The response of an insect parasitoid,Ormia ochracea (Tachinidae), to the uncertainty of larval success during infestation

Ormia ochracea is a parasitoid fly which lays its larvae on its hosts, the field crickets Gryllus integer and Gryllus rubens, in two distinct modes: (1) directly on the host and (2) around the host. In the field, 12.7% of male crickets were parasitized and 3.2% were super-parasitized. Despite the disadvantages of parasitizing infested hosts, there was no evidence that O. ochracea avoided superparasitism. This and other experiments suggest that the host assessment ability of O. ochracea is less than that reported for many hymenopteran parasitoids. by manipulating the number of larvae in each cricket, we determined that four to five larvae per host resulted in the largest number of adult flies. However, as larval number per host increased from one to six, pupal size, and hence adult size, declined. In the field, hosts were found with a mean of 1.7 ± 1.0 (SD) larvae per cricket, suggesting that there may be some selection pressure against larger clutch sizes. Nevertheless clutch sizes larger than the host can support were sometimes found in the field. During the first mode of larviposition, gravid flies deposited no more than three larvae directly onto the host. Larvae deposited directly on the host had a high probability of infesting it. During the second mode of larviposition, gravid flies laid a larger number of larvae around the host (6.1 ± 5.2). Larvae that were laid around the host were less likely to infest a cricket than were larvae that were deposited directly onto it. O. ochracea is unique in that its two different modes of larviposition have different probabilities of larval success. Even though the success rate for larvae laid during the second mode of larviposition was low, the possibility of parasitizing more hosts appears to have selected for flies laying more larvae (e.g. increasing clutch size) than is optimal if all the larvae successfully entered a single host.     

Impacts of environmental heterogeneity on alternative mating tactics in the threadtail damselfly

Environmental heterogeneity, including variation in the physical environment, may be key to understanding the evolution and maintenance of alternative mating tactics, but its influence is rarely examined. Males of the threadtail damselfly Protoneura amatoria reversibly use two alternative mating tactics (perching vs. hovering) and have previously been found to modulate their use of these tactics in response to variation in both light conditions and the density of ovipositing females. Here, I show that mating success payoffs of the two tactics are differentially influenced by these factors. The payoff of the perching tactic was greater than that of the hovering tactic under low light conditions and at low densities of ovipositing females. The payoff of the hovering tactic was greater under high light conditions and higher densities of ovipositing females. The differential success of the two mating tactics in response to light conditions is discussed in light of flight dynamics, vision, and predation.     

Test of the adaptiveness of sex ratio manipulation in a parasitoid wasp

In behavioral ecology it is generally assumed that behavior is adaptive. This assumption is tested here for sex ratio manipulation in response to host size in the parasitoid wasp Spalangia cameroni. Females produce a greater proportion of daughters on larger hosts. If this behavior is adaptive, it is not through a positive effect of host size on the fitness of daughters, as theory suggests and as found for other species. Females that developed on larger hosts were not more successful at drilling into hosts, were not more successful at interspecific competition for hosts, and did not have greater dispersal ability as measured by wing loading (weight/area of wing and thorax). The possibility that S. cameroni's sex ratio manipulation may be adaptive through a negative effect of host size on the fitness of sons cannot be ruled out. Relative to males from larger hosts, males from smaller hosts had lower wing loading and thus potentially greater dispersal ability. The actual effect of wing loading on fitness remains to be tested.     

Effect of water temperature on the courtship behavior of the Alpine newt Triturus alpestris

