Predator detection and avoidance by lotic mayfly nymphs of different size

We studied antipredatory responses of lotic mayfly (Baetis) nymphs in a factorial experiment with four levels of fish presence: (1) a freely foraging fish (the European minnow,Phoxinus phoxinus), (2) a constrained fish, (3) water from a fish stream, (4) water from a fishless stream. LargeBaetis nymphs drifted mainly during night-time in treatments involving either the chemical or actual presence of fish, whereas no diel periodicity was observed when the water was not conditioned with fish odour. The response was strongest when the fish was uncaged, which suggests that visual or hydrodynamic cues are needed in addition to chemical ones for an accurate assessment of predation risk. Fish presence had no effect on the drift rates of small nymphs. Instead, they increased their refuge use in the presence of a live fish. Chemical cues alone did not have any effect on the refuge use of any of theBaetis size classes. Our results indicate active drift entry by mayfly nymphs. Because predation pressure is spatially and temporally variable, nymphs must sample the environment in order to locate predator-free areas or areas with low predation risk. Drifting should be the most energy-saving way to do this. To avoid the risk from visually feeding fish, large individuals can sample safely (i.e. enter drift) only at night-time, while the small ones can also do this safely during the day. We suggest that, contrary to some earlier assumptions, mayfly drift is not a fixed prey response. Instead,Baetis nymphs are able to assess the prevailing predation pressure, and they adjust their foraging behaviour accordingly.     

Macro-zooplankter responses to simulated climate warming in experimental freshwater microcosms

1. We report data collected from 48 replicated microcosm communities created to mimic plant‐dominated shallow lake and pond environments. Over a 2‐year period, the microcosms were subjected to warming treatments (continuous 3 °C above ambient and 3 °C above ambient during summer only), a nutrient addition treatment and the presence or absence of fish. We tracked macro‐zooplankter dynamics, censusing cladoceran populations at the species level, copepods at the order level and ostracods as a class. 2. Responses to warming were subtle. Cladoceran diversity and overall abundance were not significantly affected by warming, although measures of community evenness increased. Warming effects on patterns of population trajectories tended to be strongly seasonal and most apparent during periods of pronounced increase. Populations of the prevalent cladocerans, Chydorus sphaericus and Simocephalus vetulus, displayed idiosyncratic patterns, with evidence in the case of S. vetulus for a negative relationship between warming and body‐size at maturity. Copepod populations were reduced in size by warming, but those of ostracods increased. 3. The effects of the nutrient addition and fish treatments were strong and consistent, interacting little with warming effects in statistical models. Zooplankter abundance tended to be the highest in the fish‐free microcosms receiving additional nutrient inputs and lowest when fish were present and no nutrients were added. Both treatments reduced cladoceran diversity and community evenness. 4. We suggest that warming, independently, is unlikely to supplant the effects of changing nutrient loading and fish predation as the major driver of zooplankter dynamics in shallow lakes and ponds. Moreover, in the situations where warming was of significant influence in our experiment, the distinction between summer‐only warming and year‐around warming was blurred. This suggests that warming effects were most pervasive during the summer, at the upper end of the temperature spectrum.     

Ingestion rate of the burrowing mayfly Hexagenia limbata as determined with14C

The effects of nymph size and temperature on the ingestion rate ofHexagenia limbata were investigated using¹⁴C labelled algae mixed into sediment. Ingestion rate increased with nymph length and temperature. Daily ingestion rates of large 19 mm nymphs burrowing in silt averaged 58 and 192% of the dry body weight at 16° and 21 °C, respectively. Ingestion of seston due to filter feeding in artificial burrows was insignificant compared to that ingested by burrowing nymphs.     

