Life histories and habitat selection in Daphnia: divergent life histories of D. magna clones differing in phototactic behaviour

To test the hypothesis of co-adaptation of life histories and daytime vertical distribution (vertical migration behaviour) in Daphnia, life history characteristics were analysed for two positively, three negatively, and four intermediately phototactic Daphnia magna clones. Clones with different phototactic behaviour were found to have divergent life history strategies, with positively phototactic clones being good exploiters under the non-limiting conditions provided in the laboratory, i.e. low density (1 ind./1), high food concentration (6,5–7 10⁵ Scenedesmus cells/ml, restored daily) and high temperature (20° C). They realized a high intrinsic rate of increase at a small adult body size through rapid development, at a cost of producing small neonates. Negatively and intermediately phototactic clones had larger adult body sizes, and produced larger neonates that were more starvation-resistant than those of positively phototactic clones. Selection for high intrinsic rate of increase in intermediately phototactic clones was mediated through the production of large clutches.     

Environmental and physiological factors influencing phototactic flight of Bemisia tabaci

Abstract. Experiments were conducted in a glasshouse and in the laboratory to determine the environmental and physiological parameters that affect flight behaviour of Bemisia tabaci (Gennadius) (Homoptera: Aleyrodidae). The number of whiteflies taking off and exhibiting a positive response to sky light in the glasshouse was greatest from 08.30 to 10.00 hours. During peak flight activity less than 5% of the population engaged in phototactic orientation. Temperature was the best single predictor for the phototactic response, accounting for 75% of the variability in whitefly ascent. Sex ratios were determined for individuals remaining on poinsettia, Euphorbia pulcherrima (Willd.), and for individuals that exhibited phototactic orientation; both groups deviated from an expected 1:1 ratio. Males were more prevalent on the plants (1:0.76), whereas females were much more prevalent (1:3.02) among the whiteflies responding to sky light. A higher percentage of the females displaying a phototactic response contained eggs when compared to females remaining on poinsettia (87 v 65%). The two groups of females did not differ significantly in their weights, but males that remained on the plants were heavier than males that responded to sky light. Mark–recapture studies and experiments where phototactic individuals were removed from the population established that the response was short-lived. Whiteflies exhibiting a phototactic response in the glasshouse were more likely to exhibit long-duration, phototactic flights in a vertical flight chamber when compared to individuals that remained on poinsettia (80.7 ± 6.7 v 36.0 ± 5.8% phototactic response; 7.0 ±3.2 v 0.7±0.2min flights). There was also less deviation in flight across the horizontal plane among the individuals that exhibited a positive response to sky light in the glasshouse in comparison to individuals that remained on their host. However, initial rates of climb were not significantly different between the two groups of whiteflies.     

Flight behaviour of Bemisia tabaci in a vertical flight chamber: effect of time of day, sex, age and host quality

Abstract. The free-flight behaviour of Bemisia tabaci (Gennadius) (Homoptera: Aleyrodidae), the sweet potato whitefly, was investigated in a vertical flight chamber. A mercury-vapour lamp presented from above induced a phototactic flight response. Although flight propensity was comparable from 06.00 to 19.00 hours, flight duration was maximum between 06.00 and 10.00 hours. Males flew longer than females and their mean flight duration remained constant throughout the day. Females flew longer from 06.00 to 13.00 hours than from 13.00 to 19.00 hours. Both sexes were capable of sustaining flight for more than 2h, although less than 5% of those tested did so.
Flight activity also was influenced by age and by host quality. The propensity to take off, proportion exhibiting phototactic orientation and flight duration varied with the age of the whitefly. Host quality influenced the timing of flight behaviour. Whiteflies reared on senescing plants exhibited greater take-off rates and initiated longer phototactic flights up to 4 days following adult eclosion when compared to individuals reared on vegetative plants. Thereafter, individuals reared on vegetative plants exhibited greater response levels. Whiteflies reared on vegetative plants weighed more and survived longer than did individuals reared on senescing plants.
Whiteflies that responded to the overhead light initially exhibited a strong photokinetic and phototactic response. Over the course of the flight, these responses declined and flight instability increased, as indicated by an overall decrease in the mean rate of climb, accompanied by an increase in the variability of this parameter and an increase in horizontal displacement. Although males and females displayed similar flight characteristics, females exhibited a greater rate of climb than did males, and for both sexes, individuals that flew longer than 25 min had a greater rate of climb than did individuals that flew for less than 25 min.     