Temperature is expected to have an effect on the behavioral patterns of all organisms, especially ectotherms. However, although several studies focused on the effect of temperature on acoustic displays in both insects and anurans, almost nothing is known about how environmental temperature may affect ectotherm visual courtship displays and sexual performance. The purpose of this study was to determine the effect of environmental temperature on the sexual behavior of Alpine newts (Triturus alpestris). We subjected T. alpestris to two different temperatures in controlled laboratory conditions. Temperature had a major effect on both male and female behaviors: at low temperature, the frequencies of several displays, including tail-raising during sperm deposition, are lowered. This variation is caused indirectly by temperature because it is due to female responsiveness, which is temperature-dependent. However, the fanning movement of the males tail during its main courtship display is independent of female behavior: at lower temperatures, the tail beats at a lower rate, but for a longer time. The similar reproductive success (i.e. sperm transfer) at the two temperature ranges indicates that breeding in cold water is not costly but instead allows males and females to mate early in the season. This is particularly adaptive because, in many habitats, the reproductive period is shortened by drying or freezing conditions, which may impair survival of branchiate offspring. This study also demonstrates the necessity of considering environmental parameters when modeling optimality and characteristics of ectotherm behaviors.Communicated by W. Cooper     

A trade-off between prey- and predator-induced polyphenisms in larvae of the salamander Hynobius retardatus

Organisms in natural habitats participate in complex ecological interactions that include competition, predation, and foraging. Under natural aquatic environmental conditions, amphibian larvae can simultaneously receive multiple signals from conspecifics, predators, and prey, implying that predator-induced morphological defenses can occur in prey and that prey-induced offensive morphological traits may develop in predators. Although multiple adaptive plasticity, such as inducible defenses and inducible offensive traits, can be expected to have not only ecological but also evolutionary implications, few empirical studies report on species having such plasticity. The broad-headed larval morph of Hynobius retardatus, which is induced by crowding with heterospecific anuran (Rana pirica) larvae, is a representative example of prey-induced polyphenism. The morph is one of two distinct morphs that have been identified in this species; the other is the typical morph. In this paper, we report that typical larval morphs of Hynobius can respond rapidly to a predatory environment and show conspicuous predator-induced plasticity of larval tail depth, but that broad-headed morphs cannot respond similarly to a predation threat. Our findings support the hypothesis that induction or maintenance of adaptive plasticity (e.g., predator-induced polyphenism) trades off against other adaptive plastic responses (e.g., prey-induced polyphenism). For a species to retain both an ability to forage for larger prey and an ability to more effectively resist predation makes sense in light of the range of environments that many salamander larvae experience in nature. Our results suggest that the salamander larvae clearly discriminate between cues from prey and those from predators and accurately respond to each cue; that is, they adjust their phenotype to the current environment.     

The effect of mirrors on African clawed frog (Xenopus laevis) larval growth, development, and behavior

We examined the behavioral and developmental responses of Xenopus laevis larvae to their mirror images in three experiments. The mirrors allowed us to visually simulate increased density, without the tadpoles’ behavior being confounded by chemical cues from additional tadpoles. In the first experiment, we demonstrated that Xenopus tadpoles have a right eye preference for mirrors, contrary to the left eye preference of all other anuran species studied to date. This lateralized eye use disappeared, however, as tadpoles approached metamorphosis. Next, we examined how mirrored aquaria walls affected tadpole growth and development. We found that tadpoles raised in aquaria with partially mirrored walls showed depressed growth compared to tadpoles raised without mirrors, despite the fact that Xenopus larvae normally thrive when raised in visual contact with conspecifics. The tadpoles raised with mirrors had, though not significantly, proportionally larger bodies relative to their tail length (d = 0.51). This suggests that a phenotypically plastic response in body proportions was induced in these tadpoles solely by the sight of other tadpoles. The third experiment established that X. laevis tadpoles are more active in front of a mirror; i.e., they turn more often and spend more time in front of mirrored surfaces. We consider this increased activity to be an aberrant behavior of the tadpoles, which were attempting to school with their own images. We suggest that this extra activity reduced the amount of energy available for growth, accounting for the depressed growth seen in our second experiment.     