Consequences of tadpole shrimp predation on mayflies in some Californian ponds

SUMMARY. 1. During two, 1-month studies, the abundance of mayfly nymphs, Callibaetis californicus Banks, in small ponds that contained the tadpole shrimp, Triops longicaudatus (LeConte), often was significantly lower than in ponds without tadpole shrimps. Mayfly abundance was either positively correlated or not significantly associated with that of predacious insects.
2. Larger mayfly nymphs were a greater fraction of the population in ponds with Triops than in ponds without tadpole shrimps. Between 2 and 3 weeks after ponds were flooded, the smallest mayfly nymphs were absent from ponds in which T. longicaudatus densities were >5 individuals per square metre during the summer and >15 individuals per square metre during the autumn.
3. In prey-choice trials, T. longicaudatus consumed predominantly small C. californicus nymphs (mean headwidth ± 1 SD: 0.44 ± 0.05 mm). After encounters with tadpole shrimps, small mayfly nymphs moved relatively shorl distances, were easily overtaken, and were, consequently, more susceptible to tadpole shrimps than larger-sized nymphs.     

Response of macroinvertebrates to warming,nutrient addition and predation in large-scale mesocosm tanks

There is increasing concern about the effect of climate change on aquatic systems. We examined changes in macroinvertebrate communities caused by increased temperature (3°C above ambient during summer only and continuous 3°C above ambient all year round), influences of fish (Gasterosteus aculeatus L.) and addition of nutrients (nitrogen and phosphorus) in 48 large-scale (3000 l) tanks over a 2 year period. While numbers of Isopoda, Chaoborus, Corixidae, Ephemeroptera, Notonectidae and Odonata were reduced by the presence of fish, nutrient addition caused isopods, corixids, mayflies and odonates to increase in abundance. Impacts of temperature increase were surprisingly low, with only gastropods increasing in heated tanks, suggesting that, overall abundances of most macroinvertebrate taxa will not be severely affected by the predicted temperature rise. To determine if taxa were sampled representatively during the experiment, net sweep samples taken towards the end of the experiment were compared with final macroinvertebrate abundances when the complete contents of each tank were harvested. We found that net sweeping is an appropriate semi-quantitative method for most taxa in mesocosm tanks. However, mites, coleopteran adults and larvae, dipterans and Chaoborus were not adequately sampled. This might explain why we could not detect any treatment effects of temperature, fish or nutrients on mites, coleopterans and dipterans and calls for different sampling techniques for these taxa, especially in ponds with vegetation stands.     

Do nymphs of Parameletus chelifer and P. minor (Ephemeroptera) reduce mortality from predation by occupying temporary habitats?

Summary The possible effect of predation on the habitat selection of nymphs of the two mayfly species Parameletus chelifer Bengtsson and P. minor (Bengtsson) was investigated for different habitats of a North Swedish boreal river. In spring, these two species migrate to river margins and subsequently, in different proportions, to adjacent seasonal ponds. Feeding rates on mayfly nymphs and mosquito larvae by some dytiscid beetles and larvae, a corixid waterbug and a caddisfly larva were studied in the laboratory. In the seasonal pond, the consumption rate of invertebrate predators on Parameletus nymphs was below that expected from their relative abundance, and mosquito larvae were preferred. At the river margin, where mosquito larvae were absent, Parameletus nymphs were consumed in agreement with their relative abundance. The absolute abundance of macroinvertebrate predators was similar at the river margin and in the pond. Fish predators visited the seasonal pond only during a short time of flood, while at the river margin their presence was continuous. In the seasonal pond, minnow and roach consumed nymphs of both P. chelifer and P. minor in a lower proportion than expected. At the river margin, nymphs of P. chelifer were consumed by minnow and ten-spined stickleback in a higher proportion than expected. Nymphs of P. minor were consumed according to expectance. Imagines of P. minor emerged early, and nymphs were equally abundant in the river and in the seasonal pond. In contrast, almost all nymphs of P. chelifer migrated to the seasonal pond. The later emergence of P. chelifer would probably result in a very high mortality from predation if it remained at the river margin.     

