Background level of risk and the survival of predator-naive prey: can neophobia compensate for predator naivety in juvenile coral reef fishes?

Neophobia—the generalized fear response to novel stimuli—provides the first potential strategy that predator-naive prey may use to survive initial predator encounters. This phenotype appears to be highly plastic and present in individuals experiencing high-risk environments, but rarer in those experiencing low-risk environments. Despite the appeal of this strategy as a ‘solution’ for prey naivety, we lack evidence that this strategy provides any fitness benefit to prey. Here, we compare the relative effect of environmental risk (high versus low) and predator-recognition training (predator-naive versus predator-experienced individuals) on the survival of juvenile fish in the wild. We found that juveniles raised in high-risk conditions survived better than those raised in low-risk conditions, providing the first empirical evidence that environmental risk, in the absence of any predator-specific information, affects the way naive prey survive in a novel environment. Both risk level and experience affected survival; however, the two factors did not interact, indicating that the information provided by both factors did not interfere or enhance each other. From a mechanistic viewpoint, this indicates that the combination of the two factors may increase the intensity, and hence efficacy, of prey evasion strategies, or that both factors provide qualitatively separate benefits that would result in an additive survival success.     

Dazzle coloration and prey movement

Many traits in animals reduce the rate of attack from visually hunting predators, including camouflage, warning signals and mimicry. In addition, some animal markings may reduce the likelihood that an attack ends in successful capture. These might include dazzle markings, high-contrast patterns that make the estimation of speed and trajectory difficult. However, until now, no study has experimentally tested whether some markings may achieve such an effect. We developed a computer 'game' where human 'predators' have to capture computer-generated prey moving across a background. In two experiments, we find that although uniform camouflaged targets were among the hardest to capture, so were a range of high-contrast conspicuous patterns, such as bands and zigzags. Prey were also more difficult to capture against more heterogeneous than uniform backgrounds, and at faster speeds of movement. As such, we find the first experimental evidence that conspicuous patterns, similar to those found in a wide range of real animals, make the capture of moving prey more challenging. Various anti-predator markings may work prey during motion, and some animals may combine such dazzle patterns with other functions, such as camouflage, thermoregulation, sexual and warning signals.     

Spots and stripes: ecology and colour pattern evolution in butterflyfishes

The incredible diversity of colour patterns in coral reef fishes has intrigued biologists for centuries. Yet, despite the many proposed explanations for this diversity in coloration, definitive tests of the role of ecological factors in shaping the evolution of particular colour pattern traits are absent. Patterns such as spots and eyespots (spots surrounded by concentric rings of contrasting colour) have often been assumed to function for predator defence by mimicking predators'' enemies'' eyes, deflecting attacks or intimidating predators, but the evolutionary processes underlying these functions have never been addressed. Striped body patterns have been suggested to serve for both social communication and predator defence, but the impact of ecological constraints remains unclear. We conducted the first comparative analysis of colour pattern diversity in butterflyfishes (Family: Chaetodontidae), fishes with conspicuous spots, eyespots and wide variation in coloration. Using a dated molecular phylogeny of 95 species (approx. 75% of the family), we tested whether spots and eyespots have evolved characteristics that are consistent with their proposed defensive function and whether the presence of spots and body stripes is linked with species'' body length, dietary complexity, habitat diversity or social behaviour. Contrary to our expectations, spots and eyespots appeared relatively recently in butterflyfish evolution and are highly evolutionarily labile, suggesting that they are unlikely to have played an important part in the evolutionary history of the group. Striped body patterns showed correlated evolution with a number of ecological factors including habitat type, sociality and dietary complexity. Our findings question the prevailing view that eyespots are an evolutionary response to predation pressure, providing a valuable counter example to the role of these markings as revealed in other taxa.     

