Gregarious behaviour of evasive prey

Gregarious behavior of potential prey was explained by Hamilton (1971) on the basis of risk-sharing: The probability of being picked up by a predator is small when one makes part of a large aggregate of prey. This argument holds only if the predator chooses its victims at random. It is not the case for herds of evasive prey in the open, where prey's gregarious behavior, favorable for the fast group members, makes it easier for the predator to home in on the slowest ones. We show conditions under which gregarious behavior of the relatively fast prey individuals leaves slowest prey with no other choice but to join the group. Failing to do so would signal their vulnerability, making them a preferred target for the predator. Analysis of an n+1 player game of a predator and n unequal prey individuals clarifies conditions for fully gregarious, partially gregarious, or solitary behavior of the prey.     

Combined harvesting of a stage structured prey–predator model incorporating cannibalism in competitive environment

In this paper, we propose a prey–predator system with stage structure for predator. The proposed system incorporates cannibalism for predator populations in a competitive environment. The combined fishing effort is considered as control used to harvest the populations. The steady states of the system are determined and the dynamical behavior of the system is discussed. Local stability of the system is analyzed and sufficient conditions are derived for the global stability of the system at the positive equilibrium point. The existence of the Hopf bifurcation phenomenon is examined at the positive equilibrium point of the proposed system. We consider harvesting effort as a control parameter and subsequently, characterize the optimal control parameter in order to formulate the optimal control problem under the dynamic framework towards optimal utilization of the resource. Moreover, the optimal system is solved numerically to investigate the sustainability of the ecosystem using an iterative method with a Runge–Kutta fourth-order scheme. Simulation results show that the optimal control scheme can achieve sustainable ecosystem. Results are analyzed with the help of graphical illustrations.     

Predation and the evolution of prey behavior: an experiment with tree hole mosquitoes

We tested for facultative changes in behavior of an aquaticinsect in response to cues from predation and for evolutionof prey behavior in response to experimental predation regimes.Larvae of the tree hole mosquito Aedes triseriatus reducedfiltering, browsing, and time below the surface in responseto water that had held a feeding larva of the predator Toxorhynchitesrutilus. We subjected experimental A. triseriatus populationsto culling of 50% of the larval population, either by T. rutiluspredation or by random removal. After two generations of laboratoryculling, behavior of the two treatment groups diverged. Aedestriseriatus in control-culled lines retained their facultativeshift from filtering to resting, but tended to lose the response of reduced browsing below the surface in water that had helda feeding predator. Predator-culled lines lost their facultativeresponse of reduced filtering in water that had held a feedingpredator and evolved toward more time resting and less timefiltering in both water that had held a feeding predator andwater that had held only A. triseriatus. Predator-culled linesretained their facultative response of reduced browsing belowthe surface in water that had held a feeding predator. Twofield populations and their reciprocal hybrids responded similarlyto cues from predation and did not differ in their evolutionaryresponse to experimental culling. We conclude that consistentpresence or absence of predation can select rapidly for divergencein prey behavior, including facultative behavioral responsesto predators.     

Behavior of the rhabdocoel flatworm Mesostoma ehrenbergii in prey capture and feeding

The prey-capture and feeding behavior of the rhabdocoel flatworm Mesostoma ehrenbergii (Focke, 1836) was analyzed using a variety of live and dead prey, including Daphnia, mosquito larvae, and tubifex annelids. Prey-capture behavior was broken down into its individual components. Mesostoma could accommodate to and change its behavior depending on the size and type of prey. Mechanical rather than chemical cues were effective in inducing prey-capture behavior. No evidence for a special chemical paralysis as suggested by other workers was found. The apparent paralysis observed in cladocera such as Daphnia and mosquito larvae was, in part a behavioral response of the prey in ‘playing possum’ and also in part due to immobilization of the prey by the flatworm with mucous threads. This revised version was published online in July 2006 with corrections to the Cover Date.     

