Effects of three phamaceuticals on the responses of tadpoles to predator alarm cues

The aim of this study is to measure the effects of two selective serotonin reuptake inhibitors, fluoxetine and sertraline, and one selective serotonin and norepinephrine reuptake inhibitor, venlafaxine, on the swimming and lateralization behaviour of Sclerophrys arabica tadpoles exposed to predator alarm cues. Tadpoles were exposed to the three pharmaceuticals either at 0.5 µg/L or 2.0 µg/L, either individually or as a mixture for 14 days. Control tadpoles and those exposed to high concentration of drugs individually or as a mixture, and those exposed to the low-concertation mixture reduced their swimming speed. Tadpoles exposed to 0.5 µg/L fluoxetine or venlafaxine did not respond to the alarm cues. Tadpoles exposed to the mixture had similar responses to that of the control. Results indicate that these drugs have an additive mode of action. Tadpoles exposed to the low-concentration mixture increased lateralization of movement.     

Sex and life-history stage alter herbivore responses to a chemically defended red alga

Intraspecific variation in resistance to herbivory among genders and life-history phases of primary producers can significantly alter the ecological and evolutionary consequences of plant-herbivore interactions. Seaweeds (macroalgae) with complex life histories have multiple distinct phases with associated variation in traits that can potentially lead to differences in resistance to consumers and provide a unique system in which to simultaneously test the effects of sex and life-history stage on herbivory. We tested the susceptibility to grazing of the three life-history stages and separate sexes of the chemically defended red alga Asparagopsis armata against the sea hare Aplysia parvula, and we related this to the plant quality traits of different stages and genders. Differences in nutrient content and halogenated secondary metabolites between life-history phases were highly sex dependent. Male gametophytes had a low concentration of secondary metabolites and the highest nutrient content. The highest secondary metabolite content was found within the female gametophyte, in the wall of the reproductive structures (cystocarps) that contain the microscopic carposporophyte phase. Feeding choices by A. parvula were consistent with differences in algal quality and defense and resulted in the haploid male gametophytes being the most preferred food type. The diploid carposporophyte found inside the chemically rich cystocarps was the least consumed life-history stage. Selective herbivory of male gametophytes by A. parvula is consistent with an observed shift in gametophyte sex ratio in the field from unity at the beginning of the reproductive season to female bias at the end. The variation in susceptibility to herbivory found between sex and life-history stages of A. armata represents the first example of sex-biased consumption in seaweeds and may contribute to the maintenance of complex life histories such as those found in red algae.     

Individual- and population-level responses of a keystone predator to geographic variation in prey

Investigating how food supply regulates the behavior and population structure of predators remains a central focus of population and community ecology. These responses will determine the strength of bottom-up processes through the food web, which can potentially lead to coupled top-down regulation of local communities. However, characterizing the bottom-up effects of prey is difficult in the case of generalist predators and particularly with predators that have large dispersal scales, attributes that characterize most marine top predators. Here we use long-term data on mussel, barnacle, limpet, and other adult prey abundance and recruitment at sites spread over 970 km to investigate individual- and population-level responses of the keystone intertidal sunstar Heliaster helianthus on the coast of Chile. Our results show that this generalist predator responds to changes in the supply of an apparently preferred prey, the competitively dominant mussel Perumytilus purpuratus. Individual-level parameters (diet composition, per capita prey consumption, predator size) positively responded to increased mussel abundance and recruitment, whereas population-level parameters (density, biomass, size structure) did not respond to bottom-up prey variation among sites separated by a few kilometers. No other intertidal prey elicited positive individual predator responses in this species, even though a large number of other prey species was always included in the diet. Moreover, examining predator-prey correlations at approximately 80, 160, and 200 km did not change this pattern, suggesting that positive prey feedback could occur over even larger spatial scales or as a geographically unstructured process. Thus individual-level responses were not transferred to population changes over the range of spatial scales examined here, highlighting the need to examine community regulation processes over multiple spatial scales.     

