mayfly (Ephemeroptera) nymphs, Heptagenia dalecarlica and Baetis rhodani, under variable fish (European minnow) predation risk were examined in a series of laboratory experiments. 2. Heptagenia dalecarlica were almost exclusively nocturnal in their use of feeding areas on stone tops. There was a sharp increase in the proportion of nymphs out of refuge at nightfall, both in the control and fish-odour treatments. In a treatment with freely moving fish, H. dalecarlica decreased their activity during both the day and night. In feeding trials with three freely foraging minnows, nymphs were completely safe when they had access to refuges beneath stones, whereas almost half the nymphs were consumed when no refuges were available. 3. Baetis rhodani nymphs reduced their use of stone tops when exposed to four caged minnows or a freely moving fish, but this occurred both day and night. In feeding trials, B. rhodani were captured only while in the water column, and their mortality risk was independent of refuge availability. 4. It is suggested that because H. dalecarlica lack efficient post-encounter defences, they must rely on pre-encounter mechanisms to reduce the threat of fish predation. It appears that in order to dwell sympatrically with fish, H. dalecarlica have evolved a coexistence by hiding strategy. In contrast, Baetis are vulnerable to fish attacks only if they enter drift in daylight, and are thus able to remain on stone tops both during the day and at night.     

Effects of a predatory fish on a tropical detritus-based food web

In contrast to that for grazing systems, relatively little information exists for trophic cascades in detritus-based stream food webs, which are predominant in forested headwater streams. Predator–prey interactions are thought to be weak in these systems, but studies are very scarce, their results are equivocal, and they do not separate the effect of direct consumption from a behavioural response of shredders. We examined the effect of predatory fish on leaf litter breakdown in headwater tropical Australian streams at three levels: (1) the behavioural response of shredder species to predator presence as indicated by chemical cues; (2) the rates of leaf breakdown resulting from shredder activity; and (3) the relationship between shredder species richness and leaf breakdown rates. Our results suggest that predatory fish can have a trait-mediated effect on detritus-based food webs in streams, by reducing consumer activity. We identified reductions in short-term overall activity in response to the presence of predatory fish cues, comparable to those found for grazers. We also observed a visible, albeit statistically non-significant, reduction in consumption rates. Shredder species richness did not affect leaf breakdown rates, and fish presence did not modify this relationship or the differences in breakdown rates among species, suggesting that the overall reduction in leaf breakdown caused by fish presence is due to a reduction in activity in every species. Thus, our laboratory studies have shown that there can be a behavioural basis for trait-mediated trophic cascades linked to fish presence in detrital food webs in streams. However, the strength of fish effects depends on environmental circumstances, and field studies of litter breakdown in streams with and without predatory fish are required if we are to elucidate the ecological significance of our observations.     

The effects of extended photoperiods on the drift of the mayfly ephemerella subvaria McDunnough (Ephemeroptera : Ephemerellidae)

Field drift studies indicated that the nocturnal drift density of E. subvaria nymphs was greater in early May than in early November.Laboratory studies showed that the number of individuals appearing in the drift was a linear function of the duration of the preceding photoperiod. Nymphs had a greater propensity to drift when they were not in a state of active growth than when they were growing. The tendency of individuals in a single laboratory population to drift was observed to change under conditions of constant temperature and randomized photoperiod. This suggests that the shift was due to some internal physiological change rather than to an external cue.It is suggested that drift in E. subvaria functions as a method relocation from fast-water areas to slow-water pools and stream margins. Redistribution to these areas may reduce mortality incurred during spring run-off and during emergence.     

