Differential learning rates of chemical versus visual cues of a northern pike by fathead minnows in a natural habitat

We stocked 39 juvenile pike, Esox lucius, into a previously pike free pond which contained a population of approximately 78 000 fathead minnows, Pimephales promelas. Fathead minnows sampled prior to pike stocking did not show a stereotypic fright response to either visual or chemical cues from pike. After stocking pike, we sampled minnows every two days for a period of two weeks. Minnows sampled six days after stocking still did not show a fright response to the sight of a pike, but those sampled eight days after stocking did exhibit a significant fright response, indicating that acquired predator recognition based on vision occurred between six and eight days. Minnows sampled two days after stocking did not show a fright response to chemical cues of a pike. Those sampled four days after did, however, exhibit a significant fright response, indicating that acquired predator recognition based on chemical cues occurred between two and four days. These data indicate that acquired predator recognition occurs very rapidly and that the rate of learning of predator identity differs for chemical versus visual cues.     

Chemical alarm signals increase the survival time of fathead minnows (Pimephales promelas) during encounters with northern pike (Esox Lucius)

We tested the hypothesis that exposure to a conspecific alarmpheromone improves survival of fathead minnows (Pimephales promelas)during staged encounters with an unfamiliar predator (northernpike: Esox luaus). Minnows exposed to the alarm pheromone survived39. 5% longer than controls. This difference in survival timeappeared to result not from direct inhibition of the pike butrather from some aspect of the minnows' antipredator behavior.Minnows exhibited significant increases in both shoaling andshelter use after exposure to the alarm pheromone. For controlminnows, the degree of shoaling was positively correlated withsurvival time, suggesting that increased shoaling is an effectiveantipredator response. This study provides the first directexperimental evidence that chemical alarm signals in fishesimprove survival of receivers.     

Effects of exposure to predatory cues on territorial behaviour of male fathead minnows

We conducted a laboratory study to determine if male fathead minnows, Pimephales promelas, altered their territorial behaviour associated with reproduction in response to combinations of visual and chemical cues from northern pike, Esox lucius. We introduced the following stimuli to a territorial male: a brick (control), fathead minnow alarm pheromone, a pike fed brook stickleback, Culea inconstans, or a pike fed fathead minnow. The territorial behaviour of males did not change when the control was added. Male minnows experiencing threat from pike fed stickleback significantly reduced the frequency at which they performed three territorial behaviours, but, within 12 h, had returned to pre-exposure activity levels. Male minnows subjected to alarm pheromone alone and to pike fed fathead minnow significantly reduced their territorial behaviour, abandoned their nests, and did not return to pre-exposure levels of activity after 24 h. We suggest that because risk of predation triggers prolonged decreases in territorial defense, it may affect competition between nesting males and female mate choice. We conclude that fathead minnows can assess the severity of predatory threat and adjust their reproductive behaviour accordingly.     

Effects of social structure on reproductive activity in male fathead minnows (Pimephales promelas)

The selection of alternative reproductive phenotypes is oftenthought to be the result of physiological state, with smallindividuals forced energetically to postpone the allocationof resources to reproduction. However, for male fathead minnows(Pimephales promelas), we show that seasonal reproductive activityis modulated by social status. In enclosure and pond experiments,small males advanced their reproductive condition, held nesting territories, and spawned earlier in the reproductive seasononly when large males were absent or removed from the population.Since differences in the timing of reproduction among smallmales were not size- or condition-dependent, the common explanationfor the selection of alternative reproductive phenotypes, basedon state-dependence, is insufficient. In the absence of large,socially dominant individuals, small males produced comparablenumbers of offspring as the treatment with large males, although the offspring of these uninhibited small males were smallerat the end of the growing season than the young of large males.Thus, interactions among conspecifics may account for muchof the phenotypic diversity observed within and among naturalfathead minnow populations, through their direct and indirecteffects on growth, recruitment and survival.     