Behavioural responses of the marine bivalve Macomona liliana exposed to copper- and chlordane-dosed sediments

Sublethal behavioural responses including avoidance and burial rate were compared with 10 d acute morbidity and mortality in bioassays using juvenile Macomona liliana 1 to 3 mm long. The bivalves showed significant movement away from all copper-dosed sediments, with maximal movement after 96 h at 25 mg Cu kg (dry wt)^-1. Increasing copper concentrations slowed the rate of burial, and above 15 mg Cu kg (dry wt)^-1, most shellfish failed to bury after 90 min. After 10 d exposure, morbidity (defined as inability to rebury) occurred at 15 mg Cu kg (dry wt)^-1 and mortality at 30 mg Cu kg (dry wt)^-1. In avoidance trials, chlordane had a significant effect on overall movement from 20 g kg (dry wt)^-1, but increasing concentrations reduced movement away from dosed sediment. Chlordance appeared to have no effect on burial rate. Survival was affected by 10 d exposure to 400 g chlordane kg (dry wt)^-1, with increased morbidity and death. The relative sensitivities of the responses shown by M. liliana to copper were avoidance > burial/morbidity > mortality, with a 6-fold difference in the response threshold between avoidance and mortality. With chlordance, an avoidance response was detected at a concentration 20-fold lower than that causing morbidity. Behavioural responses of M. liliana have the potential to provide a sensitive bioassay.     

Linking direct and indirect data on dispersal: isolation by slope in a headwater stream salamander

There is growing recognition of the need to incorporate information on movement behavior in landscape-scale studies of dispersal. One way to do this is by using indirect indices of dispersal (e.g., genetic differentiation) to test predictions derived from direct data on movement behavior. Mark-recapture studies documented upstream-biased movement in the salamander Gyrinophilus porphyriticus (Plethodontidae). Based on this information, we hypothesized that gene flow in G. porphyriticus is affected by the slope of the stream. Specifically, because the energy required for upstream dispersal is positively related to slope, we predicted gene flow to be negatively related to change in elevation between sampling sites. Using amplified DNA fragment length polymorphisms among tissue samples from paired sites in nine streams in the Hubbard Brook Watershed, New Hampshire, USA, we found that genetic distances between downstream and upstream sites were positively related to change in elevation over standardized 1-km distances. This pattern of isolation by slope elucidates controls on population connectivity in stream networks and underscores the potential for specific behaviors to affect the genetic structure of species at the landscape scale. More broadly, our results show the value of combining direct data on movement behavior and indirect indices to assess patterns and consequences of dispersal in spatially complex ecosystems.     

Population differences in male responses to chemical mating cues in the white-spotted longicorn beetle, Anoplophora malasiaca

Adult Anoplophora malasiaca beetles were collected from field populations inhabiting three different host plants: Citrus, Salix and Vaccinium. Male orientation responses towards odours of wounded branches of each host plant were examined. Males of all three populations were attracted most frequently to odours of their original hosts, indicating that male orientation behaviour was induced by the odours of the plants from which they were originally collected. We also observed the mating responses of the males to elytra extracts from young and mature females of the three populations and chemically analysed those extracts. Citrus males were highly responsive to female extracts, regardless of origin. Salix males were less reactive than Citrus males, but responded moderately to female extracts of all populations. Vaccinium males were never attracted to Citrus female extracts and responded more to extracts from mature Vaccinium and Salix females, indicating that males might distinguish female origin and maturity. All eight hydrocarbons, four ketones, and three lactones that were previously identified as female contact sex pheromones were found in the extracts of three populations and both ages. Higher amounts of lactones on mature females likely caused greater male responses. These results suggested that males would be more likely to approach a female feeding on the same host plant and subsequently recognise her sexual maturity based on the profile of contact sex pheromones on her elytra.     

Variable predation risk and the dynamic nature of mosquito antipredator responses to chemical alarm cues

Summary. Predation is a pervasive selective agent highly variable in space and time. Due to the costs associated with antipredator responses, prey would be at a selective advantage if they respond to predation threats with an intensitfy matching the threat posed by the predator. Many aquatic organisms have been shown to use chemical alarm cues present in the water to assess the level of risk in their environment. This includes mosquito larvae which show antipredator responses to conspecific alarm cues. In this study, we investigated the nature of the responses of larval mosquitoes Culex restuans to those cues. In our initial observations, we showed pond/population differences in the response intensity of C. restuans to alarm cues. In experiment 1, we showed that the response intensity to alarm cues could be increased by increasing the background level of risk in the mosquitoes’ environment (by adding salamander predators) and once turned on, the response intensity to alarm cues was likely maintained for the remainder of the mosquitoes’ aquatic life. In experiments 2 and 3, we investigated if the increase in response intensity to alarm cues was directly correlated with the level of background risk in the mosquitoes’ environment. When given increasing levels of background risk, mosquito larvae subsequently showed a graded response to conspecific alarm cues. This series of experiments demonstrates that the response intensity of larval mosquitoes to a standard concentration of alarm cues is not fixed, but rather dependent on the background level of risk in the environment. An understanding of the background level of risk is particularly important for comparing antipredator responses of prey between habitats.     