Patch departure rules in Bumblebees: evidence of a decremental motivational mechanism

The patch living rules of a pollinator, the bumblebee Bombus terrestris L., are studied here in the framework of motivational models widely used for parasitoids: The rewarding events found during the foraging process are supposed to increase or decrease suddenly the tendency of the insect to stay in the current patch and therefore to adjust the patch residence time to the patch profitability. The foraging behaviour of these pollinators was observed in two environment types to determine their patch-leaving decisions. The rich environment was composed of male-fertile flowers, offering pollen and nectar, and the poor one of male-sterile flowers, offering little nectar and no pollen. The experimental design consisted of a patch system in which inflorescences were evenly arranged in two rows (1 m distance). Residence times of foragers inside inflorescences and rows were analysed by a Cox proportional hazards model, taking into account recent and past experience acquired during the foraging bout. Most of the results showed a decremental motivational mechanism, that is, a reduction in the residence time on the inflorescence or in the row related to exploitation of flowers within inflorescences and inflorescences within rows These results indicate that bumblebees tend to leave the patch using departure rules similar to those found in parasitoids. The results also provide information on the memory, learning and evaluating capabilities of bumblebees especially when rich and poor environments were compared. The patch-leaving mechanism suggested by this study is consistent with the central place foraging theory.     

Environmental and genetic influences on mating strategies along a replicated food availability gradient in guppies (Poecilia reticulata)

Food availability is expected to influence the relative cost of different mating tactics, but little attention has been paid to this potential source of adaptive geographic variation in behavior. Associations between the frequency of different mating tactics and resource availability could arise because tactic use responds directly to food intake (phenotypic plasticity), because populations exposed to different average levels of food availability have diverged genetically in tactic use, or both. Different populations of guppies (Poecilia reticulata) in Trinidad experience different average levels of food availability. We combined field observations with laboratory “common garden” and diet experiments to examine how this environmental gradient has influenced the evolution of male mating tactics. Three independent components of variation in male behavior were found in the field: courtship versus foraging, dominance interactions, and interference competition versus searching for mates. Compared with low-food-availability sites, males at high-food-availability sites devoted more effort to interference competition. This difference disappeared in the common garden experiment, which suggests that it was caused by phenotypic plasticity and not genetic divergence. In the diet experiment, interference competition was more frequent and intense among males raised on the greater of two food levels, but this was only true for fish descended from sites with low food availability. Thus, the association between interference competition and food availability in the field can be attributed to a genetically variable norm of reaction. Genetically variable norms of reaction with respect to food intake were found for the other two behavioral components as well and are discussed in relation to the patterns observed in the field. Our results indicate that food availability gradients are an important, albeit complex, source of geographic variation in male mating strategies.     

A model for the evolution of communal foraging in hierarchically structured populations

A kin selection model is described for populations in which groups of interacting individuals (trait groups, sensu Wilson 1975) are spatially situated within larger aggregations. The model predicts the optimal foraging strategy when resources are shared with other trait group members and there is an individual risk in foraging. The ecological mechanism of variation in group fitness, differential resource accumulation, is explicitly incorporated into the model. The optimal foraging rate obtained from this model depends on the product of a benefit-to-cost ratio and a relatedness parameter. The appropriate definition of relatedness for the evolution of communal foraging is determined by the details of the ecological interaction between consumers and resources. When competition is purely intra-specific, the genetic correlation among interactants relative to other members of the local aggregation defines the relatedness parameter applicable to selection on foraging propensity. When competition is primarily inter-specific, the genetic correlation among trait group members relative to the entire population defines relatedness.     

Adaptive prey behavior and the dynamics of intraguild predation systems

Intraguild predation constitutes a widespread interaction occurring across different taxa, trophic positions and ecosystems, and its endogenous dynamical properties have been shown to affect the abundance and persistence of the involved populations as well as those connected with them within food webs. Although optimal foraging decisions displayed by predators are known to exert a stabilizing influence on the dynamics of intraguild predation systems, few is known about the corresponding influence of adaptive prey decisions in spite of its commonness in nature. In this study, we analyze the effect that adaptive antipredator behavior exerts on the stability and persistence of the populations involved in intraguild predation systems. Our results indicate that adaptive prey behavior in the form of inducible defenses act as a stabilizing mechanism and show that, in the same direction that adaptive foraging, enhances the parameter space in which species can coexist through promoting persistence of the IG-prey. At high levels of enrichment, the intraguild predation system exhibits unstable dynamics and zones of multiples attractors. In addition, we show that the equilibrium density of the IG-predator could be increased at intermediate values of defense effectiveness. Finally we conclude that adaptive prey behavior is an important mechanism leading to species coexistence in intraguild predation systems and consequently enhancing stability of food webs.     

