Selection by passerine birds is anti-apostatic at high prey density

Most of the results from past experiments with wild birds and green and brown pastry 'baits' have suggested that disproportionately more of the rare forms are eaten when bait density is high (i.e. selection is anti-apostatic). In two separate series of experiments we presented birds with dishes containing 270 baits of one colour and 30 of another. In series I, five different pairs of colours were presented simultaneously to wild birds at two sites. One colour of each pair was common at one site and the same colour was rare at the other site. After 35 days the ratios of the colours were reversed and the dishes were presented for another 35 days. There was a statistically significant tendency for the colours to be at a higher risk when rare. In series II, three caged blackbirds were offered green and brown baits in two dishes simultaneously; in one dish green was rare and in the other brown was rare. Selection over 6 days was anti-apostatic for all three birds combined but the data proved heterogeneous both between and within individuals. At any one time, each bird tended to concentrate on one colour, irrespective of whether that colour was rare or common. We believe that this behaviour leads to anti-apostatic selection, as has been observed in these and other experiments with pastry prey.     

Fast movement of densely aggregated prey increases the strength of anti-apostatic selection by wild birds

'Anti-apostatic' selection occurs when predators preferentially remove rare forms of prey, and has been demonstrated in both static (artificial) prey and moving (natural) prey. We performed 24 experiments at separate sites to test the effect of prey mobility on the strength of anti-apostatic selection by wild passerine birds. The prey were yellow and red Calliphora maggots and were presented in 9: 1 and 1: 9 ratios on a specially designed bird table. The temperature of the maggots, and hence their speed of movement, was adjusted by a combination of the ambient temperature and a candle under the bird table. Selection was anti-apostatic at all three classes of temperature and was strongest at the highest. We conclude that anti-apostatic selection on static prey is enhanced when they are made to move–possibly because the 'confusion effect' caused by the moving prey makes the birds concentrate more strongly on the rarer colours.     

Selection by wild birds on artificial dimorphic prey on varied backgrounds

Our aim was to test the effects of prey frequency and background composition on selection by free-ranging birds. We did three series of experiments with populations of grey and orange pastry prey scattered among coloured stones that made the prey either conspicuous or inconspicuous. Series 1 tested whether the predicted equilibrium frequency of the two prey types was influenced by the frequency of matching grey and orange stones. Birds at a single site were given a random sequence of combinations of prey frequency and stone frequency. Selection was dependent on background and the effect of prey frequency also varied with background. In series 2, we explored the frequency-independent effect of background: birds at five sites were given equal numbers of the two prey in three frequencies of matching stones and two of non-matching. There was a higher risk of predation for prey that matched rarer stones. In series 3 we attempted to measure, at a single site, the actual equilibrium prey frequencies in three different backgrounds: two extreme stone frequencies and one intermediate. Each experiment started with a population of equal numbers of grey and orange prey. After half the prey had been eaten we calculated the frequencies of the survivors and presented a new population of the original size but with the new prey frequencies; each experiment ran for 25 such 'generations'. The results suggested that at equilibrium the commoner 'morph' was the one that resembled the commoner colour of stone. Overall, our findings support the idea that visual selection can result in morph frequencies becoming related to the proportions of their matching background components and that this equilibrium will 'track' temporal or spatial changes in the background.     

The evolution of plumage polymorphism in birds of prey and owls: the apostatic selection hypothesis revisited

Co-evolution between phenotypic variation and other traits is of paramount importance for our understanding of the origin and maintenance of polymorphism in natural populations. We tested whether the evolution of plumage polymorphism in birds of prey and owls was supported by the apostatic selection hypothesis using ecological and life-history variables in birds of prey and owls and performing both cross taxa and independent contrast analyses. For both bird groups, we did not find any support for the apostatic selection hypothesis being the maintaining factor for the polymorphism: plumage polymorphism was not more common in taxa hunting avian or mammalian prey, nor in migratory species. In contrast, we found that polymorphism was related to variables such as sexual plumage dimorphism, population size and range size, as well as breeding altitude and breeding latitude. These results imply that the most likely evolutionary correlate of polymorphism in both bird groups is population size, different plumage morphs might simply arise in larger populations most likely because of a higher probability of mutations and then be maintained by sexual selection.     