Behavioural responses of Diana monkeys to male long-distance calls: changes in ranging,association patterns and activity

Although much is known about the relationship between vigilance, group size and predation risk, behavioural responses to predation risk and their resultant costs are less clear. We investigated the response of Diana monkeys to increased predation risk by looking at behavioural changes associated with male long-distance calls, which are reliably given to certain predators. After male long-distance calls, group spread and nearest-neighbour distance decreased whilst travel and association rates for the group increased. The average height and exposure level of individuals in the group did not change after calls. Individual Diana monkeys changed their behaviour and were more likely to be vigilant or travel and less likely to engage in social or resting behaviours after long-distance calls. In addition, movement rates increased with the number of species the Diana monkeys were associated with. Diana monkey long-distance calls facilitate the joining of groups of other species. Black and white colobus and lesser spot-nosed monkeys were more likely to be in an association following a long-distance call than before. Behavioural responses, such as increased travel or association rates, that reduce foraging efficiency are interpreted as evidence of a non-lethal impact of increased predation risk.     

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.     

Behavioral responses to a novel predator and competitor of invasive mosquitofish and their non-invasive relatives (Gambusia sp.)

Attributes of the recipient community may affect the invasion success of arriving non-indigenous organisms. In particular, biotic interactions may enhance the resistance of communities to invasion. Invading organisms typically encounter a novel suite of competitors and predators, and thus their invasiveness may be affected by how they cope with these interactions. Behavioral plasticity may help invaders to respond appropriately to novelty. We examined the behavioral responses of highly invasive mosquitofish to representative novel competitors and predators they might encounter as they spread through North America. We compared the behavior of invasive Gambusia holbrooki and G. affinis to that of two close relatives of lower invasive potential (G. geiseri and G. hispaniolae) in order to elucidate whether responses to novelty related to invasiveness. In short-term assays, female Gambusia were paired with a novel competitor, Pimephales promelas, and a novel predator, Micropterus dolomieu. Behavioral responses were measured in terms of foraging success and efficiency, activity, refuge use, predator inspections, and interspecific aggression. Contrary to a priori predictions, invasive and non-invasive responses to novel interactions did not differ consistently. In response to novel competition, both invasive species increased foraging efficiency, but so did G. geiseri. In response to novel predation, only G. holbrooki decreased consumption and activity and increased refuge use. No antipredator response was observed in G. affinis. We found consistent differences, however, between invasives and non-invasives in foraging behavior. Both in the presence and absence of the competitor and the predator, invasives foraged more efficiently and consumed more prey than non-invasives.Communicated by P. Bednekoff     

Linking movement, diving, and habitat to foraging success in a large marine predator

Establishing where and when predators forage is essential to understanding trophic interactions, yet foraging behavior remains poorly understood in large marine carnivores. We investigated the factors leading to foraging success in gray seals (Halichoerus grypus) in the Northwest Atlantic in the first study to use simultaneous deployments of satellite transmitters, time depth recorders, and stomach-temperature loggers on a free-ranging marine mammal. Thirty-two seals were each fitted with the three types of instrumentation; however, complete records from all three instruments were obtained from only 13 individuals, underscoring the difficulty of such a multi-instrument approach. Our goal was to determine the characteristics of diving, habitat, and movement that predict feeding. We linked diving behavior to foraging success at two temporal scales: trips (days) and bouts (hours) to test models of optimal diving, which indicate that feeding can be predicted by time spent at the bottom of a dive. Using an information-theoretic approach, a Generalized Linear Mixed Model with trip duration and accumulated bottom time per day best explained the number of feeding events per trip, whereas the best predictor of the number of feeding events per bout was accumulated bottom time. We then tested whether characteristics of movement were predictive of feeding. Significant predictors of the number of feeding events per trip were angular variance (i.e., path tortuosity) and distance traveled per day. Finally, we integrated measures of diving, movement, and habitat at four temporal scales to determine overall predictors of feeding. At the 3-h scale, mean bottom time and distance traveled were the most important predictors of feeding frequency, whereas at the 6-h and 24-h time scales, distance traveled alone was most important. Bathymetry was the most significant predictor of feeding at the 12-h interval, with feeding more likely to occur at deeper depths. Our findings indicate that several factors predict feeding in gray seals, but predictor variables differ across temporal scales such that environmental variation becomes important at some scales and not others. Overall, our results illustrate the value of simultaneously recording and integrating multiple types of information to better understand the circumstances leading to foraging success.     