Behavioral,morphological, and physiological traits associated with migratoryBemisia tabaci (Homoptera: Aleyrodidae)

Bemisia tabaci, that were flown for varying lengths of time (1 min-2.5 h), exhibited behavioral, morphological, and physiological differences. When whiteflies that were engaging in phototactic orientation in a vertical flight chamber were presented with a visual cue simulating their host plant (550-nm narrowband interference filter), 76% landed on this cue within three presentations. Another 18% of the whiteflies displayed an intermittent attraction to the host cue, and 6% displayed a response that, historically, has been considered to be indicative of migration. These individuals failed to respond to the target until they had flown for at least 15 min. These three categories of response were observed in both sexes, in all flight-capable individuals from 1 to 5 days old and in two groups of whiteflies that were exhibiting distinct behaviors prior to our tests (i.e., they were either settled on poinsettia or engaging in phototactic orientation). There was a trend for males that engaged in long-distance flight to have smaller wing dimensions than males that engaged in shorter flights; no trend was evident in female whiteflies. High levels of vitellogenin and vitellin did not inhibit flight activity. In fact, we generally found higher levels of egg proteins in long-distance fliers and in premigratory individuals than in short-duration fliers and settled individuals.     

Hygropreference behaviour and humidity detection in the yellow‐spined bamboo locust,Ceracris kiangsu

Water is a critical resource for all terrestrial insects. The yellow‐spined bamboo locust Ceracris kiangsu Tsai (Oedipodidae: Orthoptera), matures, copulates and reproduces in the summer and adults are observed mud‐puddling in human urine‐contaminated materials on hot days. Besides obtaining NaCl and NH₄HCO₃, the locust may also compensate for water deficiency. In a dual‐choice test in the laboratory, adults prefer to remain on wet rather than dry filter paper at high temperature (36 °C). The females are more likely to move towards and stay on wet filter paper than males. The adults chew wet filter paper in the laboratory at higher temperature and the females consume more than the males. Moreover, foam plastic containing an aqueous solution of an insecticide, bisultap, results in greater mortality in females than a dry formulation, both in the laboratory and the field. The water content of the frass from males is significantly lower than that from females and may partially explain the stronger hygropreference behaviour in females. Other possible reasons for sexually different hygropreference are also discussed. The removal of the adult antennae substantially reduces discrimination between wet and dry filter paper and mortality resulting from wet bisultap formulations in the laboratory. Hence, antennae mediate, at least partially, hygropreference behaviour.     

The effect of carbon dioxide enrichment on apparent stem respiration from <Emphasis Type="Italic">Pinus taeda</Emphasis> L. is confounded by high levels of soil carbon dioxide

Huey and Slatkin’s (Q Rev Biol 51:363–384, 1976) cost–benefit model of lizard thermoregulation predicts variation in thermoregulatory strategies (from active thermoregulation to thermoconformity) with respect to the costs and benefits of the thermoregulatory behaviour and the thermal quality of the environment. Although this framework has been widely employed in correlative field studies, experimental tests aiming to evaluate the model are scarce. We conducted laboratory experiments to see whether the common lizard Zootoca vivipara, an active and effective thermoregulator in the field, can alter its thermoregulatory behaviour in response to differences in perceived predation risk and food supply in a constant thermal environment. Predation risk and food supply were represented by chemical cues of a sympatric snake predator and the lizards’ food in the laboratory, respectively. We also compared males and postpartum females, which have different preferred or “target” body temperatures. Both sexes thermoregulated actively in all treatments. We detected sex-specific differences in the way lizards adjusted their accuracy of thermoregulation to the treatments: males were less accurate in the predation treatment, while no such effects were detected in females. Neither sex reacted to the food treatment. With regard to the two main types of thermoregulatory behaviour (activity and microhabitat selection), the treatments had no significant effects. However, postpartum females were more active than males in all treatments. Our results further stress that increasing physiological performance by active thermoregulation has high priority in lizard behaviour, but also shows that lizards can indeed shift their accuracy of thermoregulation in response to costs with possible immediate negative fitness effects (i.e. predation-caused mortality).     