Foraging among cannibals and kleptoparasites: effects of prey size on pike behavior

The northern pike (Esox lucius) is an important and selective piscivorethat chooses smaller prey than predicted from energy / timebudgets. In a laboratory experiment, we investigated pike predatorybehavior to explain this selectivity. Northern pike feedingon different prey sizes in aquaria were observed when foragingalone, when in the presence of chemical cues from similar-sizedor larger conspecifics, and when in the presence of conspecifics thatwere allowed to interact with the focal pike. The results showthat prey handling time increases with prey size and that theduration of manipulating and handling prey inflicts a risk ofexposure to cannibals and kleptoparasites on the pike. Therefore,the risk of falling victim to cannibals or kleptoparasites increaseswith prey size. Attracting and experiencing intraspecific interactorscan be regarded as major fitness costs. Chemical cues from foragingconspecifics have only minor effects on pike foraging behavior.Furthermore, the ability to strike and swallow prey head first improvespike predatory performance because failing to do so increases handlingtime. Our findings emphasize the increasing potential costswith large prey and explain previous contradictory suggestionson the underlying mechanisms of behavior, selectivity, and trophiceffects of northern pike predation.     

The effect of physical condition and shoalmate familiarity on proliferation of alarm substance cells in the epidermis of fathead minnows

Ostariophysan fishes have specialized epidermal alarm substance cells (ASCs). In this study, metabolic cost to the production and maintenance of these cells was demonstrated. Fathead minnows Pimephales promelas were maintained on high or low food rations and, to test for the effect of social context on the number of ASCs, they were held with either zero or two shoalmates. Shoalmates were familiar (from the focal fish's shoal), or unfamiliar (from a shoal separated by 1 km from the focal fish's shoal). After 16 days, epidermal thickness, number of mucus cells and number of ASCs were all significantly greater for minnows on the high ration than for those on the low ration. Within the high ration group, physical condition was positively correlated with epidermal thickness and ASC number. The shoalmate treatment had no measurable effect on the epidermis. It was concluded that food availability determines investment into ASCs, inferring a trade-off between the cost of ASCs and the fitness benefits they accrue.     

Direct measurements of metabolism, activity and feeding behaviour of pike, Esox Zucius L., in the wild, by the use of heart rate telemetry

Heart rate telemetry records of up to 5 days duration were obtained from pike living in Lochs Kinord and Davan, Scotland. Applying metabolic rate correlations it was found that mean metabolic rate (R) was 1.5 times standard metabolic rate (R_a), The fish rarely worked near their metabolic limits. Activity metabolism (R_a) was much higher than estimates based on mean swimming speed and comprised up to 10% of R. Most activity metabolism was the result of localized bursts of activity. Less than 10% of activity showed evidence of oxygen debt. Specific Dynamic Action or feeding metabolism (R_f) comprised 15–25% of R. Food intake estimated from heart rate was 1.5% wet body weight day⁻¹, consumed in the form of small items captured during the day and digested during the afternoon and night.     

Fathead minnows (Pimephales promelas) learn to recognize chemical stimuli from high-risk habitats by the presence of alarm substance

We exposed fathead minnows (Pimephales promelas) to water fromone of two distinct habitat types (an open water site or a vegetatedcover site in the same stream) that we mixed with either alarmsubstance or a distilled water control. Upon subsequent exposuresto the habitat waters alone, minnows showed a fright responseto the habitat water that they received in conjunction withalarm substance but not to the other habitat water. These resultsindicate that minnows can learn to recognize high-risk habitatsbased on the association of habitat specific chemical cues withalarm substance. The ability to recognize these habitats couldpotentially lower the minnows' risk of predation. These resultsprovide evidence of a long-term benefit to receivers of a chemicalalarm signal.     

