Density- and frequency-dependent predation of artificial bird nests

Predators are often expected to vary their relative predation rates according to the frequency of prey types in the environment (frequency-dependent predation). The underlying cause for this must lie in some dependency of absolute predation rates on the density of prey types in the environment (density-dependent predation). However, frequency-dependent predation may either be caused by 'simple' density-dependent predation, in which the absolute predation rate on a given prey type depends purely on the density of that type, or by more complex responses in which absolute rates depend also on the density of other prey types. It is usually difficult to distinguish the underlying cause of frequency-dependent predation, because frequencies tend to change as densities change. Here, we describe the results of an experiment conducted to disentangle these phenomena under two prey richness (low and high) conditions. We used artificial bird nests (placed on shrubs and on saplings) baited with quail eggs placed in natural forests as models of natural bird nests. Our results indicate that both the absolute and relative predation rates on the prey types may vary in complex ways. Predation rates depend on a complex interaction between the prey's own density, other prey density and the diversity of prey in the environment. Neglecting to include, or consider, these complexities into analyses may lead to erroneous conclusions in studies of absolute or relative predation rates.     

Ontogenetic changes in foraging tactics of the piscivorous cornetfish <Emphasis Type="Italic">Fistularia commersonii</Emphasis>

Foraging behaviors of the piscivorous cornetfish Fistularia commersonii were observed at shallow reefs in Kuchierabu-jima Island, southern Japan. This fish foraged on two types of prey fishes: one was reef fish that typically dwell on or near substrata (e.g., Tripterygiidae and Labridae), and the other was pelagic fish that shoal in the water column (e.g., Clupeidae and Carangidae). The prey sizes, prey types and foraging behaviors changed as the predator size increased. Prey sizes were largely limited by gape size of the cornetfish, and small predators consumed small prey. The small cornetfish (10–30 cm in total length) fed only on reef fish captured after stalking (where the fish slowly approaches the prey and then suddenly attacks). The stalking was done either solitarily or in foraging association with conspecifics. Large fish (30–120 cm) fed on both types of fishes by stalking and/or chasing (where the fish chases the prey using its high mobility and attacks), either solitarily or in foraging association with con- or heterospecifics. Thus, chasing was only performed by the large cornetfish against pelagic prey fish in associative foraging with other con- and heterospecific predators. As their body sizes increased, F. commersonii began to show a diversification of foraging behaviors, which was strongly related not only to the habitat types and anti-predatory behaviors of the prey fishes but also to associative foraging with con- or heterospecifics, which improves their foraging success.     

Mixed encounters, limited perception and optimal foraging

This article demonstrates how perceptual constraints of predators and the possibility that predators encounter prey both sequentially (one prey type at a time) and simultaneously (two or more prey types at a time) may influence the predator attack decisions, diet composition and functional response of a behavioural predator-prey system. Individuals of a predator species are assumed to forage optimally on two prey types and to have exact knowledge of prey population numbers (or densities) only in a neighbourhood of their actual spatial location. The system characteristics are inspected by means of a discrete-time, discrete-space, individual-based model of the one-predator-two-prey interaction. Model predictions are compared with ones that have been obtained by assuming only sequential encounters of predators with prey and/or omniscient predators aware of prey population densities in the whole environment. It is shown that the zero-one prey choice rule, optimal for sequential encounters and omniscient predators, shifts to abruptly changing partial preferences for both prey types in the case of omniscient predators faced with both types of prey encounters. The latter, in turn, become gradually changing partial preferences when predator omniscience is considered only local.     

Sensory modulation and behavioral choice during feeding in the Australian frog, Cyclorana novaehollandiae

This study investigates how visual and tactile sensory information, as well as biomechanical effects due to differences in physical characteristics of the prey, influence feeding behavior in the frog Cyclorana novaehollandiae. Video motion analysis was used to quantify movement patterns produced when feeding on five prey types (termites, waxworms, crickets, mice and earthworms). Twelve kinematic variables differed significantly among prey types, and twelve variables were correlated with prey characteristics (including mass, length, height and velocity of movement). Results indicate that C.␣novaehollandiae uses a different strategy to capture each prey type. Visual assessment of prey characteristics appeared to be more important in modulating feeding behavior than tactile cues or biomechanical effects. We propose a hierarchical hypothesis of behavioral choice, in which decisions are based primarily on visual analysis of prey characteristics. In this model, the frogs first choose between jaw prehension and tongue prehension based on prey size. If they have chosen jaw prehension, they next choose between upward or downward head rotation based on length and height of the prey. If they have chosen tongue prehension, they next choose between behavior for fast and slow prey. Final decisions may be the result of behavioral fine tuning based on tactile feedback. Accepted: 5 August 1996     

