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Clamming up: environmental forces diminish the perceptive ability of bivalve prey.

The lethal and nonlethal impacts of predators in marine systems are often mediated via reciprocal detection of waterborne chemical signals between consumers and prey. Local flow environments can enhance or impair the chemoreception ability of consumers, but the effect of hydrodynamics on detection of predation risk by prey has not been investigated. Using clams as our model organism, we investigated two specific questions: (1) Can clams decrease their mortality by responding to predators? (2) Do fluid forces affect the ability of clams to detect approaching predators? Previous research has documented a decrease in clam feeding (pumping) in response to a neighboring predator. We determined the benefits of this behavior to survivorship by placing clams in the field with knobbed whelk or blue crab predators caged nearby and compared mortality between these clams and clams near a cage-only control. Significantly more clams survived in areas containing a caged predator, suggesting that predator-induced alterations in feeding reduce clam mortality in the field. We ascertained the effect of fluid forces on clam perception of predators in a laboratory flume by comparing the feeding (pumping) behavior of clams in response to crabs and whelks in flows of 3 and 11 cm/s. Clams pumped significantly less in the presence of predators, but their reaction to blue crabs diminished in the higher velocity flow, while their response to whelks remained constant in both flows. Thus, clam reactive distance to blue crabs was affected by fluid forces, but hydrodynamic effects on clam perceptive distance was predator specific. After predators were removed, clams exposed to whelks took significantly longer to resume feeding than those exposed to blue crabs. Our results suggest that prey perception of predators can be altered by physical forces. Prey detection of predators is the underlying mechanism for trait-mediated indirect interactions (TMIIs), and recent research has documented the importance of TMIIs to community structure. Since physical forces can influence prey perception, the prevalence of TMIIs in communities may, in part, be related to the sensory ability of prey, physical forces in the environment that impact sensory performance, and the type of predator detected.

Animal Communication↗

Behavioral stochastic resonance: how the noise from a Daphnia swarm enhances individual prey capture by juvenile paddlefish.

Zooplankton emit weak electric fields into the surrounding water that originate from their own muscular activities associated with swimming and feeding. Juvenile paddlefish prey upon single zooplankton by detecting and tracking these weak electric signatures. The passive electric sense in this fish is provided by an elaborate array of electroreceptors, Ampullae of Lorenzini, spread over the surface of an elongated rostrum. We have previously shown that the fish use stochastic resonance to enhance prey capture near the detection threshold of their sensory system. However, stochastic resonance requires an external source of electrical noise in order to function. A swarm of plankton, for example Daphnia, can provide the required noise. We hypothesize that juvenile paddlefish can detect and attack single Daphnia as outliers in the vicinity of the swarm by using noise from the swarm itself. From the power spectral density of the noise plus the weak signal from a single Daphnia, we calculate the signal-to-noise ratio, Fisher information and discriminability at the surface of the paddlefish's rostrum. The results predict a specific attack pattern for the paddlefish that appears to be experimentally testable.

Animals↗

Transient division of labor and behavioral specialization in the ant Formica schaufussi.

Cooperative prey retrieval in the monomorphic ant Formica schaufussi is carried out by workers that perform functionally distinct roles which persist only for the duration of a single retrieval event. A forager (scout) that locates prey too large to retrieve individually organizes cooperative prey transport by recruiting nestmates (recruits) to assist in retrieval. The scout and recruit roles appear to be determined by whether a worker activates recruitment or is recruited from the nest. Scouts organize recruitment and play a key role in maintaining the cohesion of the retrieval group. If a scout that has initiated group transport is experimentally removed, the recruited workers composing the retrieval group typically abandon the prey and cooperative foraging is terminated. In this context, recruits are unable to function as scouts and reorganize group transport. Individuals marked as recruits in one prey retrieval, however, can switch and act as scouts in subsequent retrievals. Because the roles of individuals persist within but not between retrieval events, the specialization involved in cooperative prey retrieval cannot easily be explained as a response associated with age-related or genetic predispositions. This transient division-of-labor, in which individual roles may persist only for the duration of a single group action, represents a novel type of short-term individual specialization.