Environmental factors regulate the effects of roach Rutilus rutilus and pike Esox lucius on perch Perca fluviatilis populations in small boreal forest lakes

In this study of 18 small boreal forest lakes, the effects of abiotic and biotic factors (roach Rutilus rutilus and pike Esox lucius) on various population variables of perch Perca fluviatilis were examined. As a single variable, the gillnet catch per unit effort (CPUE) of R. rutilus was negatively related to the mean mass of small (< 200 mm) and the growth rate of young (1–2 years) P. fluviatilis. The mean mass of large (≥ 200 mm) P. fluviatilis was the highest at intermediate CPUE of R. rutilus. Redundancy analysis including environmental factors and P. fluviatilis population variables suggested that ‘predation–productivity–humus' gradient affected P. fluviatilis populations by decreasing the CPUE and mean mass of small individuals but increasing these variables of large individuals. The CPUE of R. rutilus and lake area had a negative effect on small and a positive effect on large P. fluviatilis growth rate. In small boreal forest lakes, P. fluviatilis populations are affected by the partially opposite forces of competition by R. rutilus and predation by E. lucius, and the intensity of these interactions is regulated by several environmental factors.     

Changes in the fish community and water quality during seven years of stocking piscivorous fish in a shallow lake

SUMMARY 1. Piscivores (annual stocking of 1000 individuals ha^?1 of 0+ pike and a single stocking of 30 kg ha^?1 of large 20–30 cm perch) were stocked in seven consecutive years in a shallow eutrophic lake in Denmark. The stocking programme aimed at changing food‐web structure by reducing zooplanktivorous and benthivorous fish, with resultant effects on lower trophic levels and ultimately water quality. 2. The fish community and water quality parameters (Secchi depth, concentrations of total phosphorus, chlorophyll a and suspended solids) were monitored between 1996 and 2000 and relationships were evaluated between predatory fish and potential prey and between zooplanktivorous or benthivorous fish and water quality parameters. In addition, potential consumption of piscivorous fishes was calculated. 3. The density of fish feeding on larger zooplankton or benthos (roach >15 cm, crucian carp >15 cm) declined distinctly during the study period. This effect was attributed to predation by large (>50 cm) pike. Based on scale readings, we cautiously suggest that the stocking of 0+ pike boosted the adult pike population to produce an unexpected impact in later years. Conversely, no direct impact of stocked 0+ pike was detected on 0+ roach. 4. A major decline in the recruitment strength of 0+ roach was observed in 2000. A combination of (i) the indirect effect of large pike preying on adult roach, with negative effect on roach reproduction and (ii) the direct predation effect of 0+ pike and/or 1+ and 2+ perch recruited in the lake, provides the most likely explanation of this phenomenon. 5. A marked increase in Secchi depth in 2000 and declining trends in suspended solids, chlorophyll‐a and total phosphorus concentrations were observed. These changes may also be attributable to changes in the fish community, although the relationships were not straightforward. 6. This 7‐year study indicates that piscivorous fish may be a significant structuring force in shallow eutrophic lakes, suggesting that stocking piscivores can increase predation pressure on cyprinids. However, the general lack of impact of 0+ pike points to the need of refining current stocking practices in several countries across Europe.     

Conflict between sexes in the water strider, Gerris lacustris: a test of two hypotheses for male guarding behavior

We studied the effect of operational sex ratio on female reluctanceand male persistence to mate as well as on the length of copulationand postcopulatory guarding in Gerris lacustris by adding fivesurplus males or females to the basin with a pair in tandem.In the control treatment, a pair alone was tested. Accordingto the copulatory guarding hypothesis (CGH), males should prolongmating and guard females in the presence of surplus males. Accordingto the convenience polyandry hypothesis (CPH), females shouldshow lower levels of resistance to prolonged mating in the presenceof surplus males because the mating male protects the femaleagainst harassment from other males. As expected on the basisof both the CGH and CPH, mating (copulation + guarding) averagedlonger in the male-biased treatment. The behavior of males andfemales during mating suggested that both hypotheses hold true:females showed less resistance to prolonged mating (as predictedfrom CPH), and male behavior suggested stronger efforts to stayon the female when surplus males were present (as predictedfrom CGH). Comparisons of the treatment with surplus femaleswith the results from the mating pair without surplus individualssuggested that the capabilities of water striders in tandemto assess the sex of nearby nonmating striders are limited.     