Foraging performance of diet-induced morphotypes in pumpkinseed sunfish (Lepomis gibbosus) favours resource polymorphism

Morphological plasticity can influence adaptive divergence when it affects fitness components such as foraging performance. We induced morphological variation in pumpkinseed sunfish (Lepomis gibbosus) ecomorphs and tested for effects on foraging performance. Young-of-year pumpkinseed sunfish from littoral and pelagic lake habitats were reared each on a 'specialist diet' representing their native habitat-specific prey, or a 'generalist diet' reflecting a combination of native and non-native prey. Specialist and generalist diets, respectively, induced divergent and intermediate body forms. Specialists had the highest capture success on their native prey whereas generalist forms were inferior. Specialists faced trade-offs across prey types. However, pelagic specialists also had the highest intake rate on both prey types suggesting that foraging trade-offs are relaxed when prey are abundant. This increases the likelihood of a resource polymorphism because the specialized pelagic form can be favoured by directional selection when prey are abundant and by diversifying selection when prey resources are restricted.     

Size-based Prey Selectivity and Dietary Shifts in the Jellyfish, Aurelia aurita

Gut content analyses on field-caught Aurelia aurita showed bothquantitative and qualitative change in diet as a function ofmedusa size. Larger medusae tended towards greater numbersand diversity of prey (up to 1550 individual prey representingas many as 13 different prey groups). We also found that medusasize was a good predictor of prey diversity recovered from themedusa gut. While a shift toward greater prey diversity inlarger medusae might be explained by increased contact rateswith 'rare' prey taxa, we also found size-based prey selectivitychanges in A. aurita. We used in situ gut content data to describeselectivity by A. aurita for three prey types representing varyingdegrees of swimming or escape velocity. Fish eggs were usedas a non-swiming prey, and small (     

Modulation and variability of prey capture kinematics in clariid catfishes

Species with narrow or limited diets (trophic specialists) are expected to be less flexible in their feeding repertoire compared to species feeding on a wide range of different prey (trophic generalists). The ability to modulate prey capture kinematics in response to different prey types and prey position, as well as the overall variability in prey capture kinematics, is evaluated in four clariid species ranging from trophic generalist (Clarias gariepinus) to species with morphological specializations and a narrow diet (Channallabes apus and Gymnallabes typus). High-speed video recordings were made of prey captures on two prey that differ in shape, attachment strength and hardness. While the observed amount of strike-to-strike variability in prey capture kinematics is similar for all species and not influenced by prey type, only the two less specialized species showed the ability to modulate their prey capture kinematics in function of the presented prey types. All species did, however, show positional modulation during the strike by adjusting the magnitude of neurocranial elevation. These results indicate that the narrow dietary breadth of trophic specialists is indeed indicative of functional stereotypy in this group of fishes. Although most studies focussing on prey processing found a similar result, the present study is one of the few that was able to demonstrate this relationship when focussing on prey capture mechanics. Possibly, this relationship is less frequently observed for prey capture compared to prey processing because, regardless of prey type, the initial capture of prey requires a higher amount of variability.     

Functional responses of five cyprinid species to planktonic prey

Synopsis The functional responses of five species of cyprinids (Chalcalburnus chalcoides, Vimba vimba, Abramis brama, Rutilus rutilus, and Scardinius erythrophthalmus) feeding on four planktonic prey types were measured in the laboratory. Although no alternative prey types were present, the response curves were sigmoid in most cases, because attack rates were not independent of prey density. The findings are explained as being the overt expression of the fishes& foraging tactics. The chief way of maximizing food uptake, according to our interpretation, is accelerating attack rates with increasing prey density. The ability of prey to escape or relative prey size may interfere with this strategy. C. chalcoides, the only obligatory planktivore among the species studied, attacks at higher rates and responds most markedly to changes in prey density.     