Temperature preferences and tolerances of three fish species inhabiting hyperthermal ponds on mangrove islands

The fish species Cyprinidon artifrons, Floridichthys carpio, and Gambusia yucatana inhabit shallow mangrove ponds off the coast of Belize. Portions of these ponds experience a diurnal temperature change from 26 °C at night to 40 °C and above during midday. Repeated field observations indicate Cyprinidon prefer the warmer (and much larger) portions of the ponds whereas the other two species stay in the cooler areas.The hypothesis that temperature is serving as a cue for partitioning within the ponds was supported by laboratory thermal gradient tests in which Cyprinidon preferred temperatures clearly higher than the other two species.The critical thermal maximum (CTM) was determined for the three species using members that had been acclimated to either a daily cycling temperature similar to that for the ponds, or to the mean of the 24-hour cycle (30 °C). Cyprinidon acclimated to the cycling temperature had a CTM of 45.5 °C, which apparently sets a new record for fish CTM. Acclimation to a constant 30 °C lowered the CTM to 43.7 °C. Floridichthys and Gambusia acclimated to the cycled temperature had CTMs of 43.9 and 43.3 °C respectively, and 42.5 and 42.6 °C for those acclimated to 30 °C.All three species appear to have the ability to tolerate the high temperatures throughout the ponds but only Cyprinidon utilize the whole pond during the day. This may help to explain the large populations of Cyprinodon found in these mangrove ponds compared to the other species.     

Effects of leaping grey mullet Liza saliens (Osteichthyes, Mugilidae) in the macrophyte beds of oligohaline Mediterranean coastal lagoons

Some mugilid fish are known to enhance small phytoplankton in freshwater macrophyte-free environments due to zooplankton depletion. This suggests that they may have negative effects on natural macrophyte beds of freshwater and oligohaline lagoons due to phytoplantkon enhancement. To test this hypothesis, we compared the ecosystems of control enclosures that contained no fish with those of enclosures stocked with Liza saliens at two different densities. The occurrence of L. saliens at a density of 321±92.42 kg ha^–1 reduced cladoceran density, depleted epiphytic chironomid larvae, enhanced mayfly nymphs and cyclopoid copepods and reduced the organic matter content of sediment, all in comparison with control enclosures. At a density of 673±42.04 kg ha^–1, L. saliens reduced total zooplankton density, depleted epiphytic and sediment dwelling chironomid larvae and enhanced mayfly nymphs. The organic matter contents of sediment was not affected. These results showed that L. saliens was very effective in reducing zooplankton density even when macrophyte biomass was high. However, these effects do not affect phytoplankton density, probably because zooplankton was dominated by species with low filter-feeding rates and macrophytes depleted nutrients.     

Ontogenetic Shift in Efficacy of Antipredator Mechanisms in a Top Aquatic Predator,Anax junius (Odonata: Aeshnidae)

The ability of prey to escape predation often lies in the occurrence and efficacy of their predator avoidance and antipredator behaviors, which are often coupled with specialized morphology. How the use and efficacy of these behaviors change throughout ontogeny may be indicative of the vulnerability and ecological roles these animals experience throughout their lives. We examined the antipredator behavior of a large dragonfly nymph, Anax junius, from a historically fishless pond where these animals have traditionally been classified as top predators. These dragonfly nymphs displayed a series of distinct aggressive antipredator behaviors when grasped that involved stabbing with lateral and posterior spines and seizing with labial hooks. Larger (older) nymphs displayed these aggressive behaviors significantly more than smaller (younger) animals in simulated predation trials. During encounters with live larval salamander predators (Ambystoma tigrinum), all large nymphs, but only 12.5% of small nymphs successfully escaped predation attempts by the amphibians through the use of antipredator behavior. Large nymphs were also significantly more active than smaller nymphs in the presence of salamander larvae. Despite often being considered top predators in fishless ponds, our study demonstrates that their true role is more complex, depending on ontogeny and body size, and that effective antipredator behavior is likely necessary for survival in these systems.     

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.     

Habitat-Specific variation in eyespot size of a benthic cladoceran

Summary Pigmented eyespot size of the benthic cladoceran Simocephalus exspinosus was measured in individuals sampled from four freshwater ponds that differed in the extent of visually-oriented predation. In ponds with such predation (from fish, salamander larvae, and dragonfly nymphs), eyespot size was found to be significantly smaller, relative to body size, than in a pond without visually-oriented predation. Reduction in pigmented eyespot size may represent an adaptation to reduce vulnerability to predation.     