The influence of prey identity and size on selection of prey by two marine fishes

Laboratory experiments were conducted with two predatory fishes, Lagodon rhomboides (Linnaeus) and Syngnathus floridae (Jordan & Gilbert), to determine if prey selection was a result of predator preference or prey accessibility. Prey consisted of two species of caridean shrimp, Tozeuma carolinense (Kingsley) and Hippolyte zostericola (Smith), that commonly inhabit seagrass meadows. Natural diets of both predators revealed that selection ofTozeuma and Hippolyte was not related to their field densities. My experiments demonstrated that natural prey selection was a consequence of prey accessibility, not predator preference. Experiments examining the role of prey size in predator preference revealed that large individuals were significantly preferred over small individuals. Observations of prey behavior indicated that prey motion affected predator choice. These results suggest that predator preference was primarily determined by prey visibility and that the combined effects of prey size and motion contributed to the visibility of these prey.     

Perceptual Processes and the Maintenance of Polymorphism Through Frequency-dependent Predation

One of the key challenges of both ecology and evolutionary biology is to understand the mechanisms that maintain diversity. Negative frequency-dependent selection is a powerful mechanism for maintaining variation in the population as well as species diversity in the community. There are a number of studies showing that this type of selection, where individuals of a rare type (i.e. a rare morph or a rare species) experience higher survival than those of more common type(s). However, it is still not clear how frequency-dependent selection operates. Search image formation has been invoked as a possible, proximate explanation. Although the conceptual link between search image and frequency-dependent predation is often assumed in ecological and evolutionary studies, a review of the literature reveals a paucity of evidence demonstrating the occurrence of both in a natural predator-prey system. Advances in the field of psychology strongly support the existence of search image, yet these findings are not fully recognized in the realm of ecology and evolutionary biology, in part, we feel because of confusion and inconsistencies in terminology. Here we try to simplify the language, clarify the advances in the study of frequency-dependent predation and search image, and suggest avenues for future research. We feel that the investigations of both proximate (perceptual mechanisms) and ultimate (pattern of predation) processes are necessary to fully understand the importance of individual behavioural processes for mediating evolutionary and ecological diversity.Co-ordinating editor: O. Leimar     

Influence of extraguild prey density on intraguild predation by heteropteran predators: A review of the evidence and a case study

Heteropteran predators constitute an important component of predatory guilds in terrestrial and aquatic ecosystems. Most heteropteran species have generalist diets, and intraguild predation has been documented in most heteropteran families. Zoophytophagous species also frequently engage in intraguild interactions. An increase in extraguild prey density is often predicted to reduce intraguild predation between guild members by providing abundant alternate prey. However, an increase of extraguild prey density may also be associated with an increase in the density of intraguild predators, which could instead strengthen intraguild predation. Evaluating the combined effect of these potentially opposing influences on intraguild predation is difficult. Most studies have been carried out in the laboratory, using artificially simplified communities of predators and prey and employing spatial and temporal scales that may not reflect field conditions. We review experimental studies examining how extraguild prey density influences the intensity of intraguild predation and then report an observational case study examining the influence of extraguild prey density on the intensity of intraguild predation at larger spatial and temporal scales in unmanipulated cotton fields. Fields with more abundant extraguild prey (aphids, mites) were not associated with elevated densities of intraguild predators, and were strongly associated with increased survival of intraguild prey (lacewing larvae). In this system, the ability of extraguild prey to relax the intensity of intraguild predation, as previously documented in small-scale field experiments, also extends to the larger spatial and temporal scales of commercial agriculture.     

Effects of prey density,prey size and predator size on rates of feeding by an intertidal predatory gastropod Morula marginalba Blainville (Muricidae), on several species of prey

As a prerequisite for models of foraging behaviour of the whelk, Morula marginalba Blainville (Muricidae), the effects of variation in density of prey on the rate of feeding of the predator were examined in field conditions for three coexisting species of prey. Densities of prey used were those at which the prey, two limpets and a barnacle, occurred naturally in the rocky intertidal habitat.Large limpets, Cellana tramoserica (Sowerby) can resist attacks by predatory gastropods by raising the mantle over the outside of the shell. These experiments showed that no C. tramoserica were killed by Morula marginalba even at very great densities and with no alternative prey present. For the small limpet Patelloida latistrigata (Angas), one of the whelk's most highly preferred prey, juveniles were eaten 1.4 times as fast as adults. Fitting the random predator equation gave greater attack coefficients and shorter handling times for juvenile than adult limpets.Sizes of both predator and prey affected rates of eating barnacles, Tesseropora rosea (Krauss), but not in a simple way. Whelks of 15-mm aperture length ate adult barnacles 4.2 times faster than did 12-mm whelks, but there was no significant difference in the rates at which the two sizes of snail ate juvenile barnacles.Rates of feeding on T. rosea and Patelloida latistrigata increased significantly with prey density. These results form a basis for including the density of prey in models of spatial dispersion of the predatory gastropod Morula marginalba.     