Song and immunological condition in male barn swallows (Hirundo rustica)

Male secondary sexual characters may have evolved as intra-or intersexual signals of male phenotypic or genetic quality.In birds, singing performance may have the function to honestlyreveal health and vigor of individual males. Infectious diseasesand poor body conditions would therefore be expected to negativelyinfluence singing performance. Since bird pathogens are knownto elicit both a humoral and a cell-mediated immune response,it can be predicted that a negative relationship exists betweensinging performance and activity of the immune system. Thisprediction was tested for the first time in this correlationalstudy. The relationships between song rate and features andhematological variables (concentration of leukocytes in peripheralblood, ratio of gamma-globulins to total plasma proteins, bloodcell sedimentation rate, hematocrit) and body condition wereanalyzed in a population of bam swallows (Hirundo rustica).Song rate was negatively correlated with lymphocyte concentrationand with the ratio of gamma-globulins to plasma proteins. Spectrographicanalysis showed that features of song were not significantlycorrelated with hematological variables or body condition. Thelevel of circulating testosterone was not correlated with songrate nor hematological variables. This study is the first toshow a correlation between a bird's singing performance andhematological profile and suggests that song rate of male barnswallows may reflect their health status. Song in this speciesmight thus have evolved because it allows prospecting femalesto assess aspects of phenotypic and/or genetic quality of potentialmates     

Brood discrimination, nest mate discrimination, and determinants of social behavior in facultatively quasisocial beetles (Nicrophorus spp.)

In this study we investigated ecological determinants of socialityin burying beetles (Nicrophorus spp.), potential conflicts ofinterest among reproductive females, and the effects of nestingfailure and costs of fighting on cooperation. Burying beetlesare known to form monogamous pairs when exploiting small vertebratecarcasses. More complex social behavior in this group is poorlyunderstood. We conducted experiments in which one or two females(N. defodiens, N. orbicollis) were provided small or large carcasseson which to breed. On large but not on small carcasses, twofemales often formed cooperative breeding associations (jointlyprepared a carcass and fed young). In N. defodiens, but notN. orbicollis, two females produced a larger brood than singlefemales on large carcasses. In both species, the reproductiveoutput per female was less for two than for one female. Thepresence of a second female did not decrease the preparationtime of a carcass (discovery of resource to egg hatch). Conflictwas evident between females. Trials employing females of similarsize were more likely to result in injury than trials usingfemales of dissimilar size (N. tomentosus, N. defodiens, N.orbicollis). In N. tomentosus, those associations that persistedthe longest resulted in the fewest injuries. After care of youngwas initiated, conflict among familiar nest mates was not observed.There was no evidence that breeding females could discriminatebetween brood; use of a genetic marker (N. orbicollis) demonstratedthat females fed related and unrelated young alike. Femalesof similar size (high potential cost of fighting for the dominantindividual) were not more likely to form cooperative breedingassociations than females of dissimilar size (low cost of fightingfor dominant). Females of a species subject to a high rate ofnest failure (N. defodiens) were more likely to cooperate thanfemales of a species with a low rate of nest failure (N. orbicollis).It is argued that limited reproductive opportunities, difficultyin controlling rivals' access to a large carcass, and the superabundantlarval food supply represented by a large carcass, but not kinselection, have contributed to the evolution of cooperativebehavior in this group. In addition, we hypothesize that beetlesmight initially tolerate consexual rivals on large carcasseswhen there is a high likelihood of nesting failure, therebyavoiding potentially costly conflicts.     

