Influence Of Diet-Related Chemical Cues from Predators on the Hatching of Egg-Carrying Spiders

Previous studies have shown that animals may make adaptive adjustments in response to chemical cues from predators, but hatching responses to diet-related chemical cues from predators have not been previously demonstrated. In the system studied here, the predator is an araneophagic jumping spider (Salticidae), Portia labiata, and the prey organism is a subsocial spitting spider, Scytodes pallida (Scytodidae). The spitting spider carries its eggs in its chelicerae, and carrying eggs is known to make it more vulnerable to predators. It is also known from an earlier study that the prior diet of the predator alters how dangerous the individual predator is to the spitting spider. In the experiments reported here, incubation time was shorter when volatile cues from the predator were present and longer in control tests when no chemical cues from the predator were present. The previous predators diet also influenced incubation time: when in the presence of volatile cues from individuals of P. labiata that had previously fed on individuals of S. pallida, incubation time was shorter than when in the presence of volatile cues from individuals of P. labiatathat had been feeding instead on house flies.     

Timber Rattlesnakes (Crotalus horridus) Use Chemical Cues to Select Ambush Sites

Chemicals left by organisms moving through the environment are used by other organisms to mediate interspecific interactions. Most studies of chemical eavesdropping focus on prey responding to chemical cues from predators, despite the fact that chemical cues are frequently used by predators as a source of information about prey. Crotalus horridus uses a foraging strategy that is widespread among sedentary predators: the snake chooses a site where it is likely to encounter prey and remains immobile for many hours. I investigated this ambush hunting behavior in captive-raised timber rattlesnakes and provide evidence that sit-and-wait predators may discriminate among prey chemical cues, even when they have no prior experience with the prey. Snakes explored chemical cues with chemosensory behaviors, and more frequently adopted a stereotyped ambush foraging posture toward chemical cues from prey sympatric with their population of origin than either allopatric prey or sympatric nonprey species that are eaten by other viperids. These results support the notion that intra- and interspecific variation in diet may be mediated proximally by innate recognition of cues from particular prey items. This system also describes a bioassay that may be used in the isolation and identification of prey-derived kairomones. Studies such as this can be used to determine more realistic parameters for models of predator-prey interaction and foraging behavior that involve secretive, less active predators.     

Chemical and Physical Signals Mediating Conspecific and Heterospecific Aggregation Behavior of First Instar Stink Bugs

We investigated cues that mediate the aggregation behavior of immature pentatomid bugs by using nymphs of six different pentatomid bug species (Nezara viridula, Acrosternum hilare, Chlorochroa ligata, Chlorochroa sayi, Thyanta pallidovirens, and Euschistus conspersus). When first instars of any two species were put together in a Petri dish, they readily formed heterospecific aggregations similar to their natural conspecific aggregations. The chemical profiles of first and second instar nymphs of each species were determined by solvent extraction with pentane, followed by GC-MS analysis. Immature bugs of the different species had some compounds in common, and some that were more species specific. Within a species, there were distinct differences in the profiles of compounds extracted from first and second instars. Bugs did not aggregate around untreated polysulfone beads (1 mm diam) that were glued together in groups approximating bug egg masses, suggesting that tactile cues alone were insufficient to induce aggregation. Furthermore, when tested over a range of doses, groups of polysulfone beads treated with crude whole-body extracts of bugs did not induce or maintain aggregations. However, first instar N. viridula nymphs did respond to beads treated with two of the three major components of bug extracts. 4-Oxo-(E)-2-decenal induced significant aggregations at two doses, whereas tridecane, the major component in extracts from all six species, did not, and (E)-2-decenal was repellent. The repellence of (E)-2-decenal may explain why we and previous researchers were unable to induce aggregations of first instar N. viridula using whole-body extracts.     