To eat and not be eaten: modelling resources and safety in multi-species animal groups

Using mixed-species bird flocks as an example, we model the payoffs for two types of species from participating in multi-species animal groups. Salliers feed on mobile prey, are good sentinels and do not affect prey capture rates of gleaners; gleaners feed on prey on substrates and can enhance the prey capture rate of salliers by flushing prey, but are poor sentinels. These functional types are known from various animal taxa that form multi-species associations. We model costs and benefits of joining groups for a wide range of group compositions under varying abundances of two types of prey-prey on substrates and mobile prey. Our model predicts that gleaners and salliers show a conflict of interest in multi-species groups, because gleaners benefit from increasing numbers of salliers in the group, whereas salliers benefit from increasing gleaner numbers. The model also predicts that the limits to size and variability in composition of multi-species groups are driven by the relative abundance of different types of prey, independent of predation pressure. Our model emphasises resources as a primary driver of temporal and spatial group dynamics, rather than reproductive activity or predation per se, which have hitherto been thought to explain patterns of multi-species group formation and cohesion. The qualitative predictions of the model are supported by empirical patterns from both terrestrial and marine multi-species groups, suggesting that similar mechanisms might underlie group dynamics in a range of taxa. The model also makes novel predictions about group dynamics that can be tested using variation across space and time.     

The effect of prey type and density on the foraging efficiency of juvenile perch, (Perca fluviatilis)

The foraging efficiency of juvenile perch (Perca fluviatilis), feeding on two types of prey, was studied in laboratory experiments. Waterfleas (Daphnia magna) and phantom midge larvae (Chaoborus flavicans) were offered in a range of densities, either separately or combined. Perch fed more efficiently on each prey type separately than when both were mixed. Foraging efficiency decreased with an increase of mixed prey density with both prey types present in equal numbers, but also when the proportion of Chaoborus increased. This could be caused by the existence of different hunting techniques, each of which is fully efficient in the presence of one prey type only. In the presence of two prey types, the predator constantly has to switch from one hunting technique to another.     

Does prey community composition affect the way different behavioral types interact with their environment?

We examined how different exploratory behavioral types of largemouth bass responded to differing prey communities by determining effects on growth, survival and diet in experimental ponds. We found evidence that non-explorer largemouth bass target young-of-year bluegill early on in life, but bluegill were not an important diet item by late summer. The presence of young-of-year bluegill as prey does appear to affect the foraging strategy of the two exploring types differently. In the absence of small bluegill, both behavioral types feed primarily on benthic invertebrates and zooplankton. When small bluegill were present, we saw a shift away from zooplankton as prey for largemouth bass. However, that shift was toward more benthic invertebrates for non-exploring behavioral types and toward terrestrial insects for exploring behavioral types. Thus, it appears that prey community composition can have important effects on the way in which different behavioral types interact with their environment.     

Predation risk and the structure of freshwater zooplankton communities

Summary Many predators inflict substantial mortality on their prey. The prey respond to these selective pressures with changes in their spatial and temporal overlap with the predator (density risk responses), or with changes in their vulnerability to the predator (prey vulnerability responses). Here we develop a simple predation model that permits quantification of the basic response types of the prey in nature. We then test the hypothesis that prey response will be proportional to the intensity of the predation mortality relative to all other sources of mortality and decreased natality acting on the prey. A significant regression relationship is obtained for the prey vulnerability response but not for any of the density risk responses. The individual response values and regression statistics are used to interpret the relative importance of the different response types and to assess the predator's influence on prey community structure.Supported by Lehigh University Environmental Studies Center     

Should fatty acid signature proportions sum to 1 for diet estimation?

Knowledge of predator diets, including how diets might change through time or differ among predators, provides essential insights into their ecology. Diet estimation therefore remains an active area of research within quantitative ecology. Quantitative fatty acid signature analysis (QFASA) is an increasingly common method of diet estimation. QFASA is based on a data library of prey signatures, which are vectors of proportions summarizing the fatty acid composition of lipids, and diet is estimated as the mixture of prey signatures that most closely approximates a predator’s signature. Diets are typically estimated using proportions from a subset of all fatty acids that are known to be solely or largely influenced by diet. Given the subset of fatty acids selected, the current practice is to scale their proportions to sum to 1.0. However, scaling signature proportions has the potential to distort the structural relationships within a prey library and between predators and prey. To investigate that possibility, we compared the practice of scaling proportions with two alternatives and found that the traditional scaling can meaningfully bias diet estimators under some conditions. Two aspects of the prey types that contributed to a predator’s diet influenced the magnitude of the bias: the degree to which the sums of unscaled proportions differed among prey types and the identifiability of prey types within the prey library. We caution investigators against the routine scaling of signature proportions in QFASA.     