Animals↗

Context-dependent discrimination and the evolution of mimicry.

Many mimetic organisms have evolved a close resemblance to their models, making it difficult to discriminate between them on the basis of appearance alone. However, if mimics and models differ slightly in their activity patterns, behavior, or use of microhabitats, the exact circumstances under which a signaler is encountered may provide additional clues to its identity. We employ an optimality model of mimetic discrimination in which signal receivers obtain information about the relative risk of encountering mimics and models by observing an external background cue and flexibly adjust their response thresholds. Although such flexibility on the part of signal receivers has been predicted by theory and is supported by empirical evidence in a range of biological settings, little is known about the effects it has on signalers. We show that the presence of external cues that partly reveal signaler identity may benefit models and harm mimics, harm both, or even benefit both, depending on ecological circumstances. Moreover, if mimetic traits are costly to express, or mimics are related to their neighbors, context-dependent discrimination can dramatically alter the outcome of mimetic evolution. We discuss context-dependent discrimination among signal receivers in relation to small-scale synchrony in model and mimic activity patterns.

Adaptation, Physiological↗

Oviposition habitat selection in response to risk of predation in temporary pools: mode of detection and consistency across experimental venue.

Natural selection should favor females that avoid ovipositing where risk of predation is high for their progeny. Despite the large consequences of such oviposition behavior for individual fitness, population dynamics, and community structure, relatively few studies have tested for this behavior. Moreover, these studies have rarely assessed the mode of detection of predators, compared responses in prey species that vary in vulnerability to predators, or tested for the behavior in natural habitats. In an outdoor artificial pool experiment, we tested the oviposition responses of two dipteran species, Culiseta longiareolata (mosquito) and Chironomus riparius (midge), to the hemipteran predator, Notonecta maculata. Both dipteran species have similar life history characteristics, but Culiseta longiareolata larvae are highly vulnerable to predation by Notonecta, while Chironomus riparius larvae are not. As their vulnerabilities would suggest, Culiseta longiareolata, but not Chironomus riparius, strongly avoided ovipositing in pools containing Notonecta. An experiment in natural rock pools assessing oviposition by Culiseta longiareolata in response to Notonecta maculata yielded an oviposition pattern highly consistent with that of the artificial pool experiment. We also demonstrated that the cue for oviposition avoidance by Culiseta longiareolata was a predator-released chemical: Notonecta water (without Notonecta replenishment) repelled oviposition for 8 days. Oviposition avoidance and mode of detection of the predator have important implications for how to assess the true impact of predators and for the use of commercially produced kairomones for mosquito control.

Animals↗

Social transmission of novel foraging behavior in bats: frog calls and their referents.

The fringe-lipped bat, Trachops cirrhosus, uses prey-emitted acoustic cues (frog calls) to assess prey palatability . Previous experiments show that wild T. cirrhosus brought into the laboratory are flexible in their ability to reverse the associations they form between prey cues and prey quality . Here we asked how this flexibility can be achieved in nature. We quantified the rate at which bats learned to associate the calls of a poisonous toad species with palatable prey by placing bats in three groups: (a) social learning, in which a bat inexperienced with the novel association was allowed to observe an experienced bat; (b) social facilitation, in which two inexperienced bats were presented with the experimental task together; and (c) trial-and-error, in which a single inexperienced bat was presented with the experimental task alone. In the social-learning group, bats rapidly acquired the novel association in an average of 5.3 trials. In the social-facilitation and trial-and-error groups, most bats did not approach the call of the poisonous species after 100 trials. Thus, once acquired, novel associations between prey cue and prey quality could spread rapidly through the bat population by cultural transmission. This is the first case to document predator social learning of an acoustic prey cue.

Animals↗

Neuronal organization underlying visually elicited prey orienting in the frog--II. Anatomical studies on the laterality of central projections.