Effects of Ligula intestinalis on habitat use, predation risk and catchability in European minnows

The frequency of infection with Ligula intestinalis (Cestoda) in European minnow Phoxinus phoxinus , in a subalpine lake in Eastern Norway, did not differ between vegetated shoreline, exposed shoreline and non-vegetated localities >50 m from the shoreline. There was no difference in the vertical distribution of infected and uninfected individuals. The frequency of infection was higher among minnows in brown trout Salmo trutta stomachs than among those obtained by gillnets and minnow traps, suggesting that brown trout selectively preyed on infected minnows. Prevalence of infection decreased with increasing fish size, probably due to selective mortality among parasitized individuals. Within a given length-class, minnows captured by different sections of multi-mesh gillnets showed a significant increase in the frequency of infection with increasing mesh-size. Apparently, parasitized individuals had a higher catchability in gillnets due to increased girth caused by the plerocercoid in the body cavity. This may partly explain why the observed prevalence of infection was several times higher among minnows captured by gillnets than by minnow traps.     

The benefits of stealing from a predator: foraging rates, predation risk, and intraspecific aggression in the kleptoparasitic spider Argyrodes antipodiana

In the trade-off between food and safety, the role of aggressiveintraspecific interactions has not been extensively examined.Here I present information on this system using a kleptoparasiticspider, Argyrodts antipodiana, and its host spider and potentialpredator, Eriophora pustulosa. A. antipodiana can feed eitherat a potentially dangerous site (the hub of its host's web withthe host), or at a relatively safe site (on food bundles aroundthe edge of the host's web). I found that A. antipodiana cangain food very quickly when feeding with the host, apparentlyby exploiting the host's ability to digest the prey. Thus A.antipodiana follows predictions based on foraging models inthat it accepts a higher predation risk at the hub because ofthe higher food payoff. A. antipodiana also aggressively competesfor access to more food. However, aggressive competition increasesthe predation risk from the host, especially at tile hub wherethe host is very close. Consequently, A. antipodiana modifiesits level of intraspecific aggressiveness in accordance withits position on the web: at the hub, where the cost of aggressionis high (due to predation risk), A. antipodiana reduces itsaggressiveness, but it is aggressive away from the hub whencompeting for food bundles. The ability of A. antipodiana tochange interaction intensity as a function of its position onthe web enables it to exploit a rich, but risky, food sourceand provides a new angle for examining food and safety tradeoffs in light of intraspecific competition for food     

Experiments and a model examining learning in the area-restricted search behavior of ferrets (Mustela putorius furo)

Area-restricted searches have been described as important componentsof the foraging behavior of many organisms. It is unclear, however,whether individual foragers can use learning to fine-tune theirsearches, or even whether these searches are efficiently performed.I used a simulation model to make qualitative predictions aboutsearch behavior in a laboratory system. The simulation modelindicates that the sinuosity and path length of searches stronglyaffect search efficiency. The model predicts that, for a rate-maximizingforager, path length should increase and search sinuosity shoulddecrease as prey become less clumped. Foraging animals may thereforebe selected to learn the path length and sinuosity of searchesin response to changing degrees of dumping of prey. These predictionswere tested in a laboratory system involving ferrets (Mustelaputorius furo) foraging for oil-drop "prey items." Search pathschanged in a graded manner to experimental manipulations ofthe dumping of prey. As predicted by the model, ferrets learnedto perform longer and less sinuous search paths as prey becameless clumped. This study provides the first evidence that area-restrictedsearch behavior is learned and can be fine-tuned to efficientlyexploit different spatial distributions of food.     

Black-and-white plumage in male pied flycatchers (Ficedula hypoleuca) reduces the risk of predation from sparrowhawks (Accipiter nisus) during the breeding season

Bright colors in birds might signal that they are undesirableas prey (aposematic), an idea that has been difficult to test.When stuffed pied flycatchers Ficedula hypoleuca are exposedto migrating sparrowhawks Accipiter nisus in spring or in autumn,the hawks attack cryptic females more often than bright males.To achieve better statistical control and to assess whethermale plumage also reduces predation risk in the breeding seasonand in the nesting habitat of the pied flycatcher, I placedpairs of male and female flycatcher mounts in similar positionsnear 22 nests of sparrowhawks. The hawks attacked mainly femalemounts, verifying that the preference is real. The sparrowhawkscaught at least 19 live pied flycatchers; 12 young, 5 adultmales, 1 adult female, and 1 female or young. Hawks that caughtan adult male seemed to prefer attacking female mounts. I discussthree interpretations of these results, suggesting that black-and-whitemale flycatchers may benefit from being a novel and aberrantprey, at least early in the breeding season.     