Effects of nutrients and predators on an old-field food chain: interactions of top-down and bottom-up processes

In theory, selection favours predators that select prey in order to maximise reproductive success. We studied the association between preference and performance of the generalist predator Orius laevigatus with respect to two prey species: spider mites ( Tetranychus urticae ) and western flower thrips ( Frankliniella occidentalis ). Under ample prey supply, the predators had higher maximum reproductive success (measured as intrinsic population growth rate r ) on thrips than on spider mites; hence thrips represent a higher prey quality to the bugs. This was at odds with the observed preference of the predatory bug for plants (patches) with high densities of spider mites to plants with moderate densities of thrips in release-recapture experiments. Thus, prey quality does not suffice to explain the preference of predators for plants with prey. The quality of a patch as an oviposition site (i.e. the number of eggs produced on a patch per bug per day) also did not match preference patterns. Hence, patch preference was not correlated to prey quality or oviposition rate on prey patches. However, patch productivity, i.e. the total number of offspring surviving until adulthood that can be produced by one female on a patch, was correlated with preference. This was further tested by offering the predators a choice between plants with high densities of spider mites and plants with high densities of thrips in an independent set of release-recapture experiments. These two types of prey patches were found equivalent in terms of patch productivity. Indeed, the predators showed no preference for either of the two types of patches, which is in agreement with our predictions. This suggests that the predatory bugs select patches based on patch productivity rather than on prey quality or oviposition rate on a patch.     

Cooperation and Prey Capture Efficiency in a Social Spider,Anelosimus eximius (Araneae,Theridiidae)

We studied the efficiency of the hunt and the characteristics of cooperation during the prey capture in a social spider Anelosimus eximius. Two natural types of prey of roughly the same length (20 mm) were used: grasshoppers (Orthoptera) and moths (Lepidoptera); 128 tests were made on 14 colonies, the smallest with 20 and the largest with 1,700 individuals. Test times were 12.00 h, defined as an inactive period for the spiders and 18.00 h, defined as an active period. Overall capture rate of intercepted prey was 66%: it was higher in large colonies or at 18.00 h, when more spider alerts were triggered by the struggling prey. Characteristics of cooperation during capture did not vary with colony size. Capture rate was higher for grasshoppers than moths (73%-58%) in spite of similar number of alerts (76%-87%); so moths must have been more difficult to capture. For both prey types, large colonies capture more rapidly and so had advantages in terms of time gain. We showed that cooperation depended on prey type: more spiders mobilised to attack moths and attack was faster than on grasshoppers. This may be interpreted as an adaptive response of the group to the prey type.     

Evolutionary consequences of a search image

Many predators are able to become better at spotting cryptic prey by recognising specific clues, but by concentrating on one prey type they will become worse at spotting other prey types. This phenomenon is known as the formation of a search image for a certain prey by a predator and is related to apostatic selection. Here, we study the evolution of a search image in the predator by formulating and analysing a mathematical model. The predator forages for two prey types and is able to form an independent search image for both prey. The results show that the evolutionary dynamics can be divided into two parts: a fast and a slow part. At first selection pressure will be strong towards a stable ratio of prey, which is the same as the ratio found for the unbeatable prey choice for predators with a Holling type II functional response. Following this, the slow dynamics will keep this ratio constant independent of the trait values, but the predator will slowly evolve towards a stronger search image and ultimately become a specialist predator or slowly evolve towards generalist with a weak search image. In conclusion, the formation of a search image causes the predator to control the prey densities such that the ratio of available prey is kept constant by the predator.     

Prey selection by thaidid gastropods: some observational and experimental field tests of foraging models

Summary Field observations and experiments revealed that predatory intertidal gastropods of the genus Thais (or Nucella) were able both to recognize the expected food value of encountered prey (expected energy or growth potential gained per unit handling time) and to monitor their average yield over time (average energy or growth potential gained per unit foraging time). They appeared to discriminate not only among prey species, but also among different sized individuals of the same prey species. The evidence supporting these interpretations included: 1) field observations of snails feeding preferentially on prey types of higher expected food value even though lower value prey types were available and abundant, 2) a very limited number of direct underwater observations of foraging snails rejecting encountered items that were either of lower expected value than the item finally eaten or not measurably different from it, and 3) field (=arena) experiments in which both average yield, and the distribution and abundance of potential prey were controlled: snails conditioned at a high average yield fed preferentially on high value items, while those animals conditioned at a low yield consumed prey in the proportions that they were encountered. These behaviors are all consistent with a prey-selection decision motivated by energetic considerations. Further, the field experiments indicated that these predatory gastropods could select items from a diverse array of prey so as to maximize growth in their natural environment. The behaviors were not consistent with three alternative foraging hypotheses: non-selective foraging, frequency-dependent foraging on prey types (here, sizes of particular prey species), and frequency-dependent foraging on prey species. Deviations from some of the quantitative predictions of optimal foraging theory appeared related to learning and risk.     