A complete transverse hemisection of the neuraxis just caudal to the optic tectum in the frog, Rana pipiens, results in a failure to orient toward stimuli in one visual hemifield [Kostyk and Grobstein (1986) Neuroscience 21, 41-55]. The extent of the deficit area implies disturbances in the outputs triggered by both tectal lobes. In this paper we report studies aimed at determining more precisely what damage is involved in producing the hemisection deficit, with the broader objective of identifying particular neural structures which may be important in visually elicited orienting. Small lesions at the level of the hemisection which are restricted to the ventromedial white tracts result in an orienting deficit identical to that produced by a complete hemisection. Large lesions which spare the ventromedial white tracts are without significant effect on orienting turns. The finding is consistent with the hypothesis that the hemisection deficit results from interruption of tectal outflow paths. Interestingly, partial damage of the ventromedial white tracts does not result in disconnection of any local tectal region from premotor circuitry but instead systematically alters the turns triggered from all tectal regions. Ventrolateral lesions at the same level do not produce deficits in orienting but do disturb optokinetic behavior. Introduction of horseradish peroxidase into ventromedial lesions produces retrograde labeling in a large number of structures both rostral and caudal of the lesion. Labeling patterns following introduction of horseradish peroxidase into ventrolateral lesions, which do not affect orienting turns, were qualitatively similar but differed quantitatively. The observed patterns of tectal cell labeling make it unlikely that the hemisection deficit can be accounted for in terms of interruption of direct projections deriving from complementary regions of the two tectal lobes. They also indicate that if there exists an uncrossed tectal outflow adequate to trigger orienting turns, it must be by way of an indirect projection. A more general analysis of the labeling patterns suggests that a crossed tectal projection and uncrossed projections from three midbrain tegmental nuclei (the anterodorsal tegmental nucleus, the nucleus profunds lateralis and the nucleus of the medial longitudinal fasciculus) are likely to be involved in triggering orienting turns. The three midbrain tegmental nuclei are of particular interest in that they provide possible anatomical substrates for an indirect uncrossed descending tectal outflow path.

Animals↗

Heavy metals alter the survival, growth, metamorphosis, and antipredatory behavior of Columbia spotted frog (Rana luteiventris) tadpoles.

Amphibian populations appear to be declining around the world. Although there is no single cause, one factor may be pollution from heavy metals. As a result of mining in the Silver Valley of Idaho, heavy metals have been released into habitats containing many species of sensitive organisms, including spotted frogs (Rana luteiventris). While the gross extent of pollution has been well documented, the more subtle behavioral effects of heavy metals such as lead, zinc, and cadmium are less well studied. We tested the effects of heavy metals on the short-term survival (LC50) of spotted frog tadpoles. Compared to single metals, metals presented together were toxic at lower doses. We also raised the tadpoles in outdoor mini-ecosystems containing either a single heavy metal or soil from an EPA Superfund site in the Silver Valley known to be composed of numerous heavy metals. Exposure to Silver Valley soil resulted in delayed metamorphosis. We tested the ability of metal-exposed tadpoles to detect and respond to chemical cues emanating from predacious rainbow trout. We found that high levels of Silver Valley soil, medium levels of zinc, and medium and high levels of lead resulted in a decreased fright response. Low levels of cadmium, zinc, and lead did not cause a significant effect, but low levels of soil did result in a decreased fright response. Heavy metals may alter interactions between tadpoles and their predators.

Animals↗

alpha-melanocyte-stimulating hormone and habituation of prey-catching behavior in the Texas toad, Bufo speciosus.

We investigated dose-dependent effects of alpha-melanocyte-stimulating hormone (alpha-MSH) on habituation in the Texas toad, Bufo speciosus. Additionally, we determined changes in plasma and brain levels of alpha-MSH following peripheral administration of the peptide or following exposure to an ether stressor. The ability of alpha-MSH to facilitate acquisition of habituation was dose dependent. Plasma alpha-MSH concentrations were elevated within 5 min of dorsal lymph sac injection and remained elevated up to 600% over controls after 30 min. Administration of 50 microgram alpha-MSH had no effect on plasma corticosterone levels. Radiolabeled alpha-MSH was detected in cerebrospinal fluid microdialysates within minutes of peripheral injection. Concentrations of alpha-MSH in the telencephalon and preoptic area were significantly lowered after ether exposure, whereas levels in the optic tectum, thalamus/hypothalamus, brainstem, and plasma were unchanged. We conclude that alpha-MSH administered peripherally facilitates habituation in a dose-dependent fashion. Our results confirm that the effects of alpha-MSH are independent of corticosterone secretion. The peptide is cleared rapidly into the bloodstream and enters the cerebrospinal fluid after dorsal lymph sac injection. Neuronal alpha-MSH may help toads gather information about their environment when exposed to certain stressors.