Effects of thermal regime on mayfly assemblages in mountain streams

The variation in thermal regime and elevation among streams in the Sawtooth Mountains of Idaho, USA was used to test hypotheses about forces structuring larval mayfly assemblages. Sites above and below lakes were included to maximize variation in thermal regime. Forty-five sites were sampled for mayfly larvae and their summer thermal regime was measured. Ordination methods were used to analyze variation in the mayfly assemblages. Principal components analysis showed that mayfly assemblages were strongly and consistently affected by lakes within the stream system, apparently through the effects of lakes on stream temperature. Redundancy analysis explained 51% of the variation in assemblages and identified maximum water temperature and elevation as strong predictors of mayfly assemblages. Elevation influenced assemblage structure independently of summer maximum water temperature, suggesting that air temperature or some other elevation-dependent feature is also important. As predicted by the River Continuum Concept, mayfly diversity increased with increasing maximum daily range in temperature. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: R. Bailey     

The role of trout in stream food webs: integrating evidence from field surveys and experiments

1. We evaluated the effects of brown trout on boreal stream food webs using field surveys and enclosure/exclosure experiments. Experimental results were related to prey preference of uncaged trout in the same stream, as well as to a survey of macroinvertebrate densities in streams with vs. without trout. Finally, we assessed the generality of our findings by examining salmonid predation on three groups of macroinvertebrate prey (chironomid midges, epibenthic grazers, invertebrate predators) in a meta-analysis. 2. In a preliminary experiment, invertebrate predators showed a strong negative response to trout, whereas chironomids benefited from trout presence. In the main experiment, trout impact increased with prey size. Trout had the strongest effect on invertebrate predators and cased caddis larvae, whereas Baetis mayfly and chironomid larvae were unaffected. Trout impact on the largest prey seemed mainly consumptive, because prey emigration rates were low and independent of fish presence. Despite strong effects on macroinvertebrates, trout did not induce a trophic cascade on periphyton. Uncaged trout showed a strong preference for the largest prey items (predatory invertebrates and aerial prey), whereas Baetis mayflies and chironomids were avoided by trout. 3. Densities of invertebrate predators were significantly higher in troutless streams. Baetis mayflies also were less abundant in trout streams, whereas densities of chironomids were positively, although non-significantly, related to trout presence. Meta-analysis showed a strong negative impact of trout on invertebrate predators, a negative but variable impact on mobile grazers (mainly mayfly larvae) and a slightly positive impact on chironomid larvae. 4. Being size-selective predators, salmonid fishes have a strong impact on the largest prey types available, and this effect spans several domains of scale. Discrepancies between our experimental findings and those from the field survey and meta-analysis show, however, that for most lotic prey, small-scale experiments do not reflect fish impact reliably at stream-wide scales. 5. Our findings suggest that small-scale experiments will be useful only if the experimental results are evaluated carefully against natural history information about the experimental system and interacting species across a wide array of spatial scales.     

New mayfly nymphs from the Jurassic of Northern and Northeastern China (Insecta: Ephemeroptera)

Four new species of mayfly nymphs, Furvoneta relicta sp. nov., Clavineta eximia sp. nov. (originally Mesoneta antiqua), Mesobaetis latifilamentacea sp. nov., and Stackelbergisca cylindrata sp. nov. are described from the Jurassic of China. All these species may have inhabited lakes rather than flowing water. The geological age and stratigraphical correlation of the nymph-bearing strata are briefly reviewed.     

The effect of the method of supplementary feed application on the feeding habits of carp (Cyprinus carpio L.) with regard to natural food in ponds

Gross primary production in Sathiar reservoir showed that it is comparable to and as high as in other eutrophic waters. The diel periodicity of photosynthesis which shows a peculiar decline in early afternoon, has been shown to be linked with photorespiration due to high oxygen tension. Besides temperature and transparency the trophogenic area appears to regulate gross primary production. In Sathiar nutrients are not limiting GPP and the main factors controlling it are linked with phytoplankton biomass, water level changes and therefore with precipitation. The maximum production density was always at the surface except in the months of December and January.     