The role of size-specific predation in the evolution and diversification of prey life histories

Some of the best empirical examples of life-history evolution involve responses to predation. Nevertheless, most life-history theory dealing with responses to predation has not been formulated within an explicit dynamic food-web context. In particular, most previous theory does not explicitly consider the coupled population dynamics of the focal species and its predators and resources. Here we present a model of life-history evolution that explores the evolutionary consequences of size-specific predation on small individuals when there is a trade-off between growth and reproduction. The model explicitly describes the population dynamics of a predator, the prey of interest, and its resource. The selective forces that cause life-history evolution in the prey species emerge from the ecological interactions embodied by this model and can involve important elements of frequency dependence. Our results demonstrate that the strength of the coupling between predator and prey in the community determines many aspects of life-history evolution. If the coupling is weak (as is implicitly assumed in many previous models), differences in resource productivity have no effect on the nature of life-history evolution. A single life-history strategy is favored that minimizes the equilibrium resource density (if possible). If the coupling is strong, then higher resource productivities select for faster growth into the predation size refuge. Moreover, under strong coupling it is also possible for natural selection to favor an evolutionary diversification of life histories, possibly resulting in two coexisting species with divergent life-history strategies.     

Behavioral strategies of aphid hyperparasitoids to escape aggression by honeydew-collecting ants

We analyzed the behavioral interactions between two species of honeydew-collecting ants (Lasius niger, Myrmica laevinodis) and foraging females of four species of aphid hyperparasitoids (Aphidencyrtus aphidivorus, Dendrocerus carpenteri, Pachyneuron aphidis, Asaphes vulgaris) usingAphis fabae ssp.cirsiiacanthoidis andLysiphlebus cardui on thistles as aphid and primary parasitoid, respectively. The observed interaction patterns and foraging parameters varied within hyperparasitoid species and revealed different strategies based upon behavioral and morphological constraints.D. carpenteri generally tried to avoid ant encounters. This avoidance strategy was successful in interactions withM. laevinodis but failed when encountering the more aggressiveL. niger, which caused about 26% adult mortality. In contrast,A. aphidivorus, P. aphidis, andA. vulgaris possess jumping ability and were hardly exposed to mortality risks. The escape reaction jump off was used as soon as ants made physical contact with foraging females. While the flight strategy ofP. aphidis is connected with cryptic movement patterns without avoidance behavior,A. aphidivorus first avoided ants and jumped off only as a last resort. Similar patterns, but less expressive, are displayed byA. vulgaris. We suggest that these different strategies are responsible for different foraging success in ant-attended resources in field.     

A game theoretical approach to the evolution of structured communication codes

Structured meaning-signal mappings, i.e., mappings that preserve neighborhood relationships by associating similar signals with similar meanings, are advantageous in an environment where signals are corrupted by noise and sub-optimal meaning inferences are rewarded as well. The evolution of these mappings, however, cannot be explained within a traditional language evolutionary game scenario in which individuals meet randomly because the evolutionary dynamics is trapped in local maxima that do not reflect the structure of the meaning and signal spaces. Here we use a simple game theoretical model to show analytically that when individuals adopting the same communication code meet more frequently than individuals using different codes—a result of the spatial organization of the population—then advantageous linguistic innovations can spread and take over the population. In addition, we report results of simulations in which an individual can communicate only with its K nearest neighbors and show that the probability that the lineage of a mutant that uses a more efficient communication code becomes fixed decreases exponentially with increasing K. These findings support the mother tongue hypothesis that human language evolved as a communication system used among kin, especially between mothers and offspring.     