Behavioral and antennal electrophysiological responses of a predator ant to the pygidial gland secretions of two species of Neotropical dung roller beetles

The pygidial gland secretions of Neotropical dung roller beetles appear to function as defensive agents against predator ants. Behavioral and electrophysiological tests were used to evaluate the responses of the ant Camponotus sericeiventris to pygidial compounds secreted by the dung roller beetles Canthon cyanellus cyanellus and Canthon femoralis femoralis (Coleoptera: Scarabaeidae). Our behavioral investigation revealed a lower aggressive response of C. sericeiventris to the dung roller beetles compared to their response to the fly Anastrepha ludens. Additionally, C. sericeiventris workers attacked C. f. femoralis more often than C. c. cyanellus. The behavioral aggression of C. sericeiventris was greater toward male than female C. c. cyanellus. Electroantennography showed that the antennae of C. sericeiventris react to pygidial extracts from C. c. cyanellus and C. f. femoralis. Three compounds from male and female C. f. femoralis and C. c. cyanellus elicited coupled gas chromatography-electroantennogram (GC-EAD) activity in the antennae of C. sericeiventris. Geraniol and an unknown compound (A) were specific for C. f. femoralis, whereas guaiacol and another unknown compound (B) were specific to C. c. cyanellus; phenol was shared by both beetle species. This is the first report confirming that the pygidial gland secretions of dung roller beetles play a role in their defense against predators.     

Leaf quality, predators, and stochastic processes in the assembly of a diverse herbivore community

Ecological communities are structured by both deterministic, niche-based processes and stochastic processes such as dispersal. A pressing issue in ecology is to determine when and for which organisms each of these types of processes is important in community assembly. The roles of deterministic and stochastic processes have been studied for a variety of communities, but very few researchers have addressed their contribution to insect herbivore community structure. Insect herbivore niches are often described as largely shaped by the antagonistic pressures of predation and host plant defenses. However host plants are frequently discrete patches of habitat, and their spatial arrangement can affect herbivore dispersal patterns. We studied the roles of predation, host plant quality, and host spatial proximity for the assembly of a diverse insect herbivore community on Quercus alba (white oak) across two growing seasons. We examined abundances of feeding guilds to determine if ecologically similar species responded similarly to variation in niches. Most guilds responded similarly to leaf quality, preferring high-nitrogen, low-tannin host plants, particularly late in the growing season, while bird predation had little impact on herbivore abundance. The communities on the high-quality plants tended to be larger and, in some cases, have greater species richness. We analyzed community composition by correlating indices of community similarity with predator presence, leaf quality similarity, and host plant proximity. Birds did not affect community composition. Community similarity was significantly associated with distance between host plants and uncorrelated with leaf quality similarity. Thus although leaf quality significantly affected the total abundance of herbivores on a host plant, in some cases leading to increased species richness, dispersal limitation may weaken this relationship. The species composition of these communities may be driven by stochastic processes rather than variation in host plant characteristics or differential predation by insectivorous birds.     

Behavioural and electrophysiological responses of the olive fruit fly, Bactrocera oleae (Rossi) (Diptera: Tephritidae), to male- and female-borne sex attractants

In the olive fruit fly, Bactrocera oleae, females attract males by producing 1,7-dioxaspiro[5.5]undecane (olean), the main component of the sex pheromone secreted by rectal glands. It has been recently demonstrated that males are able to produce (Z)-9-tricosene (muscalure) in rectal glands, a compound that selectively attracts females. In this study, a male grooming reaction that may transfer the male-borne compounds from rectal to urotergal glands was observed, suggesting that urotergal glands could be involved in B. oleae sexual communication. GC/MS, EAG, GC/EAD analyses and behavioural assays were carried out to compare the role of male rectal and urotergal glands during courtship. In both male glands, olean and muscalure amounts were age dependent. Extracts of rectal glands contained higher amounts of olean and/or muscalure than urotergal ones. Extracts of rectal and urotergal glands of males and females elicited EAG responses in both sexes. GC/EAD showed that female EAG response to male rectal extracts was mainly due to olean and muscalure. Synthetic compounds evoked EAG dose-dependent responses in both sexes, and the EAG response to muscalure was higher as compared to olean. Rectal and urotergal glands from old males were able to attract females, while urotergal glands from young males attracted only males. Overall, our results add knowledge to the mating system of B. oleae, giving first evidences on the electrophysiological activity of muscalure towards both sexes, as well as on the involvement of male urotergal glands in the chemical sexual communication of this pest.     