The relative importance of lethal and non-lethal effects of fish on insect colonisation of ponds

1. We hypothesised that adult insects actively monitor potential habitats for the presence of fish by means of chemical cues and avoid sites that pose significant risks. This was examined by quantifying colonisation of insects in outdoor pools with no fish (controls), fish (direct predation effect) or caged fish (chemical predator cues).
2. A significant direct effect of predation was found, but no indirect effect (avoidance of chemical cue pools), on the total biomass of colonising insects. However, predatory insects avoided fish-cue pools, thus releasing non-predatory insects from predation. This resulted in significantly greater biomass of non-predatory insects in fish-cue pools than control pools.
3. Fish reduced the number of species of colonising insects in pools through predation. This negative influence of fish implies that caution is necessary when stocking wetlands and ponds with fish if the goal is to maximise biodiversity.
4. Our data suggest that although predatory aquatic insects may use chemical signals to assess the quality of potential habitats with respect to predation risk, direct predation is the main method by which fish affect insect assemblages in ponds. Because fish and invertebrate predators may both have strong effects on prey mortality, behavioural adjustment by insects to the actual predator regime within a habitat should be more important than avoiding colonisation of habitats with fish.     

Comparing predatory drillholes to taphonomic damage from simulated wave action on a modern gastropod

Marine drilling predation, in which the predator bores a hole through shelled invertebrate prey, plays a role in the structure of benthic communities. As drilling often leaves the prey shell otherwise undamaged, the resulting holes are also an excellent proxy for drilling predation pressure in the fossil record. Considering that a large number of predation studies focus on drilling predation in the fossil record, it is crucial that we are able to distinguish true drilling predation from taphonomy. The purpose of this study is to determine damage on Olivella biplicata shells, drilled by naticid gastropods, is distinguishable from taphonomically produced damage to these shells. In addition, the potential for preferential breakage due to either the presence or whether absence of a drillhole was investigated. Drilled and non-drilled O. biplicata shells were tumbled to simulate wave action and were checked at intervals to record accumulated damage. Drilled and non-drilled shells do not show a significant difference in damage accumulated while undergoing simulated wave action. Taphonomic damage is unlikely to be mistaken for drilling damage, due to the jagged, irregular appearance of taphonomically produced holes.     

The influence of a predatory ostracod,Australocypris insularis,on zooplankton abundace and species composition in a saline lake

The influence of the predatory ostracod Australocypris insularis on zooplankton abundances and species composition in an Australian saline lake was monitored over 8 months in aquaria and tank microcosms. Control, Predator (double A. insularis Control densities) and No Predator (all A. insularis removed) treatments were set up in small 11.5l aquaria and large 883.5l tanks. Microcosm salinities were manipulated to accord with seasonal lake salinities and ranged from 35 to 110 gl^-1. Densities of the calanoid copepods Calamoecia clitetellata and C. salina, the small ostracods Diacypris compacta and D. dietzi, and juvenile ostracods (excluding A. insularis) were significantly reduced by A. insularis predation in the aquaria experiments. Daphniopsis sp. was only present (in low numbers) in the No Predator aquaria. Results from the large tank experiments were not as clear-cut, due to difficulties in the manipulation of predator numbers. Differing salinity tolerances of predator and prey may confound the predation impact of A. insularis.     

To fear or to feed: the effects of turbidity on perception of risk by a marine fish

Coral reefs are currently experiencing a number of worsening anthropogenic stressors, with nearshore reefs suffering from increasing sedimentation because of growing human populations and development in coastal regions. In habitats where vision and olfaction serve as the primary sources of information, reduced visual input from suspended sediment may lead to significant alterations in prey fish behaviour. Here, we test whether prey compensate for reduced visual information by increasing their antipredator responses to chemically mediated risk cues in turbid conditions. Experiments with the spiny damselfish, Acanthochromis polyacanthus, found that baseline activity levels were reduced by 23 per cent in high turbidity conditions relative to low turbidity conditions. Furthermore, risk cues elicited strong antipredator responses at all turbidity levels; the strongest antipredator responses were observed in high turbidity conditions, with fish reducing their foraging by almost 40 per cent, as compared with 17 per cent for fish in clear conditions. This provides unambiguous evidence of sensory compensation in a predation context for a tropical marine fish, and suggests that prey fish may be able to behaviourally offset some of the fitness reductions resulting from anthropogenic sedimentation of their habitats.     