Predator/prey interactions: A link between the individual level and both higher and lower level effects of toxicants in aquatic ecosystems

Behavior can demonstrate linkages ofcontaminant effects at different levels oforganization from the biochemical/cellular to theorganism, population, and community levels.Mummichogs, Fundulus heteroclitus, from acontaminated area were previously found to havereduced condition, growth, and longevity, comparedwith conspecifics from clean sites. Thispopulation-level observation may be due to theirimpaired predator/prey behavior, which is associatedwith altered levels of serotonin in their brains. Theyare slow, less able to capture live food, and eat muchnon-nutritious detritus, which may contribute to theirdecreased growth and condition. They are also lessable to avoid being captured by blue crabs, which canalso contribute to their reduced life span andsize-structure. Grass shrimp, Palaemonetespugio, a major prey species, are not behaviorallyimpaired at this polluted site, but are more numerousand larger in size than at the reference site,probably due to reduced mummichog predation.     

A rare predator exploits prey escape behavior: the role of tail-fanning and plumage contrast in foraging of the painted redstart (Myioborus pictus)

Escape response, triggered by an approaching predator, is acommon antipredatory adaptation of arthropods against insectivores.The painted redstart, Myioborus pictus, represents insectivorousbirds that exploit such antipredatory behaviors by flushing,chasing, and preying upon flushed arthropods. In field experimentsI showed that redstarts evoke jump and flight in prey by spreadingwings and tail: this display increased frequency of aerial chasesby redstarts. Artificial models with spread tails also elicitedescape responses more often than models with closed tails and wings.The white patches on black wings and tails additionally help:the frequency of chases decreased when the white patches werecovered with black dye. Black models also tended to elicit escaperesponse less often than black-and-white models did, at leastin some situations. Hence, the prey's ability to detect birdsand to flee could cause the evolution of predators specializedat using conspicuous behavior and contrast in body colorationto elicit and exploit such antipredatory responses. Redstartsconstitute only a small proportion of the predatory guild, andtheir adaptations to exploit the prey's behavior illustratethe theoretically modeled "rare enemy" effect present in multispeciespredator-prey systems. This is the first experimental studyof morphological and behavioral adaptations of a rare predatorthat both elicits and exploits antipredator escape behaviorof its prey against more common predators. Hence, the studydocuments a behavior that could be evolutionarily explainedonly if indirect interactions in predator-prey communities aretaken into account.     

The influence of distance to burrow on flight initiation distance in the woodchuck, Marmota monax

We used woodchucks (Marmota monax) to test predictions of acost-benefit model of antipredator behavior that flight initiationdistance would increase with distance to refuge and with predatorapproach velocity. We also examined the effects of distanceto refuge and predator approach velocity on escape velocityand on both temporal and spatial margin of safety (expectedtime and distance between predator and burrow at the time ofthe woodchuck's arrival). The observer, assumed to be perceivedas a potential predator, approached juvenile woodchucks fromthe direction opposite to the burrow at a slow (1.24 m/s) orfast (1.79 m/s) walking pace. When the woodchuck started toflee, the observer recorded the woodchuck's distance from theobserver and from its burrow, the time spent running, and whetherthe woodchuck stopped before reaching its burrow. Flight initiationdistance increased consistendy with distance to the burrow overthe entire observed range (0–25 m) but was not significantlyaffected by observer approach velocity. Escape velocity wasnot significantly influenced by the observer approach velocityand was approximately constant over the range of 2–25m, but was slower for woodchucks less than 2 m from their burrows.Both temporal and spatial margins of safety increased with distancefrom the burrow. The temporal margin of safety increased withdistance from the burrow more rapidly for slow than for fastobserver approach velocity. Woodchucks fleeing from greaterthan 2 m usually stopped near the burrow before entering, butthose from closer distances usually entered directly. Theseresults support the assumption that antipredator behavior issensitive to the costs and benefits of alternative escape decisions.     