The effect of predator and prey density on the induced defence of a ciliate

1. The level of antipredator defence should be proportional to the actual attack probability to minimize the cost of defence and maximize the net benefit.
2. The hypothesis that the induced antipredator morphology of Euplotes octocarinatus is a graded response to the actual risk of predation by Stylonychia mytilus was tested by manipulating the density of both prey and predator populations.
3. The magnitude of the response was graded according to both predator and prey density. A dense prey population may be protective since a prey is more exposed to a predator's attack as a solitary individual.
4. The results suggest that Euplotes is able to 'estimate' the real risk of predation and respond appropriately, without mobilizing more resources than needed.
5. Separation of the prey and predator with a nylon net revealed that the response was not induced by a water-transmitted factor but that direct cell-to-cell contacts were important. This finding departs from those of other studies.     

Feeding by the aquatic heteropteran,Diplonychus rusticum (Belostomatidae): an effect of prey density on meal size

The behaviour of Diplonychus rusticum feeding on chironomid larvae has been investigated under laboratory conditions. Changes in the percentage of material extracted from prey indicated that feeding for the first two minutes enabled the predator to obtain approximately 33% of the available food; feeding for 10 minutes resulted in only 60% extraction. Comparing the percentage of each prey consumed by D. rusticum exposed to various prey densities, it was apparent that predators were more wasteful and ate less of each prey as chironomid density increased. Because the rate of food intake declined as a greater proportion of each prey was extracted, predators exposed to high chironomid densities reduced the amount of each prey consumed thereby conforming to a simple optimal feeding model.     

Prey Selection by the Lady Beetle Harmonia axyridis: The Influence of Prey Mobility and Prey Species

The influence of prey mobility and species on prey selection by the coccinellid Harmonia axyridis Pallas was determined under laboratory conditions for two prey species, Hyaliodes vitripennis (Say) and Tetranychus urticae Koch. Prey selection was influenced by prey mobility. In the presence of active prey, the coccinellid selected T. urticae while in presence of immobilized prey, H. vitripennis was preferred. Harmonia axyridis searching time was longer in the presence of active H. vitripennis than in the presence of active T. urticae. Moreover, the coccinellid capture rate was lower for active H. vitripennis caused by effective defensive mechanisms. Prey suitability was affected by prey mobility and species. Immobilized H. vitripennis were the most profitable prey, i.e. induced a shorter developmental time and no mortality. However, active H. vitripennis were not a suitable food source for H. axyridis. Our results suggested that three factors are involved in prey selection by H. axyridis: (i) prey mobility; (ii) prey defensive mechanisms; and (iii) prey species.     

The influence of prey hardness on Daphnia's selectivity for large prey

Two comparable methods were used to study the feeding of four species of Daphnia on large spherical particles which differed in size and hardness. The first method used gut analysis to estimate the selectivities of daphnids feeding in a broad size range of a single particle type, including polystyrene beads (4–60 µm diameter) in the laboratory and Eudorina colonies (10–90 µm) in the field. In the second method, Daphnia of different sizes fed in a mixture of 6.5 µm Chlamydomonas and one of eight test particles. Smaller daphnids were less effective in feeding on large test particles. Nonlinear regression was therefore used to estimate the Daphnia body size at which the clearance rate on a test particle was reduced to 50% of that for Chlamydomonas. The results of both methods show that prey size and hardness are both very important in determining daphnid feeding selectivity. For a given particle size, soft algae (naked and gelatinous flagellates) are more readily ingested than hard algae (diatoms and dinoflagellates), and hard algae are more readily ingested than polystyrene beads. Daphnia can feed effectively on algae that are 2–5 times larger than the largest ingestible bead.     

The influence of prey size and abundance, and individual phenotype on prey choice by the three-spined stickleback, Gasterosteus aculeatm L.

Prey selection behaviour of three-spined sticklebacks, Gasterosteus aculeatus L., was studied in two experiments. Where possible, the experimental apparatus satisfied the assumptions of the simplest optimal diet model (the basic prey model); prey were presented sequentially, the fish could not search for and handle prey at the same time, and net energy gain, handling time and encounter rate were fixed. Experiment 1 presented fish with a range of Asellus sizes so that pursuit ( p ) and handling ( h ) time could be related to prey size. Published energy values of Asellus together with pursuit and handling times were used to calculate E /( p+h ) for Asellus measuring 3,4,5,6,7 and 9 mm. Pursuit times did not differ with prey size but handling times did. E /( p+h ) was very variable particularly at the larger prey sizes. Experiment 2 presented fish with two sequences of prey differing in the encounter rate with the most profitable prey sizes. Fish did not select the diet predicted by the basic prey model tending to always ignore the largest prey even when net energy gain would have been maximized by including them in the diet. Further analysis showed that the probability of a prey size being taken was a function of prey size, fish stomach fullness and encounter rate. It is concluded that the basic prey model is too simple to capture the behaviour of the fish. One of its main faults is that the changing state of the fish through the feeding bout is ignored.     