Density-dependent and frequency-dependent selection by bumblebees Bombus terrestris (L.) (Hymenoptera: Apidae)

The behaviour of bumblebee workers foraging on arrays of artificial flowers of two colour morphs was observed. Experiments were conducted on arrays of varying morph frequencies and at three different total flower densities. Bumblebees consistently showed a preference for the commonest colour morph, and this behaviour was not significantly affected by changing density. In contrast, frequency-independent preferences changed significantly with density. At low densities, there was a strong bias towards the more conspicuous colour, whereas at higher densities there was no overall colour bias. Flight distances between flowers decreased significantly at high density. Bumblebees also visited flowers of similar colours sequentially, but this behaviour was not density-dependent. It is suggested that as densities increase, there is an increased probability that bumblebees detect yellow flowers, which were probably less conspicuous compared with blue flowers, and that this might be caused by changes in flight speed with flight distance. Where there is a positive relationship between pollinator visitation and the relative fitness of a floral morph, the observed behaviour would induce positive frequency-dependent selection on a plant population with two corolla colour morphs on which the bumblebees were foraging, which would result in stabilizing selection for a single corolla colour, irrespective of density. There was no indication that rare colour morphs would be preferred at high density. The probability of different corolla colour morphs going to fixation would, however, be affected by density.     

The Zonal Practice of Second-year Students in the Biological Faculty of Moscow University

Gametophytes are easily cultured, even with limited facilities. They can he used to illustrate not only the alternation of generations in plant life cycles but also aspects of reproduction and development not easily studied with other plant material. The approach can be adjusted to the needs of biology and botany classes over a wide range of experience from secondary school to university. Information is presented on: the choice of species; the collection, storage and sowing of spores; the composition of the medium; light and temperature requirements; sampling and staining techniques; and methods for determining quantitative parameters of cell and gametophyte development. The basic fern life cycle, and modifications such as apogamy, are outlined. Attention is drawn to the main features of the reproductive biology of ferns and appropriate investigations are suggested. The advantages of gametophytes as experimental material for studying cellular aspects of development in intact green plants are explained. They include striking but consistent responses to a range of cultural conditions and environmental treatments. Suggestions are given for investigations to stimulate further discussion of a wide range of developmental events.     

Factors regulating predation by first-instar spined assassin bugs [Sinea diadema (Fabricius)] (Hemiptera: Reduviidae)

Selected nutritional and developmental factors regulating the predatory behavior of first-instar spined assassin bugsSinea diadema (Fabricius) (Hemiptera: Reduviidae) were investigated. The longevity of unfed nymphs provided with free water, bean pod sections, or glucose solutions was not significantly greater than that of unfed nymphs which were not provided with a source of water. First-instarS. diadema that were provided with larvae ofEphestia kuehniella Zeller (Lepidoptera: Pyralidae) as prey began feeding 1.9 (±0.9) days after hatching. In contrast, first-instarS. diadema that were provided with conspecifics of the same age did not begin feeding until they were 3.9 (±0.9) days old. These results suggest that the potential nutritional benefits to be gained from feeding equal or outweigh the risk of attacking prey capable of effective self-defense only when nymphs have not fed for 4 days after hatching. Providing the nymphs with water or glucose solutions significantly delayed the onset of conspecific predation. Additional data are presented which demonstrate that first-instarS. diadema are not restrained from preying on siblings by kin recognition.     

Defensive ant,aphid and caterpillar mimicry in plants?

Here we describe three apparently novel types of visual insect mimicry in plants. In the first type, plants of Xanthium trumarium L. have dark spots and flecks that resemble ants (Formicidae) in size and shape in the epidermis of stems, branches and petioles, and plants of Arisarum vulgare Targ.-Tozz. have them on petioles and inflorescence stems. In the second type, the dark anthers of Paspalum paspaloides (Michaux) Scribner (= P. distichum ) are the size, shape and colour of aphids (Homoptera; Aphidoidea) and they sway in the wind like swivelling aphids. Similarly, the stems of Alcea setosa (Boiss.) Alef. are covered with dark flecks that look like aphids. Finally, immature pods of three wild annual legumes ( Lathyrus ochrus (L.) DC.; Pisum fulvum Sm.; Vicia peregrina L.) have conspicuous reddish spots, arranged along the pods, that appears to mimic lepidopteran caterpillars. In one of the species ( V . peregrina ) two different mimicking morphs were found. We propose that these morphological traits may serve as herbivore repellent cues and are part of the defence system of the plants.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 393–398.     