Egg envelopes of Baetis rhodani and Cloeon dipterum (Ephemeroptera, Baetidae): a comparative analysis between an oviparous and an ovoviviparous species

A comparative ultrastructural investigation on the eggshell (vitelline and chorionic envelopes) has been carried out in the nymphs of two mayfly species encompassed into the Baetidae, namely Baetis rhodani and Cloeon dipterum. During oocyte differentiation in the meroistic telotrophic ovarioles, gametes are connected to discrete nurse cells by trophic cords. In B. rhodani, each ovariole contains several eggs arranged in sequence, whereas in C. dipterum each contains a single egg. Follicle cells are competent for vitelline and chorionic envelope synthesis. Baetis rhodani is an oviparous species and the chorion is fairly thick, formed by an alveolate endochorion and a fibrillar exochorion delimited by a honey‐comb roof. Cloeon dipterum stands out among Ephemeroptera for being ovoviviparous. In B. rhodani, ovulation starts in the older nymphs with dark wing‐pads, whereas in C. dipterum choriogenesis ends in the imaginal stage. Here the chorion is very thin and laid eggs hatch almost immediately, allowing the larvulae to move out. The maturation of a single egg per ovariole is synchronized with the achievement of the adult stage. The acquisition of ovoviviparity has led to remarkable changes in the ovariole organization along with a simplification of the eggshell structure.     

Balancing risks? Responses and non-responses of mayfly larvae to fish and stonefly predators

Summary In a series of laboratory experiments we examined the hypothesis that larvae of stream mayflies would respond to the presence of two different types of predators in such a way as to minimize their risk of being consumed by each. Positioning of larvae (whether they frequent the top, sides, or bottom of stones) of Baetis tricaudatus and Ephemerella subvaria was altered by the presence of predaceous stoneflies (Agnetina capitata) with a larger proportion of the population occurring on the upper surfaces, where the probability of encountering the predator was lowest. The presence of a benthivorous fish (Cottus bairdi) had no significant effects on positioning of the mayfly larvae. Lack of fish effects may reflect an inability of the mayflies to detect or respond to sculpins, or alternately may indicate that sculpins do not normally present a important predation risk for these mayflies. Failure of mayfly prey to account for fish predators when responding to the presence of stoneflies appcars to explain facilitation previously observed between stoneflies and sculpins.     

Effects of hydroperiod and predation on a Mississippi River floodplain invertebrate community

 The objective of this study was to determine if pond permanence and vertebrate predation (by fish and waterfowl) affect invertebrate community structure in the mudflat habitat of floodplain ponds. Invertebrate communities were studied for 1 year in four Mississippi River floodplain ponds with different hydroperiods. Pond 1 experienced five dry periods, pond 2 experienced four, pond 3 dried once, and standing water remained in pond 4 for the entire year. Vertebrate predator exclusion treatments (all access, no access, small-fish access and cage controls) were placed in all ponds. As pond duration increased, predatory invertebrate richness and abundance increased while overall invertebrate richness and abundance decreased. With the exception of the cladoceran Diaphanosoma, all commonly encountered taxa were strongly affected by pond permanence in terms of abundance, biomass and, generally, individual biomass. Taxa were nearly early divided between those that were more abundant in less permanent ponds and those that were more abundant in longer-duration ponds. Invertebrate taxa richness, abundance, and total biomass were lower in the all-access treatment than in the treatments that restricted predator access, and these effects were stronger in the more permanent ponds. In general, there were no significant differences in responses to the treatments with small-fish access and no access. These results support models that predict relatively weak effects of predation in frequently disturbed habitats. Received: 30 May 1995 / Accepted: 21 June 1996     

Distribution and ecology of Demospongiae from the circalittoral of the islands of the Aegean Sea (Eastern Mediterranean)