Discrete prey availability promotes foraging segregation and early divergence in Arctic charr, <Emphasis Type="Italic">Salvelinus alpinus</Emphasis>

Many animal species show individual foraging specialisms when potential prey requires prey-specific foraging strategies. Arctic charr are often found as benthic (macroinvertebrate) or pelagic (plankton) foraging specialists. Here, we tested specifically if given a choice of prey with different characteristics individuals would specialise in a single prey type and if individuals would chose prey based on their expressed trophic morphology, in a laboratory experiment and in a field observation. When offered a choice of benthic and pelagic prey most individuals (73%) showed that 100% fidelity to a single foraging source. Naïve individuals (not previously exposed to natural prey) with more robust head and mouth shape were more likely to forage on a benthic prey source (chironomids). In contrast, individuals with a more fusiform body, larger eye, but more slender head shape were more likely to specialise on pelagic prey (Artemia). Field observations of a natural population of Arctic charr from Loch Doine identified specialists foraging on either plankton or macrobenthos (on the basis of stomach contents) and some generalists. Morphological analysis showed that significant differences in shape reflecting recent foraging history. These results support the hypothesis that the availability of discrete, different prey types results in discrete foraging specialisms which in turn may result in the expression of discrete alternative phenotypes through subsequent plastic ontogenetic process. We conclude that this provides a partial explanation for why ecologically driven evolution processes are particularly prevalent in fishes from post-glacial lake systems.     

Prey capture and selection throughout the breeding season in a deep‐diving generalist seabird,the thick‐billed murre

Generalist seabirds forage on a variety of prey items providing the opportunity to monitor diverse aquatic fauna simultaneously. For example, the coupling of prey consumption rates and movement patterns of generalist seabirds might be used to create three‐dimensional prey distribution maps (‘preyscapes’) for multiple prey species in the same region. However, the complex interaction between generalist seabird foraging behaviour and the various prey types clouds the interpretation of such preyscapes, and the mechanisms underlying prey selection need to be understood before such an application can be realized. Central place foraging theory provides a theoretical model for understanding such selectivity by predicting that larger prey items should be 1) selected farther from the colony and 2) for chick‐feeding compared with self‐feeding, but these predictions remain untested on most seabird species. Furthermore, rarely do we know how foraging features such as handling time, capture methods or choice of foraging location varies among prey types. We used three types of animal‐borne biologgers (camera loggers, GPS and depth‐loggers) to examine how a generalist Arctic seabird, the thick‐billed murre Uria lomvia, selects and captures their prey throughout the breeding season. Murres captured small prey at all phases of a dive, including while descending and ascending, but captured large fish mostly while ascending, with considerably longer handling times. Birds captured larger prey and dove deeper during chick‐rearing. As central place foraging theory predicted, birds travelling further also brought bigger prey items for their chick. The location of a dive (distance from colony and distance to shore) best explained which prey type was the most likely to get caught in a dive, and we created a preyscape surrounding our study colony. We discuss how these findings might aid the use of generalist seabirds as bioindicators.     

Ecological and evolutionary implications of diet in monitor lizards

A survey of 35 species indicates that monitor lizards (Varanus) typically hunt over large areas, search in particular microhabitats, and feed frequently on a wide variety of prey, many of which are relatively small. There is ontogenetic, seasonal, and geographic variation in diet. With some exceptions, invertebrates are the predominant prey, but rare predation on vertebrates is often energetically significant. A few monitors specialize on prey types that occur as occasional items in the diet of species with more generalized diets; these include crabs, snails, orthopterans, lizards, and large mammals. For most species, prey specialization occurs via habitat selection and a variety of prey types and sizes are eaten, as expected for widely searching predators. Comparisons with other anguimorphans suggest that derived features of Varanus are associated with high body temperature and activity levels; specialized chemoreception; and rapid, skillful capture of hidden and/or potentially hard to catch prey. Occasional ingestion of moderately large prey is primitive for Varanoidca (Helodermatidae +Varanidae), accentuating a trend that is perhaps primitive for anguimorphan lizards. Reduction of very large prey prior to ingestion is a derived attribute within Varanus , seen infrequently in several larger species and commonly in V. komodoensis. This study illustrates the synthesis of comparative natural history in a phylogenetic context, a method that addresses the history of organismal change.