Ultrastructure of the epidermis of Meara stichopi (Platyhelminthes,Nemertodermatida) and associated extra-epidermal bacteria

This preliminary mechanistic model of normal swimming and phototactic behaviour in individual Daphnia was constructed using data and assumptions based on experiments and observations. Swimming under constant light intensity is characterized by short periods of upward movements alternating with equal periods of downward movements. Two oscillators are proposed that generate these phases in swimming. Unexpected shifts in depth, as observed in D. magna and D. longispina, are also present in the swimming of the computer daphnid and thus seem to be inherent to the underlying mechanism. As in real daphnids, during relative decreases in light intensity of low velocity, positive phototactic upward swimming is stepwise. With increasing velocity in the change in light, these steps disappear. When the model is triggered by a natural increase in light at dawn, a small downward movement results. Migration distance can be increased to commonly found depths of migrating Daphnia by the introduction of a fish exudate factor into the model, which enhances the phototactic response. Since attenuation of light in the water affects the phototactic swimming response, it also influences migration distance. The results of model calculations agree quite well with an empirical relationship between Secchi disc depth and amplitude of diel vertical migration in a number of lakes.     

Response of Eretmocerus eremicus to skylight and plant cues in a vertical flight chamber

In the southwestern United States, Eretmocerus eremicus (Hymenoptera: Aphelinidae) is a native parasitoid of the Bemisia complex (Homoptera: Aleyrodidae). However, little information currently exists on its potential as a biological control agent of whiteflies or on the factors that influence its tendencies to disperse. In this study, we examined the flight behavior of male and female E. eremicus in response to skylight (here simulated by a mercury-vapor lamp) and plant cues (a 550-nm filtered light) in a vertical flight chamber. Approximately 90% of the parasitoids took off in response to the skylight cue. Both sexes were capable of sustained flights in excess of 60 min; however, males had higher rates of climb than females (3.31±0.17 and 2.63±0.19 cm s^–1, respectively). When a plant cue was presented during the parasitoid's phototactic flight, four relatively distinct responses were observed. Fifty-one percent of the individuals responded to the plant cue throughout their flight by flying toward or by landing on the cue. The majority of these parasitoids were females. Approximately 12% of the wasps exhibited an intermittent, positive response to the plant cue. Twenty percent exhibited a `migratory' response. These parasitoids, which were predominantly males, failed to respond to the plant cue until they had flown for a considerable period. Finally, 17% failed to respond to the target during their flight. Approximately 37% of the individuals that showed a positive response to the plant cue actually landed on it and the majority of these were female. The differential response to the plant cue by male and female parasitoids could be, in part, because females are driven to locate hosts in which to oviposit, and males are driven to find mates.     

Foraging behaviour of Brachionus calyciflorus (Pallas): variations in the swimming path according to presence or absence of algal food (Chlorella)

Both swimming and attachment behaviour of immature female Brachionus calyciflorus (Pallas) from a young orthoclone were analyzed under four experimental conditions: with and without algae (Chlorella), and with fed and starved individuals. Eighteen hours before the recording, two random groups were formed. To the first group, Chlorella was added. The second was placed in pure water. Both were filmed in red light to avoid a phototactic effect. Video-recorded swimming behaviour of B. calyciflorus was analyzed by automatic tracking. Each B. calyciflorus was filmed alternately in the presence or absence of Chlorella for five periods of 15 min. Under indentical light and temperature conditions the number of animals attached to the culture dish was measured with both starved and fed females. We observed a significant difference in swimming paths depending on the presence or absence of food, and the physiological state of the individuals. In the presence of Chlorella the linear speed was lower and the angular speed was greater. This increase in angular speed was more pronounced when females were subjected to a period of starvation before the experimentation. The attachment behaviour increased in the algal environment. This modification of swimming behaviour enables females to increase the time spent in a favorable food environment.     