Animals↗

Parasite infection attenuates nonopioid mediated predator-induced analgesia in mice.

Parasites have been shown to have a broad range of effects on host behavior, including alterations of host responses to predators. Response to the threat of predation consist of a number of defensive behaviors, including a reduction in pain sensitivity and the induction of analgesia. The present study examined the relationships between subclinical (i.e., nonpathological) infection with the naturally occurring, enteric, sporozoan (coccidian) parasite, Eimeria vermiformis, predator exposure, and nociceptive responses in male mice. Brief (30 s) exposure of nonparasitized mice to a predator (a cat) induced marked, relatively short-lived analgesia that was insensitive to naloxone and blocked by the serotonin-1A (5-HT1A) agonist, 8-OH-DPAT. In contrast, mice acutely infected for 6 days with E. vermiformis, failed to show a predator-induced analgesia. The parasitized mice did display a naloxone-sensitive hypoalgesia or analgesia. However, restraint-stressed mice, which displayed a naloxone-sensitive hypoalgesia similar in amplitude to that of the infected mice, still exhibited a nonopioid mediated, predator-induced analgesia. These observations indicate that parasite infection attenuates 5-HT1A-sensitive predator-induced analgesia and likely reduces the accompanying fear and anxiety related anticipatory defense reactions of the host to the predator.

Animals↗

Talking helps: evolving communicating agents for the predator-prey pursuit problem.

We analyze a general model of multi-agent communication in which all agents communicate simultaneously to a message board. A genetic algorithm is used to evolve multi-agent languages for the predator agents in a version of the predator-prey pursuit problem. We show that the resulting behavior of the communicating multi-agent system is equivalent to that of a Mealy finite state machine whose states are determined by the agents' usage of the evolved language. Simulations show that the evolution of a communication language improves the performance of the predators. Increasing the language size (and thus increasing the number of possible states in the Mealy machine) improves the performance even further. Furthermore, the evolved communicating predators perform significantly better than all previous work on similar prey. We introduce a method for incrementally increasing the language size, which results in an effective coarse-to-fine search that significantly reduces the evolution time required to find a solution. We present some observations on the effects of language size, experimental setup, and prey difficulty on the evolved Mealy machines. In particular, we observe that the start state is often revisited, and incrementally increasing the language size results in smaller Mealy machines. Finally, a simple rule is derived that provides a pessimistic estimate on the minimum language size that should be used for any multi-agent problem.

Algorithms↗

A complex, cross-taxon, chemical releaser of antipredator behavior in amphibians.

Prey species show diverse antipredator responses to chemical cues signaling predation threat. Among terrestrial vertebrates, the red-backed salamander, Plethodon cinereus, is an important species in the study of these chemical defenses. During the day and early evening, this species avoids rinses from garter snakes, Thamnophis sirtalis, independent of snake diet, but late at night. avoids only those rinses from garter snakes that have recently eaten P. cinereus. We tested whether the selective, late-night response requires the ingestion or injury of salamanders. In three experiments, we tested P. cinereus for their responses to separate or combined rinses from salamanders (undisturbed, distressed, and injured P. cinereus) and snakes (unfed, earthworm fed, and salamander-fed T. sirtalis). When paired against a water control, only rinses from salamander-fed snakes were avoided. When salamander treatments (undisturbed or distressed) were combined with the snake treatments (unfed or earthworm-fed) and tested against a water control, the combinations elicited avoidance. When selected treatments were paired against the standard rinse from salamander-fed snakes, only the combined rinses from salamanders and snakes nullified the avoidance response to the standard rinse. These data reveal a prey defense mechanism involving chemical elements fromboth the predatorand prey that does not require injury or ingestion of the prey in the formation of the cue.