Novel cost of a sexually selected trait in the rubyspot damselfly Hetaerina americana: conspicuousness to prey

Conspicuousness to predators frequently has been invoked asa cost of sexually selected traits, but conspicuousness to preyhas not We tested for the latter using rubyspot damselflies(Hetaerina americana) as the predator. Previous work on thisspecies showed that the red spots on male wings are intrasexuallyselected and reduce survival. Since female wings lack red spots,we first compared male and female weight gain rates per unithunting time. Females gained weight significandy faster thanmales in both mg per hour and relative to body weight. We thencompared the weight gain rates of females painted with red wingspots to those of control females painted with clear ink ornot manipulated. Controls gained weight significandy fasterthan red-painted females. Behavioral observations revealed thatred females attempted to capture prey at normal rates and experiencednormal rates of agonistic interference from conspecifics ofboth sexes. Nevertheless, red females captured fewer prey perminute and per capture attempt than did sham-manipulated andunmanipulated controls. We infer that the red spots reducedfemale weight gain rates by increasing their visibility to prey.Close similarity between male and red female weight gain ratesrelative to unmanipulated females suggests that red spots mayalso be a hunting handicap for males. [Behav Ecol 7: 465–473(1996)]     

Major claws make male fiddler crabs more conspicuous to visual predators: a test using human observers

One of the possible costs of the male fiddler crabs enlarged claw can be conspicuousness to predators. This hypothesis was tested using human observers as a model of visual predators. In the European fiddler crab, Uca tangeri Eydoux, the males' major claw is white contrasting with the orange-brownish colour of the carapace and of the feeding claw, and the mudflat background. The following morphotypes were created from close-up photographs taken in nature using an image processing software: male, male without claw, female, female with enlarged claw, male with enlarged claw of the same colour of the feeding claw, male with 75% sized claw, male with 50% sized claw. These morphotypes were then presented in a randomised order to students, using a psychology test software, which allows the measurement of response time in msec. The subjects were allowed to look at the images for an unlimited amount of time, until they detected the individual or until they decided to pass on to another image. Backgrounds (i.e. mudflat picture) without individuals were also presented as a control. Male crabs were detected significantly sooner than females. When we compared males with the claw removed with females with an enlarged claw added, the pattern is reversed and the latter are detected significantly faster. Thus, the enlarged claw seems to be the key feature that makes the individuals more conspicuous. Size and colour seem to be the main aspects of the claw's conspicuousness. The data of these experiments support the initial prediction of males being more conspicuous than females because of their enlarged claw. The possible costs and benefits of this trait, related to predation, are discussed.     

Odour intensity learning in fruit flies

Animals'' behaviour towards odours depends on both odour quality and odour intensity. While neuronal coding of odour quality is fairly well studied, how odour intensity is treated by olfactory systems is less clear. Here we study odour intensity processing at the behavioural level, using the fruit fly Drosophila melanogaster. We trained flies by pairing a MEDIUM intensity of an odour with electric shock, and then, at a following test phase, measured flies'' conditioned avoidance of either this previously trained MEDIUM intensity or a LOWer or a HIGHer intensity. With respect to 3-octanol, n-amylacetate and 4-methylcyclohexanol, we found that conditioned avoidance is strongest when training and test intensities match, speaking for intensity-specific memories. With respect to a fourth odour, benzaldehyde, on the other hand, we found no such intensity specificity. These results form the basis for further studies of odour intensity processing at the behavioural, neuronal and molecular level.     