Differentiated phenotypic plasticity in larvae of the cannibalistic salamander Hynobius retardatus

Alternative phenotypes in natural populations can arise from either genetic polymorphism or an environmentally induced phenotype, that is, polyphenism. Evolutionary models of polyphenism developed by theoretical studies predict that polyphenism is favored when there are environment-dependent fitness trade-offs between alternatives and that the threshold frequency for a facultative switch between alternative phenotypes is adjusted in accordance with different selection regimes. The broad-headed (alternative) larval morph of Hynobius retardatus, which is induced by crowding with conspecifics or heterospecific anuran (Rana pirica) larvae, is a representative example of cannibalistic polyphenism. Morph induction by such proximate factors must reflect evolutionary (conditional frequency-dependent) processes. To clarify the role of frequency-dependent processes in polyphenism, I investigated the occurrence rate of the broad-headed morph under experimental crowding conditions (low conspecific, high conspecific, and high heterospecific densities) using larvae from eight natural populations with different larval densities of conspecifics and heterospecifics, and found interpopulational differences in the expression of the morph. Thus, there is a larval density-dependent equilibrium frequency of the morph in each pond, suggesting that the local switch point for morph induction was modified by selection to produce evolved differences between ponds. The evolution of such interpond differences has three necessary conditions: (1) There are pond-dependent fitness trade-offs between alternatives, (2) The maintenance of the morph is costly, and (3) The presence of conspecific or, especially, heterospecific larvae provides a reliable cue to the receiver.     

Effect of decision rules in choice experiments on hunting and bushmeat trade

Providing insight on decisions to hunt and trade bushmeat can facilitate improved management interventions that typically include enforcement, alternative employment, and donation of livestock. Conservation interventions to regulate bushmeat hunting and trade have hitherto been based on assumptions of utility- (i.e., personal benefits) maximizing behavior, which influences the types of incentives designed. However, if individuals instead strive to minimize regret, interventions may be misguided. We tested support for 3 hypotheses regarding decision rules through a choice experiment in Tanzania. We estimated models based on the assumptions of random utility maximization (RUM) and pure random regret maximization (P-RRM) and combinations thereof. One of these models had an attribute-specific decision rule and another had a class-specific decision rule. The RUM model outperformed the P-RRM model, but the attribute-specific model performed better. Allowing respondents with different decision rules and preference heterogeneity within each decision rule in a class-specific model performed best, revealing that 55% of the sample used a P-RRM decision rule. Individuals using a P-RRM decision rule responded less to enforcement, salary, and livestock donation than did individuals using the RUM decision rule. Hence, 3 common strategies, enforcement, alternative income-generating activities, and providing livestock as a substitute protein, are likely less effective in changing the behavior of more than half of respondents. Only salary elicited a large (i.e. elastic) response, and only for one RUM class. Policies to regulate the bushmeat trade based solely on the assumption of individuals maximizing utility, may fail for a significant proportion of the sample. Despite the superior performance of models that allow both RUM and P-RRM decision rules there are drawbacks that must be considered before use in the Global South, where very little is known about the social–psychology of decision making.     

The dynamics of social learning in an insect model,the bumblebee (<Emphasis Type="Italic">Bombus terrestris</Emphasis>)

Bumblebees (Bombus terrestris) are attracted to those particular inflorescences where other bees are already foraging, a process known as local enhancement. Here, we use a quantitative analysis of learning in a foraging task to illustrate that this attraction can lead bees to learn more quickly which flower species are rewarding if they forage in the company of experienced conspecifics. This effect can also be elicited by model bees, rather than live demonstrators. We also show that local enhancement in bumblebees most likely reflects a general attraction to conspecifics that is not limited to a foraging context. Based on the widespread occurrence of both local enhancement and associative learning in the invertebrates, we suggest that social influences on learning in this group may be more common than the current literature would suggest and that invertebrates may provide a useful model for understanding how learning processes based on social information evolve.     

