Prey–predator dynamics with predator switching regulated by a catabolic repression control mode

Since many predators can live under certain circumstances as saprophytes or consume more than one prey, and different enzymes are generally required for each prey or nutrient digestion, the predator must be sufficiently adaptive for effective utilization of the prey mass. Control modes as induction and repression, however, act at the level of genes and cause changes in the biosynthesis rate of these enzymes. In this work, an extension of the catabolic repression control mode from the level of genes to the level of the behavior of the predator is proposed, in order to model the balanced attack of the predator on the prey. It is demonstrated that, when the prey population has the competitive advantage over the predator (in using the common substrate), the catabolic repression mechanism favors the prey population, which dominates over the predator even at low specific dilution rate values, whereas, the stable steady or periodic coexistence state is not favored. When the predator has the competitive advantage at low substrate concentrations and the prey at high substrate concentrations, the introduction of the catabolic repression mechanism in the model again favors the stable steady state of the prey, while the coexistence region is dramatically reduced. Conversely, when the prey population has the competitive advantage at low and the predator at high substrate concentrations, dominance of prey and coexistence steady state could be favored by the catabolic repression mechanism. It is concluded that the catabolic repression control favors dominance of the prey population and, under certain circumstances, coexistence of both prey and predator populations.     

Prey density and the behavioral flexibility of a marine predator: the common murre (Uria aalge)

Flexible time budgets allow individual animals to buffer the effects of variable food availability by allocating more time to foraging when food density decreases. This trait should be especially important for marine predators that forage on patchy and ephemeral food resources. We examined flexible time allocation by a long-lived marine predator, the Common Murre (Uria aalge), using data collected in a five-year study at three colonies in Alaska (USA) with contrasting environmental conditions. Annual hydroacoustic surveys revealed an order-of-magnitude variation in food density among the 15 colony-years of study. We used data on parental time budgets and local prey density to test predictions from two hypotheses: Hypothesis A, the colony attendance of seabirds varies nonlinearly with food density; and Hypothesis B, flexible time allocation of parent murres buffers chicks against variable food availability. Hypothesis A was supported; colony attendance by murres was positively correlated with food over a limited range of poor-to-moderate food densities, but independent of food over a broader range of higher densities. This is the first empirical evidence for a nonlinear response of a marine predator's time budget to changes in prey density. Predictions from Hypothesis B were largely supported: (1) chick-feeding rates were fairly constant over a wide range of densities and only dropped below 3.5 meals per day at the low end of prey density, and (2) there was a nonlinear relationship between chick-feeding rates and time spent at the colony, with chick-feeding rates only declining after time at the colony by the nonbrooding parent was reduced to a minimum. The ability of parents to adjust their foraging time by more than 2 h/d explains why they were able to maintain chick-feeding rates of more than 3.5 meals/d across a 10-fold range in local food density.     

Habitat use by juvenile Atlantic cod (Gadus morhua) in the presence of an actively foraging and non-foraging predator

Experiments were conducted in the autumn and winter of 1992/93 to examine habitat use by juvenile (age 0+) Atlantic cod, Gadus morhua L., before, during and following exposure to a passive or actively foraging predator (age 3+ cod). Experiments presented groups of juvenile cod (n=5 fish/group) with one of two combinations of three substrates; (1) gravel, sand, and a patch of artificial kelp (kelp), or (2) cobble, sand, and kelp. Cobble is known to provide juvenile cod with a refuge from predation. Kelp was used to test the hypothesis that juvenile cod associate with fleshy macroalgae in nature because of the safety it provides from predators. There was little difference in habitat use by juvenile cod before, during or following exposure to a passive predator. Under these conditions, juvenile cod appeared to prefer finer grained mineral substrates and avoided the kelp. The extent of the juvenile response to a passive predator was to avoid the predator's location in the experimental tank. In contrast, juvenile cod showed a significant shift in habitat use when exposed to an actively foraging predator, hiding in cobble or, when cobble was not available, in kelp. Use of both these habitats resulted in a significant reduction in predation risk to the juvenile cod. Our results suggest that: (1) an association with kelp provides safety from predation to juvenile cod, and (2) juvenile cod are capable for assessing the risk a predator represents and adjust their response accordingly.     