The influence of density on frequency-dependent food selection: a comparison of four experiments with wild birds

We compare the results of four experiments, conducted at different times and with different protocols, that explored the relationship between frequency-dependent selection and prey density in wild birds feeding on artificial populations of coloured baits. One (experiment 4) used pastry baits that differed only in the presence or absence of a red stripe, and this experiment provided no evidence for any kind of selective behaviour. The other three experiments used green and brown baits, and they all provided evidence for a trend towards increasing anti-apostatic selection with high densities (>100 baits m^–2). However, one of these (experiment 3) provided no evidence for frequency-dependent selection at low densities (0.5–20 baits m^–2), while the other two experiments concurred in suggesting a trend towards increasing apostatic selection with low densities (down to 2 baits m^–2). Together, these experiments both support and qualify the published findings of experiment 1 that frequency- dependent selection by wild birds on bait populations is modified by density. Experiment 4 indicates that frequency-dependent selection may break down entirely if bait types are too similar, while experiment 3 indicates that some details of this trend with density will depend either on the protocol used or on exogenous changes in the birds’ feeding behaviour. Received: 1 September 1999 / Accepted: 22 March 2000     

The biology of ant-like jumping spiders (Araneae, Salticidae): prey and predatory behaviour of Myrtnarachne with particular attention to M. lupata from Queensland

Myrtnarachne is a genus of ant-like salticids. Eight species were observed feeding, in nature, in Australia, Kenya, Malaysia and Sri Lanka, on varied types of insects but not ants. The behaviour of M. lupata , from Australia, was studied in the laboratory. Predatory sequences were found to differ considerably from those of typical salticids. Attacking by lunging instead of leaping and the pronounced use of pre-attack tapping are especially noteworthy. The unusual behaviour of M. lupata when preying on insects is consistent with maintenance of ant mimicry. Myrmarachne lupata also preys on the eggs of other spiders which it extracts from their nests. The males of many species have very large chelicerae, and the large chelicerae of M. lupata males influence the course of predatory sequences, with insects and with eggs.     

The smell of danger: chemical recognition of fish predators by the invasive crayfish Procambarus clarkii

1. Since avoiding predation can compromise animal fitness, prey are expected to respond to different predator species with an intensity appropriate to the level of risk. In fresh waters, the threat of predation is typically assessed by chemical cues, in particular by odours released by either injured/disturbed conspecifics (conspecific alarm odour) or predators (predator odours). Here, we used the most widely distributed crayfish in the world, the invasive North American Procambarus clarkii, to investigate the relative effectiveness of odours emitted by fish predators compared with conspecific alarm odour. We also tested whether P. clarkii is able to discriminate between fish predators of which it has ‘experience’ (either recent, via introduction to the same water body, or old, by sharing a native range), as well as between fish predators that pose low or high risk. 2. The study was carried out on introduced populations of P. clarkii from two sites, characterised by different fish assemblages: the Malewa River (a tributary of Lake Naivasha, Kenya) and Lake Trasimeno (Italy). Laboratory experiments consisted of three sequential phases (‘water’, ‘food’ and ‘smell’ phases) and five treatments. Treatments differed in the odour presented during the smell phase, i.e. no odour (plain water) and odours from either injured conspecifics or three fish species per site. Crayfish from the Malewa River population were confronted with the odours of largemouth bass (Micropterus salmoides), common carp (Cyprinus carpio) and tilapia (Tilapia zillii) (all introduced to Lake Naivasha but absent from the Malewa River), and those from the Lake Trasimeno population with the odours of the introduced largemouth bass and carp and the native chub (Squalius cephalus). Largemouth bass is the only predator that imposes a high risk to crayfish, and it also shares its native range with P. clarkii. We analysed the time spent by crayfish feeding, in locomotion and in adopting a raised or lowered posture. A reduction in the time spent feeding and in locomotion, and an increase in the time spent in the lowered posture were considered to indicate alarm. 3. Crayfish from both populations responded with a more pronounced reduction in feeding to conspecific alarm odour rather than to predator odours. Crayfish from the Malewa River reacted with the same intensity to the odours of the three fish species tested, whereas, in Lake Trasimeno, the odour of largemouth bass was significantly more threatening than the odours of the other two species. 4. Procambarus clarkii seems to perceive a general fish odour that alerts it to possible predation risk without the need of either a direct recent experience or via sharing a common native range. However, where they coexist with fish, crayfish become able to distinguish among species, adapting the intensity of their response to the effective risk. Our results confirm the relatively high learning capacity of P. clarkii reported in previous studies and suggest the existence of mechanisms that make predator recognition particularly efficient in this extraordinarily successful invader.     