Patchiness and bistability in the comprehensive cyanobacterial model (CCM)

We analyzed spatiotemporal dynamics of cyanobacteria using a four-component mathematical model with nutrients, unicellular algae, colonial algae and herbivorous zooplankton. One of the astonishing abilities of cyanobacteria is the morphological change from the unicellular type to the colonial or filamentous type in response to grazing activities of zooplankton, the phenomenon known as phenotypic plasticity. Our model, referred to as comprehensive cyanobacterial model (CCM), includes the effects of phenotypic plasticity. Depending on parameter values, CCM exhibits not only bistability but also limit cycle oscillations without showing the paradox of enrichment, which has been a controversy among mathematical ecologists. Further, CCM is modified to reaction–advection–diffusion equations, the simulation results of which indicate that the ratio of the lateral diffusivity to the turbulent velocity significantly affects the appearance of patchiness patterns.     

基于食物链关系评价转Bt基因棉对寄生性天敌的非靶效应

按照“棉花 -棉大卷叶螟 (Syleptaderogata)幼虫 -螟蛉绒茧蜂 (Apantelesruficrus)”三级营养食物链关系 ,对田间转Bt基因棉苏抗 310对非靶物种螟蛉绒茧蜂种群的影响进行了调查。发现转Bt基因棉苏抗 310棉田间螟蛉绒茧蜂种群生长受到了不良影响 ,种群数量显著下降 ,棉大卷叶螟幼虫发生高峰期高龄幼虫被寄生率降低。     

Behavioral diversification in a young species flock of pupfish (<Emphasis Type="Italic">Cyprionodon</Emphasis> spp.): shoaling and aggressive behavior

Increasing empirical and theoretical evidence supports the idea that sympatric speciation is operating, for example, in species flocks comprising several closely related fish species within one lake. Divergent natural selection (promoting spatial and food niche partitioning) and sexual selection (assortative mating) have been identified as key selection factors in intralacustric adaptive radiations. However, the evolution of social behaviors accompanying such adaptive radiations is less well understood. Using a phylogenetically young species flock of pupfish (Cyprinodon spp.) as a model, we examined differences among six sympatric species and compared their shoaling, aggressive, and territorial behaviors with that of a sister species (C. artifrons). Despite an estimated age of the species flock of less than 8,000 years, pronounced behavioral differentiation was found. C. simus, the smallest species in the flock, shoaled more than the other species and was less aggressive and less territorial than C. beltrani. F₁-hybrids between C. simus males and C. beltrani females showed an intermediate expression of shoaling and aggressive behavior. Niche partitioning among the members of this species flock appears to be accompanied by rapid divergent evolution of social behaviors. We discuss the potential role of phenotypic plasticity and within-species variation of social behaviors for such rapid behavioral diversifications in sympatric speciation processes.     

Meta-analysis of foraging and predation risk trade-offs in terrestrial systems

Although there is ample evidence for the generality of foraging and predation trade-offs in aquatic systems, its application to terrestrial systems is less comprehensive. In this review, meta-analysis was used to analyze experiments on giving-up-densities in terrestrial systems to evaluate the overall magnitude of predation risk on foraging behavior and experimental conditions mediating its effect. Results indicate a large and significant decrease in foraging effort as a consequence of increased predation risk. Whether experiments were conducted under natural or artificial conditions produced no change in the overall effect predation had on foraging. Odor and live predators as a correlate of predation risk had weaker and nonsignificant effects compared to habitat characteristics. The meta-analysis suggests that the effect of predation risk on foraging behavior in terrestrial systems is strongly dependent on the type of predation risk being utilized.     