Oil-induced disruption of foraging behaviour of the asteroid keystone predator, Coscinasterias muricata (Echinodermata)

A prey-localization test allowed the observation of the foraging behaviour of Coscinasterias muricata in the presence of Mytilus edulis and the disruption of this behaviour following oil exposure. Asteroids were exposed to dilutions of water-accommodated fractions (WAF) of Bass Strait stabilised crude oil (control, 2%, 10%). Effects of exposure (4 d) and depuration (20 d) were quantified using circular statistical analyses. Observations suggested that disruption of behaviour might be concentration-dependent. Control asteroids successfully located mussels during tests. A small proportion (3 of 16) of asteroids exposed to 2% WAF also located the mussels, while asteroids exposed to 10% WAF did not. Following depuration, exposed asteroids recovered their chemoreception capacities. It is concluded that (1) Coscinasterias muricata is able to locate prey mussels through chemoreception, (2) exposure to oil disrupts its foraging behaviour, and (3) the effect is reversible. Received: 4 March 1998 / Accepted: 25 November 1998     

Hunger-dependent predator inspection and foraging behaviours in the threespine stickleback (Gasterosteus aculeatus) under predation risk

Individual fish commonly leave the relative safety of the shoal to approach potential predators at a distance. Not all members of a shoal are equally likely to initiate such predator inspection visits. Here, we show for the first time that the current hunger state of individual fish strongly influences their predator inspection behaviour, as well as their foraging rate, in the face of predation hazard. When all members of threespine stickleback (Gasterosteus aculeatus) test shoals were in a similar hunger state, they were equally likely to inspect a trout predator model alone and did not differ in the frequency of their inspection visits or foraging rate. However, when individual sticklebacks in a shoal differed in their hunger state, the food-deprived (i.e. hungrier) member of the shoal fed at a higher rate, was significantly more likely to initiate solitary predator inspection visits, and inspected the predator model significantly more often than its less hungry (i.e. well-fed) shoal mates. Individual fish which inspected the predator model more frequently also tended to have higher feeding rates. The results indicate that the hungrier fish in a shoal are more willing to take greater risks to inspect a potential threat at a distance, compared with their well-fed shoal mates, and suggest that they may gain a foraging benefit in doing so. If marked asymmetries in hunger state exist among members of fish shoals, then mutual cooperation during predator inspection visits may be difficult to achieve because well-fed individuals are not as likely to initiate or participate in inspection visits as are hungry individuals.Correspondence to: J.-G.J. Godin     

The sex in short supply for matings varies over small spatial scales in a katydid (Kawanaphilanartee, Orthoptera: Tettigoniidae)

In katydids such as Kawanaphilanartee, a female bias in the operational sex ratio (OSR) results in female competition for mates and male choice of mates. Previous work showed that the excess of sexually active females occurs when food availability is low, in part because less food increases the propensity of females to mate as they forage for the large edible spermatophores produced by males. In this study with K.nartee, a pollen-feeding species, we estimate natural variation in numbers of sexually active males and females by assessing male calling activity and the propensity of females to respond to experimental calling males. We found an excess of sexually active males at a site with many flowers and an excess of sexually active females at a site with few flowers about 900 m away. Between-site differences in gut masses of calling males were consistent with the hypothesis that pollen availability controls OSR. Finally, at a third site where flowers were at first scarce, we found that the initial excess in sexually active females changed to an excess of sexually active males after a clump of grass-trees flowered. The mean gut mass of all sampled males from this site increased after flowering. The large variation in OSR that we document for K. nartee highlights the importance of identifying the appropriate spatial and temporal scales over which OSRs are measured in studies of factors controlling sexual selection. Received: 13 May 1997 / Accepted after revision: 27 October 1997     

Environmental and social factors influence chorusing behaviour in a tropical frog: examining various temporal and spatial scales