Diel vertical migrations of invertebrate predators (Leptodora kindtii ,Thermocyclops taihokuensis, andMesocyclops sp .) in a shallow, eutrophic lake

We studied vertical distribution patterns of three invertebrate predators – Leptodora kindtii, Mesocyclopssp., and Thermocyclops taihokuensis– in a shallow eutrophic lake, Lake Suwa , Japan. From June to October in 2000 and 2001, we collected samples in the lake center in order to examine the vertical distribution patterns and the densities of the predators in the water column during the day (0900) and at night (2330). We also examined phototactic behavior of Leptodora in the laboratory. The three invertebrate predators showed clear migration patterns. Leptodora and Thermocyclops displayed a typical migration, avoiding the surface and maintaining a high abundance in deeper water during the day, and being distributed uniformly during the night. Mesocyclops, on the other hand, showed no clear vertical distribution pattern in the water column. However, Mesocyclops showed higher densities in the water column during the night than during the day. It suggests that they stayed just above the bottom during the day and migrated upward during the night. Leptodora also showed such a density difference between day and night. In the laboratory, Leptodora showed strong negative phototactic behavior. The observed density changes between day and night in Leptodora and Mesocyclops suggests the possible underestimation of their population density by usual sampling methods, and thus the impact of predation on populations of prey zooplankton species may also be underestimated in shallow water bodies.     

Local enhancement of arrivals of Ceratitis capitata females on fruit mimics

Mature female Mediterranean fruit flies, Ceratitis capitata (Wiedemann), isolated from conspecifics for 7 days prior to testing in laboratory cages, were attracted to yellow plastic fruit-mimicking hemispheres or spheres in significantly greater numbers when groups of conspecific females were present than absent on fruit mimics. Attraction of females to conspecifics occurred for females deprived of protein or continuously provided with protein since eclosion, for fruit mimics that were unbaited or baited with odor of food or fruit, and under single-choice as well as multiple-choice test conditions. Attraction proved just as great to fruit mimics having dead conspecifics as to mimics having live conspecifics, was evident also when fruit mimics were orange or white, and occurred also in response to presence of live Bactrocera dorsalis (Hendel)females on fruit mimics. We discuss our findings in relation to local enhancement, a form of social facilitation in insects and other animals.     

Paravortex karlingi sp. nov. from Cerastoderma edule L., in Britain

Paravortex karlingi sp. nov. collected from the intestine of the bivalve mollusc Cerastorderma edule from the Ythan Estuary, N. E. Scotland, and elsewhere, is distinguished from a closely related species, P. cardii, also occurring in this host, on the basis of differences in habitat occupied by the two species as well as behavioural and morphological differences. P. karlingi is smaller, has fewer embryos in the gravid adult and shows a different behaviour pattern when released from the host intestine. It is also negatively phototactic whereas P. cardii is initially positively phototactic, only later becoming negatively phototactic. The occurrence of both species in Britain is briefly described.Abbreviations bc brood capsules - c core of indigestible particles - ca common atrium - ch chamber - e eye - fa female atrium - fol fibre optic light - gc gut caecum - gp genital pore - j juvenile worm - m mouth - o ovary - oe oesophagus - p pharynx - pe penis - sg shell gland - sv seminal vesicle - t testis - u uterus - vg vitelline gland - w window     

Life history characteristics of Daphnia magna clones differing in phototactic behaviour

In an analysis of a life table experiment involving positively, intermediately and negatively phototactic Daphnia magna clones, life history traits such as the average duration of the adult instar, neonate and adult body size were found to be correlated with phototactic behaviour. The size of the eggs and neonates was positively correlated with adult body size, and with egg development time. Adult body size was positively correlated with the size of the second and subsequent clutches. I argue that the intrinsic positive correlation between offspring size and egg development time is a key factor structuring the differences in life history patterns observed between the positively and intermediately phototactic Daphnia genotypes, and that the two life history patterns are to be considered alternatives suited for different environmental conditions (e.g. habitats with and without strong predation pressure on adults).     