Simulation of gamete behaviors and the evolution of anisogamy: reproductive strategies of marine green algae

In marine green algae, isogamous or slightly anisogamous species are taxonomically widespread. They produce positively phototactic gametes in both sexes. We developed a new numerical simulator of gamete behavior using C++ and pseudo-parallelization methods to elucidate potential advantages of phototaxis. Input parameters were set based on experimental data. Each gamete swimming in a virtual rectangular test tank was tracked and the distances between the centers of nearby male and female were measured at each step to detect collisions. Our results shed light on the roles of gamete behavior and the mechanisms of the evolution of anisogamy and more derived forms of sexual dimorphism. We demonstrated that not only gametes with positive phototaxis were favored over those without, particularly in shallow water. This was because they could search for potential mates on the 2-D water surface rather than randomly in three dimensions. Also, phototactic behavior clarified the difference between isogamy and slight anisogamy. Isogamous species produced more zygotes than slightly anisogamous ones only under the phototactic conditions. Our results suggested that sperm limitation might be easily resolved particularly in the slightly anisogamous species. Some more markedly anisogamous species produce the smaller male gametes without any phototactic devices and the larger positively phototactic female gametes. In such species, female gametes attract their partners using a sexual pheromone. This pheromonal attraction system might have played a key role in the evolution of anisogamy, because it could enable markedly anisogamous species achieve 2-D search efficiencies on the water surface. The mating systems appear to be tightly tuned o the environmental conditions of their habitats.     

Democracy in animals: the evolution of shared group decisions

A 'consensus decision' is when the members of a group choose, collectively, between mutually exclusive actions. In humans, consensus decisions are often made democratically or in an 'equally shared' manner, i.e. all group members contribute to the decision. Biologists are only now realizing that shared consensus decisions also occur in social animals (other than eusocial insects). Sharing of decisions is, in principle, more profitable for groups than accepting the 'unshared' decision of a single dominant member. However, this is not true for all individual group members, posing a question as to how shared decision making could evolve. Here, we use a game theory model to show that sharing of decisions can evolve under a wide range of circumstances but especially in the following ones: when groups are heterogeneous in composition; when alternative decision outcomes differ in potential costs and these costs are large; when grouping benefits are marginal; or when groups are close to, or above, optimal size. Since these conditions are common in nature, it is easy to see how mechanisms for shared decision making could have arisen in a wide range of species, including early human ancestors.     

The continuous public goods game and the evolution of cooperative sex ratios

Some animals, such as Melittobia wasps and surface-living mites, have extremely female-biased sex ratios that cannot be explained by the existing local mate competition (LMC) theories. The restricted production of sons may entail cooperation among mothers, enabling the production of more daughters and avoiding severe competition among sons for insemination access. These unusual examples are characterized by the long-term cohabitation of egg-layers (foundresses) on resource patches and possible contact with oviposited eggs. By applying the logic of mutual policing, we develop a novel game theoretical model for the evolution of cooperation in sex-ratio traits. This is the first attempt to model the evolution of sex ratios based on iterated games. We assumed that foundresses have two abilities to enable mutual policing: they can monitor the sex ratio in the resource patch, and they can punish defectors that produce an overabundance of males. Numerical analysis and evolutionary simulations demonstrate that cooperative low sex ratios can evolve when the number of foundresses per patch is sufficiently small. Our model predicts a slight, but steady increase in oviposition sex ratios with an increase in the number of foundresses, which mimics the phenomenon observed in several animals with extremely female-biased sex ratios. We also discuss the relationship between our model and other models of sex-ratio evolution.     

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.     

Role of prey and intraspecific density dependence on the population growth of an avian top predator