Interacting effects of predation risk and male and female density on male/female conflicts and mating dynamics of stream water striders

We used a factorial experiment to examine interacting effectsof male density, female density, and sunfish (predation risk)on mating dynamics of the stream water strider (Aquarius remigis).Many of our results corroborated earlier studies on the isolatedeffects of each factor on mating behavior. The effect of eachfactor, however, depended on the other factors. For example,in low density pools, predation risk decreased male generalactivity, male/female harassment rates, mating activity, andmating duration and increased the large male mating advantage.At higher densities, however, water striders apparently enjoyed"safety in numbers" and did not alter their mating dynamicsin response to the presence of predators. Female activity showeda particularly complex response to male density and fish. Whenmales were scarce, fish caused females to reduce their activity.However, when males were abundant, fish increased female activity,probably because fish decreased male activity thus releasingfemales from harassment by males. The three treatment factorsalso had interacting effects on male mating success. In theabsence of fish, when females were scarce, increased male densityresulted in a decrease in mean male mating success; however,when females were abundant, increased male density enhancedmean male mating success. In contrast, in the presence of fish,male density had little effect on male mating success. Manyof the observed mating patterns can be explained by the effectsof ecological and social factors on male/female conflicts; thatis, on male harassment of females and female reluctance to mate.     

The effect of group size manipulations on the foraging behavior of black-tailed prairie dogs

Individuals foraging in large groups are thought to benefitbecause they are better able to detect and avoid predators.As a consequence, individuals in groups can adopt more risky,but rewarding, foraging behaviors without exposing themselvesto excessive danger. I experimentally manipulated the size ofblack-tailed prairie dog (Cynomys ludovicianus) groups to determineif individuals in large groups do forage in a more risky manner.I found that prairie dogs foraged more alertly and in less riskylocations (nearer to burrows, nearer to the center of the group,and in shorter vegetation) when group size was reduced. Effectsof group-size reductions were reversed when removed individualswere replaced, and persisted for at least three weeks in experimentswhere group size was permanently reduced. My results provideevidence that the relationships between group size and bothalertness and risk-place foraging are causal.     

Sexual selection for large male size in a polyandrous butterfly: the effect of body size on male versus female reproductive success in Pieris napi

Sexual selection theory predicts that the larger sex shouldbe that for which fitness increases at the faster rate withsize. In butterflies, as in most invertebrates, females areusually the larger sex, but previous comparative analysis hasshown that relative male size increases with female polyandryamong butterflies. In agreement with this pattern, males arelarger than females in the strongly polyandrous green-veinedwhite butterfly, Pieris napi L., and in this article we assessthe size dependence of reproductive success in both sexes. Inan experiment where virgin males and females were released inthe field, we found no strong association between size and malemating success. However, laboratory experiments showed thatthere was a strong correlation between size and the ejaculatethat the male delivered to the female at mating and that largeejaculates delayed female remating for a longer time comparedto small ejaculates. Moreover, female P. napi utilize male-derivednutrients received at mating to increase their fecundity. Hence,large males sire more offspring both by way of donating morenutrients to female egg production and by way of delaying femaleremating (given that the last male to mate with the female willfather most of the offspring). Laboratory experiments showedthat the association between size and fecundity was low, ornonexistent, among P. napi females allowed to mate only once.However, weak size dependence was found for polyandrous females.We hypothesize that size dependence of female fecundity maybe especially weak among polyandrous butterflies because a fundamentalsource of variation in fecundity relates to their ability tofind nutrient giving males, an ability which may be unrelatedto female size. According to this hypothesis there is a causalassociation between weak size dependence of female fecundityand polyandry, and a strong size dependence of male reproductivesuccess that may underlie the comparative pattern of positivecorrelation between relative male size and polyandry.