Many animals use conspicuous display to attract mates, and there should be selection for displays to occur at times and places that maximise the probability of mating, while minimising energetic costs and predator attraction. To select the best times for display, individuals may use environmental cues, the presence of other individuals, or both, but few studies have examined these sources of variation in display activity. In this study, we examined physical environmental and social factors triggering displays in a tropical, terrestrially breeding frog, Cophixalus ornatus. To measure the influence of physical environmental conditions on calling activity, we recorded temperature, rainfall, moon illumination/visibility, humidity, barometric pressure and intensity of calling activity throughout a breeding season at six locations along a 560-m transect. The intensity of calling varied daily, seasonally, and at a small spatial scale. Variation in calling activity from day to day was large. There was also a strong seasonal trend in calling activity: few males called at the start of the season, activity peaked shortly after the beginning of the season, and then declined linearly from the peak to the end of the season. There was also consistent variation among sites along the transect, which may have been due to variations in frog density at each site, or to consistent microscale variations in physical conditions, or both. After statistically removing consistent local variation among sites, a principal components analysis suggested that a maximum of 35.8% of the variation in calling activity among days was due to factors common to all sites, such as weather, moon illumination, or large-scale social facilitation (e.g. of choruses by other choruses). The remainder of the variation among sites (64.2%) was due to site-specific factors, such as small-scale social facilitation or unmeasured, apparently stochastic effects, such as microenvironmental physical factors that do not vary consistently over sites. Regressions of environmental variables on residual calling activity (after removing consistent effects of site and season), alone or in combination, accounted for very little of the variation in the number of calling males (maximum 10%). Thus, our data, showing strong seasonal effects and consistent variation among sites combined with large amounts of variation in the number of calling males at small spatial scales, suggest that environmental conditions, such as temperature, rainfall, moon illumination and barometric pressure, which act over large spatial scales, may determine the overall environmental envelope within which calling can occur but do not account for most of the variation in the number of calling males on a day-to-day or site-to-site basis. Similarly, variations in the number of calling males at small spatial scales suggest that social facilitation is a relatively unimportant trigger for displays on a large scale in these frogs. On the other hand, our data suggest that social facilitation may have important effects on variation in the number of calling males on a day-to-day and site-to-site basis. We used playback experiments to assess whether the sound of calling could initiate displays. We played either a taped chorus or white noise in areas where few (zero to two) males were calling. The number of calling males increased both during and after the chorus stimulus, whereas there was no increase in calling in response to white noise. These data suggest that examining variation in calling activity at small spatial scales can reveal the sources of variation for the number of calling males, and indicate that, in these frogs, males tend to use the calling of other individuals as a cue to determine when to display. Received: 19 October 1999 / Revised: 30 June 2000 / Accepted: 26 August 2000     

Pelagic Sargassum community change over a 40-year period: temporal and spatial variability

Pelagic forms of the brown algae (Phaeophyceae) Sargassum spp. and their conspicuous rafts are defining characteristics of the Sargasso Sea in the western North Atlantic. Given rising temperatures and acidity in the surface ocean, we hypothesized that macrofauna associated with Sargassum in the Sargasso Sea have changed with respect to species composition, diversity, evenness, and sessile epibiota coverage since studies were conducted 40 years ago. Sargassum communities were sampled along a transect through the Sargasso Sea in 2011 and 2012 and compared to samples collected in the Sargasso Sea, Gulf Stream, and south of the subtropical convergence zone from 1966 to 1975. Mobile macrofauna communities exhibited changes in community structure and declines in diversity and evenness within a 6-month time period (August 2011–February 2012). Equivalent declines in diversity and evenness were recorded in the same region (Sargasso Sea, 25°–29°N) in 1972–1973. Recent community structures were unlike any documented historically, whether compared to sites of the same latitude range within the Sargasso Sea, or the broader historical dataset of sites ranging across the Sargasso Sea, Gulf Stream, and south of the subtropical convergence zone. Recent samples also recorded low coverage by sessile epibionts, both calcifying forms and hydroids. The diversity and species composition of macrofauna communities associated with Sargassum might be inherently unstable. While several biological and oceanographic factors might have contributed to these observations, including a decline in pH, increase in summer temperatures, and changes in the abundance and distribution of Sargassum seaweed in the area, it is not currently possible to attribute direct causal links.     

Individual variation in foraging behavior reveals a trade-off between flexibility and performance of a top predator

There is increasing evidence that behavioral flexibility is associated with the ability to adaptively respond to environmental change. Flexibility can be advantageous in some contexts such as exploiting novel resources, but it may come at a cost of accuracy or performance in ecologically relevant tasks, such as foraging. Such trade-offs may, in part, explain why individuals within a species are not equally flexible. Here, we conducted a reversal learning task and predation experiment on a top fish predator, the Northern pike (Esox lucius), to examine individual variation in flexibility and test the hypothesis that an individual’s behavioral flexibility is negatively related with its foraging performance. Pikes were trained to receive a food reward from either a red or blue cup and then the color of the rewarded cup was reversed. We found that pike improved over time in how quickly they oriented to the rewarded cup, but there was a bias toward the color red. Moreover, there was substantial variation among individuals in their ability to overcome this red bias and switch from an unrewarded red cup to the rewarded blue cup, which we interpret as consistent variation among individuals in behavioral flexibility. Furthermore, individual differences in behavioral flexibility were negatively associated with foraging performance on ecologically relevant stickleback prey. Our data indicate that individuals cannot be both behaviorally flexible and efficient predators, suggesting a trade-off between these two traits.     