The influence of flood cycle and fish predation on invertebrate production on a restored California floodplain

Although floodplains are known to be tightly controlled by the flood cycle, we know comparatively little about how flooding influences predators and their consumption of secondary production, particularly in highly seasonal floodplains typical of Mediterranean climates. In this study, we investigate how the seasonal dynamics of a central California floodplain influence the timing and magnitude of fish predation and the abundance and composition of invertebrates. For 3 years (2000–2002), we compared changes in abundances and size distributions of invertebrates through the flood season (January–June) with seasonal changes in the abundance of larval and juvenile fishes. Using diet analysis of fishes and manipulative feeding experiments with fishes in field enclosures, we link specific changes in invertebrate populations directly to feeding preferences of seasonally abundant fish. Early in the flood season prior to March, we found little influence of fish predation, consistent with the near absence of larval and juvenile fishes during this period. Coinciding with the midseason increase in the abundance of larval and juvenile fishes in April, we found significant declines in zooplankton abundance as well as declines in the size of zooplankton consistent with fish feeding preferences. Our results were consistent with results from feeding enclosure experiments that showed that fish rapidly depressed populations of larger cladocerans with much less effect on smaller cladocerans and calanoid copepods. At the end of the flood season, zooplankton abundances rapidly increased, consistent with a switch in the feeding of juvenile fish to aquatic insects and subsequent fish mortality. We also found that zooplankton biomass on the floodplain reached a maximum 2–3 weeks after disconnection with the river. We suggest that floodplain restoration in this region should consider management strategies that would ensure repeated flooding every 2–3 weeks during periods that would best match the peaks in abundance of native fishes. Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users.     

The biology of Tauala lepidus,a jumping spider (Araneae: Salticidae) from Queensland: display and predatory behaviour

Abstract

The display and predatory behaviour of Tauala lepidus Wanless, an abundant salticid in north Queensland rainforests, was investigated in the laboratory and, to a lesser extent, in nature. T. lepidus leapt and walked into alien webs to catch spiders and insects, and was captured and fed on by other spiders. Females ate each other’s eggs. During intraspecific interactions, a complex repertoire of displays was used. Courtship versatility occurred, each individual male having a conditional strategy of different behaviours depending on whether the female is at or away from her nest, and whether she is adult or subadult. Yet other combinations of displays occurred during male-male and female-female interactions. Apparently, pheromones on nests and draglines of females released male courtship. Abdomen twitching, a behaviour common to the display repertoires of many salticids, was an especially complex and pervasive behaviour of T. lepidus. T. lepidus also twitched its abdomen when it contacted alien webs and preyed on other species of spiders. The behaviour of T. lepidus is compared to that of Jacksonoides queenslandica Wanless, a species from the same group (Astieae).     

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.