Learning capabilities enhanced in harsh environments: a common garden approach

Previous studies have suggested that the ability to inhabit harsh environments may be linked to advanced learning traits. However, it is not clear if individuals express such traits as a consequence of experiencing challenging environments or if these traits are inherited. To assess the influence of differential selection pressures on variation in aspects of cognition, we used a common garden approach to examine the response to novelty and problem-solving abilities of two populations of black-capped chickadees (Poecile atricapillus). These populations originated from the latitudinal extremes of the species''s range, where we had previously demonstrated significant differences in memory and brain morphology in a multi-population study. We found that birds from the harsh northern population, where selection for cognitive abilities is expected to be high, significantly outperformed conspecifics from the mild southern population. Our results imply differences in cognitive abilities that may be inherited, as individuals from both populations were raised in and had experienced identical environmental conditions from 10 days of age. Although our data suggest an effect independent of experience, we cannot rule out maternal effects or experiences within the nest prior to day 10 with our design. Nevertheless, our results support the idea that environmental severity may be an important factor in shaping certain aspects of cognition.     

Cannibalism and Intraguild Predation Among Phytoseiid Mites: Are Aggressiveness and Prey Preference Related to Diet Specialization?

We tested whether specialist and generalist phytoseiid mites differ in aggressiveness and prey choice in cannibalism and intraguild predation. Specialists tested were Galendromus occidentalis, Neoseiulus longispinosus, Phytoseiulus persimilis, and P. macropilis; tested were Amblyseius andersoni, Euseius finlandicus, E. hibisci, Kampimodromus aberrans, Neoseiulus barkeri, N. californicus, N. cucumeris, N. fallacis, and Typhlodromus pyri. Aggressiveness of cannibalistic females against larvae was not related to diet specialization except that highly aggressive species were exclusively generalists. Seldom to moderately cannibalizing species occurred in both generalist and specialist phytoseiids. In contrast to aggressiveness in cannibalism, generalists and specialists differed in aggressiveness in intraguild predation. Adult females of specialists were only slightly aggressive against heterospecific larvae, whereas adult females of all generalists except T. pyri were highly aggressive. Adult females of generalists were able to discriminate between con- and heterospecific larvae and preferentially consumed the latter when given a choice. Adult females of specialists except G. occidentalis showed no preference when given a choice between con- and heterospecific larvae. We conclude that aggressiveness in intraguild predation, species recognition and subsequent preferential consumption of heterospecifics when given a choice is common in generalist but not specialist phytoseiids. We discuss the evolutionary pathways that might have led to the difference between specialists and generalists in species discrimination.     

Avoidance of aposematic prey in European tits (Paridae): learned or innate?

Predators may either learn to avoid aposematic prey or may avoidit because of an innate bias. Learned as well as innate avoidancehas been observed in birds, but the existing evidence is basedon experiments with rather few unrelated model species. We comparedthe origin of avoidance in European species of tits (Paridae).First, we tested whether wild-caught birds (blue tits, greattits, crested tits, coal tits, willow tits, and marsh tits)avoid aposematic (red and black) adult firebugs Pyrrhocorisapterus (Heteroptera) more than nonaposematic (brown painted)ones. Larger proportion of birds avoided aposematic than brown-paintedfirebugs in majority of species (except coal tits). Second,we tested whether naive hand-reared birds of 4 species (bluetits, great tits, crested tits, and coal tits) attack or avoidaposematic and nonaposematic firebugs, both novel for them.Behavior of the naive blue tits and coal tits was similar tothat of the wild-caught birds; majority of them did not attackthe firebugs. Contrastingly, the naive great tits and crestedtits behaved differently than the wild-caught conspecific adults;majority of the wild-caught birds avoided the aposematic firebugs,whereas the naive birds usually did not show any initial avoidanceand had to learn to avoid the aposematic prey. Our results showthat the origin of avoidance may be different even in closelyrelated species. Because blue tits and coal tits avoided notonly aposematic firebugs but also their brown-painted form,we interpret their behavior as innate neophobia rather thaninnate bias against the warning coloration.