Size-selective predation by fish is often considered to be a primary driver of seasonal declines in large-bodied Daphnia populations. However, large Daphnia commonly exhibit midsummer extinctions in ponds lacking planktivorous fish. A number of empirical and theoretical studies suggest that resource competition and its interaction with nutrient enrichment may determine variable dominance by large Daphnia. Low resource levels may favor competitive dominance by small-bodied taxa while large Daphnia may be favored under high resource conditions or following a nutrient/productivity pulse. Nutrient enrichment may also influence the strength of invertebrate predation on Daphnia by affecting how long vulnerable juveniles are exposed to predation. We investigated these hypotheses using an in situ mesocosm experiment in a permanent fishless pond that exhibited seasonal losses of Daphnia pulex. To explore the effects of nutrient enrichment, Daphnia plus a diverse assemblage of small-bodied zooplankton were exposed to three levels of enrichment (low, medium, and high). To explore the interaction between nutrient enrichment and invertebrate predation, we crossed the presence/absence of Notonecta undulata with low and high nutrient manipulations. We found no evidence of competitive reversals or shifts in dominance among nutrient levels, Daphnia performed poorly regardless of enrichment. This may have been due to shifts in algal composition to dominance by large filamentous green algae. Notonecta had significant negative effects on Daphnia alone, but no interaction with nutrient enrichment was detected. These results suggest that Daphnia are not invariably superior resource competitors compared to small taxa. Though predators can have negative effects, their presence is not necessary to explain poor Daphnia performance. Rather, abiotic conditions and/or resource-based effects are probably of greater importance.     

Low Temperature Tolerance of Pacu, Piaractus mesopotamicus

Pacu, Piaractus mesopotamicus (=Colossoma mitrei), is a South American warm water fish species found in the temperature range of 15–35°C. The culture of a warm water species in temperate regions demands knowledge on its temperature requirements. Pacu introduction into the Israeli fish culture system is being considered. Temperature range in the region is 8–33°C, thus the minimum winter water temperatures might be a limiting factor. To determine what is the minimum temperature pacu would tolerate, and hence which overwintering operations in warm-temperate regions are required for this warm water species, low temperature tolerance tests in the laboratory and observations in the field were carried out. Laboratory experiments reducing temperature by 1–3°C per day were carried out with fish of 150–200g, about the size pacu reach after one culture season. The field observations compared survival of two-year-old pacu of 1.3kg mean weight overwintered in outdoors and in greenhouse ponds. For one-year-old fish 7.5°C was found to be the lower temperature tolerance limit. Two-year-old fish withstand short exposures to this temperature rather well and their lower tolerance limit might be lower. This indicates that in warm-temperate regions pacu should survive in outdoors ponds. In this case some loss of weight should be expected, and suspension of feeding when temperature drops below 16–18°C is recommended to avoid wasting feed that the fish will not consume anyway. To be in the safe side, inflow of the warmest available water into the ponds is recommended if maximum water temperature drops to 10°C or below. Overwintering in greenhouses or other heated facilities would be recommended if an exceptionally cold winter is expected and for regions with lower winter minimum temperatures.     

Effect of larval fish and nutrient enrichment on plankton dynamics in experimental ponds

The hypotheses that larval fish density may potentially affect phytoplankton abundance through regulating zooplankton community structure, and that fish effect may also depend on nutrient levels were tested experimentally in ponds with three densities of larval walleye, Stizostedion vitreum (0, 25, and 50 fish m^–3), and two fertilizer types (inorganic vs organic fertilizer). A significant negative relationship between larval fish density and large zooplankton abundance was observed despite fertilizer types. Larval walleye significantly reduced the abundances of Daphnia, Bosmina, and Diaptomus but enhanced the abundance of various rotifer species (Brachionus, Polyarthra, and Keratella). When fish predation was excluded, Daphnia became dominant, but Daphnia grazing did not significantly suppress blue-green algae. Clearly, larval fish can be an important regulator for zooplankton community. Algal composition and abundance were affected more by fertilizer type than by fish density. Inorganic fertilizer with a high N:P ratio (20:1) enhanced blue-green algal blooms, while organic fertilizer with a lower N:P ratio (10:1) suppressed the abundance of blue-green algae. This result may be attributed to the high density of blue-green algae at the beginning of the experiment and the fertilizer type. Our data suggest that continuous release of nutrients from suspended organic fertilizer at a low rate may discourage the development of blue-green algae. Nutrient inputs at a low N:P ratio do not necessarily result in the dominance of blue-green algae.     