Feeding behavior of the sea star Astropecten articulatus (Echinodermata: Asteroidea): an evaluation of energy-efficient foraging in a soft-bottom predator

The energy efficiency of the foraging behavior of Astropecten articulatus (Say) was evaluated in the laboratory. Individuals utilized in the study were collected in the northern Gulf of Mexico from 1990 to 1992. Sea stars presented with equal numbers and weights of low-quality and high-quality prey consistently selected prey of the higher quality. Choice of prey appeared to be mediated by contact chemoreception. Individuals presented with equal weights or equal numbers of different-sized prey models demonstrated a significant preference for smaller prey. Size-selective feeding may be attributable to the ease of manipulation and ingestion of smaller prey, which mazimizes food intake per unit time. In the absence of prey, A. articulatus exhibited a directional pattern of movement. However, as prey were encountered, both the frequency and magnitude of angular deviations in the foraging path increased, resulting in increased foraging in areas of higher prey density. This response to prey availability may increase foraging efficiency by maximizing the rate of prey encounter. Like four other species of the genus Astropecten, A. articulatus exhibited two peaks of activity corresponding with dawn and dusk. Diurnal activities with periods of increased prey availability or periods during which predators are diminished or absent. The sea star A. articulatus exhibits foraging behaviors consistent with the maximization of net energy intake per unit foraging time.     

Influence of human activity and monsoon dynamics on spatial and temporal hydrochemistry in tropical coastal waters (Sanya Bay,South China Sea)

Human activities and oceanic influences like mixing and/or upwelling define the hydrochemical and biological characteristics of coastal regions. In Sanya Bay, northern South China Sea, the balance between these two influences on spatial and temporal scales is poorly understood. The influence from human activities was due to discharge from Sanya River, whereas the most important marine influence was related to seasonal changes in hydrodynamics. Spatial differences (p<0.05) in total nitrogen (TN) and chlorophyll-a (Chl-a) were observed. Seasonal differences were observed for temperature and nutrients. Human activities are the dominant factor influemcing hydrochemistry in the inner bay. This region exhibited the maximum influence of discharge from Saya River as estimated by higher nutrient levels. Oceanic influences like upwelling and mixing caused by monsoons are the dominant factors influencing hydrochemistry in the central and outer bay. Both human activities and oceanic influences play important roles in coastal ecosystems in Sanya Bay. This study has defined these characteristics so that better management policies can be enacted.     

How resources and encounters affect the distribution of foraging activity in a seed-harvesting ant

We examined how the foraging ecology of the seed-harvesting ant Messor andrei depends upon the distribution of resources and the presence of conspecifics. Bait experiments showed that colonies can recruit to high-density patches of seeds. However, at the seasonal scale, natural resource distribution did not affect the distribution of foraging activity. We conducted the study in years of high rainfall and thus seed availability may not have been a limiting factor. Colonies always preferred to forage in areas closer to their nest, which may reduce travel time between the nest and foraging sites. On a day-to-day scale, encounters between neighboring colonies at a site increased the probability that colonies would return to forage at that site; this was true both for natural and experimental encounters. In the summer, this resulted in colonies foraging at the sites of intraspecific encounters on more days than in areas where no encounter had occurred. Encounters between colonies included fighting, and there was little overlap between the foraging areas of neighboring colonies: both results suggest that one function of encounters is to defend foraging space. The high probability of return to the site of an encounter between colonies suggests that encounters may have a second function: to indicate the presence of resources. Received: 28 June 1999 / Received in revised form: 12 October 1999 / Accepted: 16 October 1999     

Population dynamics of the barnacle <Emphasis Type="Italic">Chthamalus montagui</Emphasis> at two spatial and temporal scales in northern Spain

This study describes spatial and temporal patterns of variability in population parameters in the barnacle Chthamalus montagui Southward in three localities of northern Spain and evaluates whether density-dependent settlement may regulate population dynamics. The sampling design considered two spatial scales, localities and sites within localities, and two temporal scales, years and six month intervals. Density, amount of free space, mortality, growth rate and magnitude of settlement (both absolute and per unit of free space) were obtained from photographs of permanent quadrats and from direct counts in the field. The number of settlers in scraped and untouched quadrats was used to estimate the importance of the presence of conspecifics in settlement. Significant variation at the two spatial and temporal scales was found for most parameters. Large spatial and temporal variations in adult mortality rate, density, and settlement were observed. Patterns of mortality were not consistent with differences in density among localities. Differences in settlement among localities were maintained through time. We suggest that magnitude of settlement is regulated by persistent features such as topography or local water circulation. We assume that early post-settlement mortality does not differ among localities. In the absence of differential mortality, settlement determines average population density within localities. Within localities, settlement was independent of density and free space. No consistent evidence was found on preferential settlement at the vicinity of conspecifics. The main conclusion is that density-dependent settlement is not relevant for the regulation of the populations of C. montagui in the northern Spain. Regulation might occur by density-dependent processes within the adult fraction of the population and/or the larval phase before settlement.     