Animals↗

Further research at the Oldowan site of Ain Hanech, North-eastern Algeria.

Further investigations were carried out at Ain Hanech, Algeria in 1998 and 1999 to explore its potential for investigating early hominid behavioral patterns and adaptation. Research concentrated on the stratigraphy and dating, identifying new archaeological deposits, and excavating the Ain Hanech and El-Kherba localities. To enhance the chronological control within a biostratigraphic framework, the Ain Boucherit fossil-bearing stratum, yielding a Plio-Pleistocene fauna, is correlated with the regional stratigraphy. In the stratigraphic sequence, the Ain Boucherit stratum, located 13m below the Ain Hanech Oldowan occurrences, is found in Unit Q of the Ain Hanech Formation. Unit Q shows a paleomagnetically reversed polarity, which may be correlated with an age earlier than the Olduvai normal subchron (1.95-1.77Ma). Based on test trenches and stratigraphic analyses, additional Oldowan deposits A, B, and C are identified at Ain Hanech. All three deposits and the El-Kherba site contain Mode I technology artefacts associated with an Early Pleistocene fauna. El-Kherba is stratigraphically equivalent to Ain Hanech. These two archaeological sites are estimated to be dated to about 1.8Ma.

Adaptation, Physiological↗

Is sleep's 'supreme mystery' unraveling? An evolutionary analysis of sleep encounters no mystery; nor does life's earliest sleep, recently discovered in jellyfish.

Biotelemetry has revealed daily 15-h behavioral sleep periods in a cubomedusan jellyfish, Chironex fleckeri. Its sleep is expected to be phylogenetically most primitive, since jellyfish possess only two germ layers. They belong to the phylum Cnidaria, the 'simplest' multicellular organisms with an organized nervous system. Cubomedusae have a complex visual system with 24 eyes of four different types, each type specialized for a different task. Input to these eyes during visually guided fast-swimming predation requires enormous amounts of neural processing, possibly nearly saturating the capacity of their comparatively simple nervous system. These heavy neural demands may account for the need for fifteen hours of sleep. C. fleckeri is the only animal known for which sleep may be either present or absent, dependent on lifestyle. Limited knowledge of behavior of some other cubomedusae suggests that they also possess this faculty. The finding of sleep in C. fleckeri supports current proposals of sleep's origin and basic function. Evolutionary analyses link sleep to a conflict produced by excessive processing demands on multifunctional neural circuitry for detailed focal vision by complex lensed eyes. The conflict arises between the enormous demands of complex visual analysis and needs for split-second control of actions, on the one hand, and non-urgent processing of memories of ongoing and stored events, on the other. Conflict is resolved by deferring the non-urgent processing to periods of sleep. Without sleep, selection would favor the evolution of circuitry 'dedicated' to single or but few tasks, with corresponding lesser efficiency. Had complex lensed eyes of medusae originated as a consequence of selection for increased mating success of males pursuing females, it could have occurred before the evolution of fast-swimming bilateral (three-germ-layered) prey. But if it was a consequence of selection for increased prey-hunting success, the origin of such eyes probably awaited the coexistence of bilateral prey.

Animals↗

Phoretic nest parasites use sexual deception to obtain transport to their host's nest.

Cooperative behaviors are common among social insects such as bees, wasps, ants, and termites, but they have not been reported from insect species that use aggressive mimicry to manipulate and exploit prey or hosts. Here we show that larval aggregations of the blister beetle Meloe franciscanus, which parasitize nests of the solitary bee Habropoda pallida, cooperate to exploit the sexual communication system of their hosts by producing a chemical cue that mimics the sex pheromone of the female bee. Male bees are lured to larval aggregations, and upon contact (pseudocopulation) the beetle larvae attach to the male bees. The larvae transfer to female bees during mating and subsequently are transported to the nests of their hosts. To mimic the chemical and visual signals of female bees effectively, the parasite larvae must cooperate, emphasizing the adaptive value of cooperation between larvae. The aggressive chemical mimicry by the beetle larvae and their subsequent transport to their hosts' nests by the hosts themselves provide an efficient solution to the problem of locating a critical but scarce resource in a harsh environment.