Exploring predator–prey systems in diverse ecosystems increases our knowledge about ecological processes. Predator population growth may be positive when conspecific density is low but predators also need areas with prey availability, associated with competition, which increases the risk of suffering losses but stabilises populations. We studied relationships between European rabbits Oryctolagus cuniculus (prey) and adult eagle owls Bubo bubo (predators) in south-western Europe. We assessed models explaining the predator population growth and stability. We estimated the abundance of rabbits and adult eagle owls during three years in eight localities of central-southern Spain. We explored models including rabbit and adult eagle owl abundance, accounting for yearly variations and including the locality as a random variable. We found that population growth of adult eagle owls was positive in situations with low conspecific abundance and tended to be negative but approaching equilibrium in situations of higher conspecific abundance. Population growth was also positively related to previous summer rabbit density when taking into account eagle owl conspecific abundance, possibly indicating that rabbits may support recruitment. Furthermore, abundance stability of adult eagle owls was positively related to previous winter–spring rabbit density, which could suggest predator population stabilisation through quick territory occupation in high-quality areas. These results exemplify the trade-off between prey availability and abundance of adult predators related to population growth and abundance stability in the eagle owl–rabbit system in south-western Europe. Despite rabbits have greatly declined during the last decades and eagle owls locally specialise on them, eagle owls currently have a favourable conservation status. As eagle owls are the only nocturnal raptor with such dependence on rabbits, this could point out that predators may overcome prey decreases in areas with favourable climate and prey in the absence of superior competitors with similar foraging mode.     

On the impossibility of coexistence of infinitely many strategies

We investigate the possibility of coexistence of pure, inherited strategies belonging to a large set of potential strategies. We prove that under biologically relevant conditions every model allowing for coexistence of infinitely many strategies is structurally unstable. In particular, this is the case when the "interaction operator" which determines how the growth rate of a strategy depends on the strategy distribution of the population is compact. The interaction operator is not assumed to be linear. We investigate a Lotka-Volterra competition model with a linear interaction operator of convolution type separately because the convolution operator is not compact. For this model, we exclude the possibility of robust coexistence supported on the whole real line, or even on a set containing a limit point. Moreover, we exclude coexistence of an infinite set of equidistant strategies when the total population size is finite. On the other hand, for infinite populations it is possible to have robust coexistence in this case. These results are in line with the ecological concept of "limiting similarity" of coexisting species. We conclude that the mathematical structure of the ecological coexistence problem itself dictates the discreteness of the species.     

Indirect food web interactions affect predation of Tengmalm's Owls Aegolius funereus nests by Pine Martens Martes martes according to the alternative prey hypothesis

Although population cycles of rodents are geographically widespread and occur in a number of rodent species, higher‐order food web interactions mediated by predator–rodent dynamics have primarily been described from boreal and arctic biomes. During periods of low rodent abundance, predators may switch to alternative prey, which may affect other predators directly or indirectly. Using a long‐term dataset, we assessed the frequency of Pine Marten Martes martes predation on the nests of Tengmalm's Owl Aegolius funereus during periods of fluctuating rodent abundance in Central Europe. The number of nests predated by Pine Martens was positively correlated with the annual number of nests available. The probability of predation by Pine Martens on Tengmalm's Owl nests decreased with increasing spring abundance index of Apodemus mice, but was not related to the abundance index of Myodes and Microtus voles, pooled rodent abundance or age of the nestbox. Additionally, we found no relationship between the breeding frequency (i.e. the number of nesting attempts per nestboxes available) and an abundance index of Microtus and Myodes voles, Apodemus mice or overall rodent abundance. Our results demonstrate, for the first time in a temperate area, that during periods of low Apodemus mouse abundance, the switching response of an opportunistic mammalian predator can lead to indirect food web interactions through an increase in nest predation on a sympatric avian predator.     

On the evolution of arginases and related enzymes

Sequence analysis of the Agnase/agmatine ureohydrolase family, important enzymes in arginine/agmatine metabolism and the urea cycle, reveals the similarity of arginases to formiminoglutamate hydrolase (hutG) in Klebsiella aerogenes and to a previously unidentified open reading frame adjacent to the HMf locus of the archaebacterium Methanothermus fervidus. The gene structure and distribution of these homologous proteins across primary kingdoms suggest that this family is another example of a primordial enzyme possibly present in the universal common ancestor and that can be used as phylogenetic marker. Correspondence to: C.A. Ouzounis