The impact of predation by burbot (<Emphasis Type="Italic">Lota lota</Emphasis> L.) on the macroinvertebrate community in the littoral zone of a large lake

We studied the effects of predation by juvenile burbot (Lota lota) on the macroinvertebrate community in mesocosm experiments in the stony littoral zone of Lake Constance, a large prealpine lake in Central Europe. Although the growth data of the burbot suggest that the benthivorous fish exerts strong predation pressure on the invertebrate community, the predicted level of consumption is only poorly reflected by changes in biomass and abundance of most invertebrate prey taxa. High exchange rates of the prey between the mesocosms and the ambient littoral environment apparently masked the true predation effects of fish. Also, life-history events such as hatching or synchronised emergence of larvae led to temporal effects that obscured the impact of predation. However, for the dominant prey organism, the freshwater amphipod Gammarus roeseli, direct lethal effects appeared when its migration was limited. When exchange with the ambient littoral zone was possible, abundance and biomass of G. roeseli were unexpectedly high in the mesocosms stocked with burbot, indicating behavioural responses to fish presence. During the experiment, the burbot gained ca. 15% in length and about 60% in body mass. According to stomach content analyses at the end of the experiment, the ingested prey consisted mainly of relatively large and abundant invertebrates. Our study indicates that predation by juvenile burbot should be an important factor in structuring the benthic invertebrate community in terms of qualitative and quantitative effects.     

Temperature and oxygen in an Everglades alligator pond

The Everglades is a large subtropical marsh ecosystem. Ponds, maintained through the activity of alligators, are abundant and ecologically important components of this system. Temperature in a 18 x 12 m study pond was controlled primarily by ambient air temperature, with significant temporal lags because of thermal resistance caused by plant growth. Temperature changed seasonally being 10° to 15°C lower in summer than winter. Diurnal temperatures fluctuated 7°C in summer and 4°C in winter. Maximum temperature exceeded 34°C. Dissolved oxygen fluctuated between 50 to 85% saturation in winter to supersaturation in summer. During the dry season water levels fell and oxygen fluctuated markedly, for example from 4 to 200% saturation within a diurnal cycle.The relation of water temperature to air temperature was similar to other reported ponds but continued fluctuation in winter differed from some results from other ponds. The distinct seasonality has important effects on biological components. Fluctuation of dissolved oxygen can result from high animal density and plankton in the dry season and can lead to massive mortality of aquatic animals. The physio-chemical conditions of Everglades alligator ponds are controlled by a combination of seasonal temperature and insolation, water level changes and biologic factors such as fish density, bird predation and alligator activity.     

The effect of mermithid parasitism on predation of nymphal Baetis bicaudatus (Ephemeroptera) by invertebrates

We investigated how infection by the mermithid nematode Gasteromermis sp. affected predation on its nymphal mayfly host, Baetisbicaudatus, by two invertebrate predators – the stonefly nymphs of Kogotusmodestus and the caddisfly larvae of Rhyacophilahyalinata. Predation trials and behavioral observations were conducted in stream-side, flow-through experimental chambers. When parasitized and unparasitized prey were offered in equal numbers, K. modestus consumed significantly more parasitized than unparasitized nymphs. R. hyalinata consumed equal numbers of both prey types. Behavioral observations of foraging K.␣modestus on parasitized and unparasitized prey suggested that the increased consumption of parasitized nymphs was due to differences in the behavior of infected mayflies in response to the predator. Specifically, parasitized nymphs drifted less often to escape an approaching predator (non-contact encounters) compared to unparasitized nymphs, which increased the number of contact encounters and attacks that occurred between K.␣modestus and parasitized prey. Because all hosts are castrated, these behavioral alterations affect only the fitness of the parasite, which is killed along with its host by invertebrate predation. We present a number of hypotheses to explain why the parasite causes increased predation on its host. These include the large size of the parasite affecting the sensory abilities of the host, the larger energetic costs of escape behavior for parasitized individuals, and natural selection from fish predation against drifting behavior by parasitized individuals. Received: 27 May 1996 / Accepted: 30 September 1996