Adaptation, Biological↗

Prey catching in archer fish: the role of posture and morphology in aiming behavior.

Archer fishes (Toxotidae) bring down aerial prey with a squirt of water from their mouth. How they aim, however, is not clear. Measurements of the angle between the squirt and the fish's body axis (the mouth angle) already yielded agreement that pointing the body plays a part in aiming, and it has been suggested that more accurate aiming would occur by means of lips and tongue. In a previous study, however, we found that the mouth angle is an individual property, with a much smaller range than reported by others. Moreover, we found that archer fishes do not possess movable lips, and that their tongue seems too short to direct the squirt. Presently, we found that the tongue cannot even reach the rostral part of the mouth cavity, that affects the squirt's direction, and that the mouth angle is an individual property remaining constant, even across a wide range of target distances. These findings suggest that aiming the body is the only way of aiming the squirt. We also found that during aiming, the body rotates around the eyeballs which remain fixed in space. The position of the eyeballs in the head (eye angle) at the moment of squirting was also found to be an individual property. However, the eye angle is less constant than the mouth angle. The systematic variance of the eye angle probably is related to the refraction effect.

Accommodation, Ocular↗

Post-fledging survival of northern goshawks: the importance of prey abundance, weather, and dispersal.

Effective wildlife conservation strategies require an understanding of how fluctuating environmental conditions affect sensitive life stages. As part of a long-term study, we examined post-fledging and post-independence survival of 89 radio-marked juvenile Northern Goshawks (Accipiter gentilis) produced from 48 nests in northern Arizona, USA, during 1998-2001. Information-theoretic methods were used to examine within- and among-year variation in survival relative to environmental (prey abundance, weather), territory (hatching date, brood size), and individual (gender, body mass) sources of variation. The results support age- and cohort-specific differences in survival that were best explained by behaviors occurring at distinct stages of juvenile development, annual changes in the density of primary bird and mammal prey species, and gender-related differences in body mass. Survival between fledging and independence increased linearly with age and varied among annual cohorts of radio-marked juveniles from 0.81 (95% CI = 0.60-0.93) to 1.00 (95% CI = 0.95-1.00) in association with annual differences in prey density; the slope coefficient for the additive effect of prey density on survival was 1.12 (95% CI = 0.06-2.19). Survival declined to 0.71 (95% CI = 0.60-0.93) shortly after juveniles initiated dispersal (weeks 8-12 post-fledging) and moved to more open habitats at lower elevations. Survival was not closely associated with weather or territory-level parameters. A comparison of the predictions of environmental-, territory-, and individual-based models of survival demonstrated that food availability was the primary factor limiting juvenile survival. This finding indicates that forest management prescriptions designed to support abundant prey populations while providing forest structural conditions that allow goshawks to access their prey within breeding areas should benefit juvenile survival.

Animals↗

The buccal buckle: the functional morphology of venom spitting in cobras.

Multiple radiations of Asiatic and African cobras have independently evolved the ability to expel their venom as a pressurized horizontal stream, a behavior commonly referred to as spitting. Though the unique fang morphology of spitting cobras is well known, the functional bases of venom spitting have received little attention. The combined results of gross and microscopic morphology, high-speed digital videography, experimental manipulations of anesthetized cobras and electromyography reveal a two-part mechanism for spitting venom. Contraction of the M. protractor pterygoideus (PP) causes displacement and deformation of the palato-maxillary arch and fang sheath; ultimately this displacement removes soft tissue barriers to venom flow that are normally present within the fang sheath. The M. adductor mandibulae externus superficialis (AMES) is activated simultaneously with the PP; the AMES increases venom pressure within the venom gland, propelling a stream of venom through the venom duct and out the fang. The displacements of the palato-maxillary arch, which form the first part of the spitting mechanism, are very similar to the motions of these bones during prey ingestion (the pterygoid walk), suggesting that venom spitting may have evolved from a specialization of prey ingestion, rather than prey capture.

Animals↗