Social,environmental and genetic factors in the ontogeny of phenotypic differentiation in a lizard with alternative male reproductive strategies

Summary Adult male tree lizards, Urosaurus ornatus, practise alternative (territorial or sneaker/satellite) reproductive strategies that are correlated with differences in throat color and body size. In this study we raised tree lizards from hatching in the laboratory to examine the question of whether the phenotypic expression of secondary sex coloration and body size can be facultatively influenced by social or abiotic environmental factors. We compared males reared in the laboratory under different social and environmental conditions to males in the field and found no effect of different conditions on phenotypic differentiation (Figs. 2–4). Thus, phenotypic differences between morphs probably result largely from nonfacultative expression of different genotypes. This suggests that alternative male morphs practise a mixed evolutionary stable strategy (ESS) rather than one morph making the best of a bad situation. However, in the context of ESS theory it is difficult to explain our further result that the nonterritorial morph in this species grows faster and reaches a larger adult body size than the territorial morph (Fig. 5).     

Effects of hydrolysable tannins on a herbivorous insect: fate of individual tannins in insect digestive tract

Summary. We investigated the effects of four chemically characterised galloylglucoses (GGs, a subgroup of hydrolysable tannins) and their hydrolysis product, gallic acid (GA), on consumption and performance of larvae of the autumnal moth Epirrita autumnata. Larvae were fed with birch (Betula pubescens) leaves that had been painted individually with each of the compounds at two levels, 5 and 20 mg/g. In addition, we investigated the fates of the leaf-painted GGs and GA in the E. autumnata digestive tract by comparing phenolics in leaves consumed and in faeces. In general, GGs reduced leaf consumption by E. autumnata during the second and fourth instars, although there was high compound- and instar-specific variation. However, GGs did not affect the leaf consumption rates by the most voracious fifth instar larvae. This resulted in approximately the same loss of total biomass by the experimental tree, regardless of the nature and level of GGs enriched to its foliage. The characteristic fate of hydrolysable tannins, i.e. hydrolysis, was evidenced in the larval digestive tract for three of the four leaf-painted GGs. In addition to hydrolysis, the almost total absence of GGs in larval faeces was presumably related to the oxidation of GGs. The dose-dependent excretion percentage of ingested GA showed that it's faecal content should not be used, although it commonly is, to calculate the level of GG hydrolysis. Moreover, by comparing the non-uniform appearance of faecal tetragalloylglucoses, whether ingested as such or hydrolysed from pentagalloylglucose, we concluded that a major part of oxidation of GGs occurs before their hydrolysis in the digestive tract of E. autumnata. Criticism against the common use of tannic acid, a heterogeneous mixture of GA and GGs, in ecological studies is presented. Received 15 May 2002; accepted 16 July 2002     

Effects of elevated yolk testosterone levels on survival, growth and immunity of male and female yellow-legged gull chicks

Androgen hormones of maternal origin contained in the eggs of avian species are considered to have positive effects on offspring characteristics and performance. However, negative consequences have also been reported, suggesting that mothers may experience a trade-off between beneficial and detrimental effects of egg androgens to offspring fitness. We studied the effects of elevated yolk testosterone (T) concentration on survival, development and phenotype of male and female yellow-legged gull (Larus michahellis) chicks by injecting egg yolks with physiological doses of the hormone. Elevated yolk T resulted in a male-biased post-hatching sex ratio, T-treated clutches producing a greater proportion of males compared to control ones at day 4 post-hatching, likely resulting from a reduction of female embryonic survival, whereas no effect of hormone treatment on hatching success or short-term chick survival was observed. In addition, T depressed post-hatching body mass in both sexes but had no effects on the intensity of the cell-mediated immune response or skeletal growth. No sex differences in egg characteristics or chick phenotype were detected. Time to hatching was not affected by T, but females originating from first laid eggs hatched earlier than males of the same laying order, independently of hormone treatment. However, the implications of sex differences in hatching times are unclear in the study species. Taken together, our results suggest that female yellow-legged gulls may be constrained in transferring androgens to their eggs by negative consequences on the viability of female offspring and growth of chicks of the two sexes.