Sex recognition in zebrafish (<Emphasis Type="Italic">Danio rerio</Emphasis>)

We investigated sex recognition in female zebrafish (Danio rerio) to better understand the underlying sensory mechanisms and identify male secondary sexual traits. Females were simultaneously presented with two fish, a male and a female, in a flow-chamber apparatus, and females’ relative attraction towards males was observed under different conditions. With domesticated fish, females were more attracted to males when presented with both visual and chemosensory cues from stimulus fish. They still discriminated the sexes when only visual cues were provided, but not when white ambient light was changed to yellow, indicating that colour plays a role. Sex discrimination under yellow light was improved when chemosensory cues were also provided. Surprisingly, females’ attraction to males was not more pronounced in the morning when mating occurs. Domesticated females discriminated the sexes when presented with wild-derived, as well as domesticated fish, whereas wild-derived females did not show any biases for domesticated or wild-derived males. Behavioural observations indicated that the wild-derived females were distressed, which explains their lack of attraction to males. In summary, domesticated female zebrafish discriminated the sexes using both visual (body colour) and olfactory cues; however, wild-derived zebrafish were too distressed for behavioural experiments under these laboratory conditions.     

Evolution of predator avoidance in a Daphnia population: evidence from the egg bank

Dormant propagule pools (egg banks) of zooplankton populations that accumulate in sediments provide biological archives of past conditions and enable the investigation of evolutionary changes in populations over relatively long periods of time (many decades). This study examined the egg bank of a Daphnia pulicaria population in a lake that has been stocked annually with rainbow trout (a zooplanktivore) since 1961. Resting eggs from sediments from the 1920s to 2001 (determined by ²¹⁰Pb dating) were hatched and established as isofemale clonal lines. The phototactic behavior (a proxy for vertical migration behavior) of clones was assessed in the presence and absence of fish kairomones to evaluate the hypothesis that the heavy and consistent level of predation imposed by the stocked trout would have selected for clones that are more negatively phototactic. In addition, exposure to kairomones was expected to induce stronger negative phototaxis for all clones relative to control conditions. The behavior of clones derived from resting eggs from the trout era (after 1961) was significantly more negatively phototactic than it was for pre-trout era clones. Kairomone exposure induced a more negative phototactic response in clones from both eras, but the response was much greater for the pre-trout era clones. These results suggest that the consistently high level of predation by trout over several decades has selected for a conservative (less plastic) vertical migration strategy in which Daphnia maintain a deep daytime distribution with or without the presence of chemical cues from fish predators.     

Phototactic behaviour of Eucryptorrhynchus scrobiculatus and E. brandti (Coleoptera: Curculionidae) adults*

Eucryptorrhynchus scrobiculatus and E. brandti (Coleoptera: Curculionidae) are destructive weevils on Ailanthus altissima in China. This study examined phototactic behaviour of E. scrobiculatus and E. brandti in response to eight light-emitting diodes (LEDs) in the laboratory and field. Effects of gender, starvation, and light and dark experience on the phototactic behaviour of the insects were evaluated. The results demonstrated that, the two species of weevil were phototactic insects and most sensitive to violet light (400–405?nm), followed by blue–violet light (420–430?nm). They were less sensitive to red (655–660?nm), white (6000–6500?k), blue–green (470–480?nm), yellow (590–595?nm), blue (450–455?nm), and green (515–530?nm) light. In the light intensity range of 200–1000?lux, the light intensity had no significant effect on the phototatic behaviour of E. scrobiculatus and E. brandti. Phototactic behaviour of the insects was affected by gender. The phototaxis indices of the two species of weevil increased with starvation, reaching a plateau after 2 or 3?d of starvation. The phototaxis indices of E. scrobiculatus and E. brandti were significantly affected by various wavelengths of light following exposure to the light for 3?h or in different dark experience time. In the mark-release-recapture test, the number of E. scrobiculatus and E. brandti adults trapped by violet (400–405?nm) light traps is the largest. The information provided here provides a basis for survey and control of E. scrobiculatus and E. brandti.     