Transient dynamics of pelagic producer-grazer systems in a gradient of nutrients and mixing depths

Phytoplankton-grazer dynamics are often characterized by long transients relative to the length of the growing season. Using a phytoplankton-grazer model parameterized for Daphnia pulex with either flexible or fixed algal carbon:nutrient stoichiometry, we explored how nutrient and light supply (the latter by varying depth of the mixed water column) affect the transient dynamics of the system starting from low densities. The system goes through an initial oscillation across nearly the entire light-nutrient supply space. With flexible (but not with fixed) algal stoichiometry, duration of the initial algal peak, timing and duration of the subsequent grazer peak, and timing of the algal minimum are consistently accelerated by nutrient enrichment but decelerated by light enrichment (decreasing mixing depth) over the range of intermediate to shallow mixing depths. These contrasting effects of nutrient vs. light enrichment are consequences of their opposing influences on food quality (algal nutrient content): algal productivity and food quality are positively related along a nutrient gradient but inversely related along a light gradient. Light enrichment therefore slows down grazer growth relative to algal growth, decelerating oscillatory dynamics; nutrient enrichment has opposite effects. We manipulated nutrient supply and mixing depth in a field enclosure experiment. The experimental results were qualitatively much more consistent with the flexible than with the fixed stoichiometry model. Nutrient enrichment increased Daphnia peak biomass, decreased algal minimum biomass, decreased the seston C:P ratio, and accelerated transient oscillatory dynamics. Light enrichment (decreasing mixing depth) produced the opposite patterns, except that Daphnia peak biomass increased monotonously with light enrichment, too. Thus, while the model predicts the possibility of the "paradox of energy enrichment" (a decrease in grazer biomass with light enrichment) at high light and low nutrient supply, this phenomenon did not occur in our experiment.     

Size-dependent, spatial and temporal genetic variation at a leucine aminopeptidase (LAP) locus among blue mussel (Mytilus galloprovincialis) populations along a salinity gradient

Biochemical genetic variation at a leucine aminopeptidase (LAP) locus is related to salinity variation in several marine bivalve molluscs. This paper details an investigation of the Long Island Sound model of LAP selection (LAP genotype-dependent mortality occurring among newly settled Mytilus edulis mussels) carried out in 1997 among three M. galloprovincialis mussel populations along the salinity gradient of Wellington Harbour, New Zealand. Significant LAP genotypic heterogeneity was observed at the LAP locus between small (<25 mm shell length) and large (>25 mm shell length) M. galloprovincialis from Petone and Eastbourne (the two sites experiencing the greatest salinity variation), whereas genotypic heterogeneity was not significantly different between small and large mussels from Seatoun (the site experiencing the least salinity variation). The Lap ³ allele decreased in frequency and the Lap ⁴ allele increased in frequency at Petone and Eastbourne, whereas the Lap ³ and Lap ⁴ allele frequencies remained effectively constant at Seatoun. Both these findings are consistent with the Long Island Sound model of selection. At all three locations, the Lap ^3,3 genotype decreased in frequency from small to large mussels, whereas the Lap ^3,4 genotype increased in frequency from small to large mussels. All other LAP genotypes occurred at low frequencies (<0.10) at all three locations and showed no evidence of frequency change from small to large-size mussels nor evidence of clinal change among the three locations. These genotype frequency data possibly indicate that the Lap ^3,3 genotype is at a selective disadvantage compared to the Lap ^3,4 genotype at all three locations, and that this selective disadvantage is related to the extent of salinity variation which exists at each location. Further investigation is required before it can be determined if the Long Island Sound model of selection best describes the size-dependent and location-specific changes in LAP allele and genotype frequencies along this salinity gradient. Comparison of the population genetic structure at the LAP locus in 1995 and in 1997 revealed a profound change from heterozygote excesses to heterozygote deficiencies for all three M. galloprovincialis populations. The reason for the change is unknown, but the change indicates that population genetic structure at the LAP locus is highly variable in time, but consistent in space, among these M.␣galloprovincialis populations. Received: 5 February 1998 / Accepted: 27 May 1998     

Vertical distributions of decapod crustacean larvae and pelagic post-larvae over Great Sole Bank (Celtic Sea) in June 1983

Variations in the vertical distributions of pelagic stages of decapod crustaceans are described from five hauls taken over the Great Sole Bank in June 1983. Analyses of the results discriminated between the distributions mainly according to the time of day at which the hauls were taken, suggesting that diel variations were their most significant feature. Substantial ontogenetic variations were observed in the distributions of Processa canaliculata.