Patterns of prey selectivity in the cyclopoid copepod Mesocyclops thermocyclopoides

In the shallow and eutrophic subtropical aquatic ecosystems, which it generally inhabits, the omnivorous copepod Mesocyclops thermocyclopoides encounters a wide variety of animal prey types including ciliates, rotifers, and cladocerans. We studied prey selectivity in laboratory-reared adult females of this species given a choice of (i) prey types belonging to different taxa (ciliates, rotifers, cladocerans, and cyclopoid nauplii), and (ii) different prey species within a taxonomic group differing in body size, morphology or behaviour. We also tested the effect of different proportions of prey species on its selectivity. Prey type proportion had no significant effect on selectivity of the copepod, nor was there any evidence of switching based on the relative abundance of prey. Among the ciliate prey species tested, the largest species, Stylonychia mytilus was positively selected regardless of its relative abundance, while the smallest, S. notophora was selected only when its density was higher. Offered a choice of three species of a brachionid rotifer differing in size, the copepod selected the largest of them, Brachionus calyciflorus, and avoided the smallest B. angularis. The evasive rotifer Hexarthra mira was also avoided. When prey choice included three cladoceran species Daphnia similoides, Moina macrocopa and Ceriodaphnia cornuta, the copepod selected the intermediate-sized M. macrocopa regardless of the abundance of the other two species. Although it fed on Mesocyclops nauplii when there was no choice, M. thermocyclopoides avoided them when alternative food was available. In a multispecies prey choice test, the copepod selected predominantly the rotifer B. calyciflorus and the cladoceran M. macrocopa. We suggest that the prey selectivity patterns shown by M. thermocyclopoides are adaptive in that they lead to ingestion of the most profitable prey.     

Selection of waterbirds by Geoffroy's cat: Effects of prey abundance, size, and distance

In spite of its wide distribution, Geoffroy's cat (Oncifelis geoffroyi) has been recently upgraded from Least Concern IUCN category to Near Threatened because of lack of knowledge. In this study, we examine at the Mar Chiquita Biosphere Reserve (37° 46`S, 57° 27`W; Argentina) their seasonal diet, and their feeding preference in relation to waterbird abundance, size, and distance of waterbird prey before attack. The diet, evaluated by feces analysis, was comprised of 42% waterbirds and 40% small mammals. The relative importance of waterbirds in the diet changed throughout the year associated with seasonal fluctuations in their availability. Predation on waterbirds increased from autumn to summer, while predation on small mammals decreased during this period. Trophic-niche breadth for prey occurrence was higher in autumn and lower in spring. We found that abundance, distance of waterbird prey before attack, and prey size were significant predictors of waterbird consumption by the Geoffroy's cat. This small cat appears to change its kill rate in relation to waterbird abundance changes. The overconsumption of prey close to vegetation and big prey indicates selective predatory behavior by this cat.     

Prey Selection by the Water Bug Aphelocheirus aestivalis Fabr. (Heteroptera: Aphelocheiridae)

Feeding experiments were conducted in situ in flow through boxes to determine the vulnerability of different prey types (the mayfly larvae Baetis, Ephemerella and Ecdyonurus spp., the caddisfly larva Hydropsyche spp. and the crustacean Gammarus spp.) to the predaceous water bug Aphelocheirus aestivalis and to estimate the predation impact of Aphelocheirus on prey populations. Experiments with equal densities of mixed prey and experiments where each prey was tested individually revealed that Baetis and Ephemerella were most vulnerable to Aphelocheirus predation: Hydropsyche, Ecdyonurus and Gammarus were little/not preyed upon. The present results suggest that vulnerability of prey depends mainly upon prey mobility and microhabitat overlap between predator and prey and that feeding behaviour of Aphelocheirus resembles more that of megalopterans than of stoneflies. Experiments with different prey densities (120–720 prey m⁻²) showed that the predation rate of Aphelocheirus increased with increasing prey density. Experiments with different substrates documented that mortality rates of prey decreased with increasing substrate complexity. When substrate conditions were complex mortality rates in the control and in the experimental boxes were the same which suggest little importance of Aphelocheirus predation on mayflies in the study site.