Transient nestmate recognition cues contribute to a multicolonial population structure in the ant,Leptothorax curvispinosus

Summary Individual nests of the facultatively polygynous and polydomous ant, Leptothorax curvispinosus, were mapped at two sites, collected, and maintained under uniform laboratory conditions. Tests of worker acceptability between nests were conducted 2–4 weeks and 13–17 weeks after collection. Nests collected near to one another (0.09–1.87 m) were sometimes nonaggressive, and were significantly less aggressive than those from different sites (7 km apart); and there was no significant difference in aggressiveness between tests for these distance categories. However, aggression between nests collected farther away from one another at the same site (1.52–4.65 m) decreased significantly between tests: initially, the level of aggressiveness was equivalent to that between nests from different sites but later it was reduced to that between near nests. These results indicate that polydomous colonies of this species occur within multicolonial populations; and that colony segregation within local populations is largely maintained by transient environmentally-based nestmate recognition cues. More stable cues of genetic or environmental origin (or both) are also present and contribute to discrimination even after extended periods of culture under uniform conditions. These results suggest that the maintenance of colony autonomy within genetically highly interrelated populations may be the prime function of environmentally-based nestmate recognition cues. Colony autonomy under such circumstances may be important to maintain a relatively small but optimal colony size, or because the mechanisms which regulate colony growth, development, etc., require a limited colony size.     

Induced responses to herbivory in the Neotropical ant-plant association between Azteca ants and Cecropia trees: response of ants to potential inducing cues

Plant defense against herbivores often involves constitutive and inducible mechanisms of resistance. Obligate ant-plants, which provide food and housing for ants, are thought to primarily rely on ants for defense against herbivores. This form of plant defense has largely been viewed as static. We have been investigating the dynamic nature of Azteca ants as an inducible defense of Cecropia trees. Ants rapidly recruit to and patrol sites of foliar damage. We propose that Azteca ants can be viewed as an inducible defense for Cecropia trees because of their sensitivity to cues associated with herbivory, their rapid and aggressive recruiting ability, and their reclaimable and redeployable nature as a plant defense. In this study, we examine ant behavior following plant damage, and the potential cues that indude ant recruitment. We found that ants present on leaves when the plant is damaged leave the damaged leaf and recruit other ants to it, presumably by laying recruitment trails. Volatile leaf cues associated with herbivory were important in eliciting an induced response in two experiments. However, we found that cues associated with a congeneric plant elicited a much stronger ant response than conspecific cues. Although the type of leaf damage (gaping wounds versus leaf edge wounds) did not affect the level of ant recruitment, the extent of damage did. Leaves with one hole punched showed a 50% increase in ants, while leaves with five holes punched in them elicited a 100% increase in ant numbers. In sum, it appears that multiple plant-related cues associated with herbivory are involved in induction of ant recruitment in the Cecropia-Azteca system. We discuss the generality of ant responses to herbivory in obligate ant-plant systems, and in facultative ant-plant associations, which may be more common. Received: 23 March 1998 / Accepted after revision: 5 July 1998     

Adaptive responses to local mate competition by the parasitoid,Telenomus remus

Summary The parasitic wasp, Telenomus remus, lays her eggs in diserete patches of moth eggs, where her offspring develop and mate before dispersal, satisfying conditions for local mate competition (LMC). In the presence of other ovipositing females, wasps lay a higher sex ratio (proportion males), as predicted by LMC theory, and achieve this by a combination of two mechanisms, (1) avoidance of superparasitism and a sequence of sex allocation initially biased towards males and (2) a direct increase in sex ratio in the presence of other wasps, sex ratio increases with the proportion of previously parasitized hosts, as predicted by LMC theory. In both cases, chemical traces left by foraging wasps are indicated as the stimuli causing wasps to increase the proportion of males allocated to hosts.