Multiple matings increase the fecundity of the yellow swallowtail butterfly,Papilio xuthus L., in summer generations

Females of the yellow swallowtail butterfly, Papilio xuthus,were reared in the laboratory. They were divided into four groups held under different mating conditions: nonmating (virgin) and mated once, twice, and three times. The number of eggs in the ovaries was counted by dissection. Virgin females produced increasing numbers of mature eggs, up to about 30, in the week following emergence. When the female had mated once, the number of mature eggs was significantly higher than that of virgin females by the second day after emergence. However, the double- and triplemated females did not increase the number of eggs in each state further than the singlemated females. The double-mated females deposited significantly more eggs than the singlemated females in the laboratory. The triplemated females also deposited more eggs on the day after the third mating than the doublemated females. Thus, multiple matings increased the number of eggs deposited. The change in the hatchability and the morphology of the spermatophore in the bursa copulatrix suggested that the sperm from the last mating had precedence.     

Relevance of visual and olfactory cues for host location in the mustard leaf beetle Phaedon cochleariae

The relevance of visual and olfactory cues for host‐plant location is investigated in males and females of the oligophagous mustard leaf beetle Phaedon cochleariae Fabricius (Coleoptera: Chrysomelidae). Different objects are offered in a walking arena and the behaviour of beetles is observed. Beetles orient toward vertically or horizontally striped black and white pattern independent of stripe orientation. The results suggest that contrast facilitates orientation in the field, whereas the pattern itself may be less important for host location in dense vegetation. The response to green and yellow objects is tested to investigate discrimination abilities between young (green) and mature (yellow) leaves. Beetles prefer green over yellow independent of material (cardboard or leaves of Nasturtium officinale R. Br., Brassicaceae). Preference behaviour tested in a dual‐choice contact assay coincides with visual preferences, where adults prefer young, more nutritious leaves for feeding and oviposition. Furthermore, females discriminate between visual cues of green leaves and green cardboard, whereas males do not, indicating that females are more sensitive in colour discrimination. Differences in colour wavelength influence the choice of beetle behaviour more strongly than differences in intensity. Both sexes of P. cochleariae prefer volatiles of the host plant N. officinale, whereas only females respond to the main volatile compound 2‐phenylethyl isothiocyanate. Given a choice between visual and olfactory cues, males orientate towards the colour cues, whereas females do not show any preferences. In males, visual cues may thus override olfactory cues, whereas, in females, both are equally important, which may reflect different ecological requirements and/or physiological abilities.     

光谱和光强度对棕榈蓟马雌成虫行为反应的影响

室内研究了光谱、光强度对棕榈蓟马雌成虫的趋、避光行为的影响。结果显示:在340—605 nm波谱内棕榈蓟马雌成虫对14个单色光刺激的趋光行为反应为多峰型。其中蓝光483 nm处峰最高,趋光反应率达34.96%,其次为绿光498—524 nm、562—582 nm、紫外光340 nm处;其避光行为反应共有3个峰,其中紫外光380 nm处最高,避光率18.08%,另外2个峰分别在橙光605 nm、紫光420 nm处。在趋光率较高的单色光(340、483、524、582 nm)和避光率较高的单色光(380、605 nm)以及白光刺激下,棕榈蓟马雌成虫的趋光率随光强增强的增强而提高,而避光率则随着光强的增强而降低;光强最弱时仍均有一定趋光率,最强时均未出现高端平台。因此:棕榈蓟马雌成虫对不同单色光具有明显的选择性,光谱和光强度对其趋光行为和避光行为有较大影响,光强度的影响作用与波长因素有关。     

An analysis of the phototactic behaviour of Daphnia magna clones and their sexual descendants

Daphnia magna clones, isolated from different natural populations, were inbred and crossbred. Some aspects of the heredity of the phototactic behaviour, quantified by an index, were analysed. Heritability in the broad sense was determined for several assemblages of clones through calculation of the clonal repeatability. Heritability sensu strictu was determined by offspring-parent regressions. Frequency distributions of a phototactic index of several inbred and crossbred families are given.The results indicate a significant contribution of additive genetic variance to the total phenotypic variance. Comparison of heritability estimates suggests an important genetic interaction component.Some implications of these results on the variability of vertical migration patterns in natural populations are discussed.