Ethanol and Methanol as Possible Odor Cues for Egyptian Fruit Bats (Rousettus aegyptiacus)

Frugivorous bats from the Old and New World use odor cues to locate and assess fruit condition. We hypothesized that Egyptian fruit bats (Rousettus aegyptiacus) use as odor cues those volatile compounds that increase in emission rate as fruit ripens. We examined whether the smell of fermentation products may indicate the degree of ripeness to fruit bats. We analyzed volatile compounds in the headspace (the gas space above a fruit in a closed container) of dates (Phoenix dactylifera) and rusty figs (Ficus rubiginosa), both of which are consumed by fruit bats, to elucidate which compounds originate from fermentative pathways and to determine which change in emission rate during ripening. Ethanol, acetaldehyde, and acetic acid were the only volatile compounds detected as products of fermentation in both fruits. In dates, emission rates of these compounds increased during maturation, whereas in rusty figs, they decreased or remained constant. Methanol, although not a fermentation product, increased in emission rate during ripening in both fruits. We found that R. aegyptiacus was neither attracted nor deterred by the smell of methanol at any of the concentrations used. Although the odor of ethanol emanating from food containing concentrations similar to those found in ripe fruit did not attract the bats, at relatively high concentrations (≥1%), the smell of ethanol deterred them. Thus, ethanol at high concentrations may serve as a signal for bats to avoid overripe, unpalatable fruit.     

Learned Recognition of Predation Risk by Enallagma Damselfly Larvae (Odonata, Zygoptera) on the Basis of Chemical Cues

We studied two populations of damselfly larvae (Enallagma boreale): one population cooccurred with a predatory fish (northern pike, Esox lucius); the other did not. Damselflies that cooccurred with pike adopted antipredator behavior (reduced activity) in response to chemical stimuli from injured conspecifics, and to chemical stimuli from pike, relative to a distilled water control. Damselflies from an area where pike do not occur responded only to chemical stimuli from injured conspecifics. In a second set of experiments, we conditioned pike-naive damselflies to recognize and respond to chemical stimuli from pike with antipredator behavior. Damselfly larvae that were previously unresponsive to pike stimuli learned to recognize pike stimuli after a single exposure to stimuli from pike and injured damselflies or pike and injured fathead minnows (Pimephales promelas). The response to injured fathead minnows was not a general response to injured fish because damselfly larvae did not respond to chemical stimuli from injured swordtails (Xiphophorus helleri), an allopatric fish. Taken together, these data suggest a flexible learning program that allows damselfly larvae to rapidly acquire the ability to recognize local predation risk based on chemical stimuli from predators, conspecifics, and heterospecific members of their prey guild.     

Chemical Defense in the Plant Bug Lopidea robiniae (Uhler)

Secretions from the metathoracic glands (MTG) of the black locust bug, Lopidea robiniae (Uhler) (Heteroptera: Miridae) contained six major compounds, including (E)-2-hexenal, (E)-2-hexen-1-ol, (E)-2-octenal, (E)-2-octen-1-ol (E)-2-heptenal, and (Z)-3-octen-1-ol. Males and females did not differ significantly in the relative compositions of identified compounds. In feeding trials, six bird species [robin (Turdus migratorious), blue jay (Cyanocitta cristata), brown thrasher (Toxostoma rufum), killdeer (Charadrius vociferus), starling (Sturnus vulgaris), and house wren (Troglodytes aedon)] demonstrated feeding aversions towards L. robiniae, implying that black locust bugs are chemically defended. Bugs discharged the liquid contents of their MTG when attacked, thereby producing a strong and distinct odor. Some birds immediately ejected bugs out of their mouth after biting them, suggesting that the MTG secretion was a deterrent.     

Alarm response by a plethodontid salamander (Desmognathus ochrophaeus): Conspecific and heterospecific “Schreckstoff”

The detection of chemical alarm cues plays an important role for predator avoidance in many taxonomic groups, but little is known about the presence of such chemical cues in adult or caudate amphibians. We investigated the response (i.e., aversion or nonaversion) to chemical cues from damaged salamander skin and mealworms (Tenebrio molitor) in the plethodontid salamander,Desmognathus ochrophaeus. Avoidance responses were demonstrated to skin extracts of both conspecific and heterospecific salamanders. However, salamanders (D. ochrophaeus) did not avoid heated conspecific skin, fresh conspecific viscera, fresh mealworm, or freshPlethodon richmondi skin extracts. These results indicate that chemical alarm cues are: (1) present in the skin ofDesmognathus salamanders, (2) not present in mealworm or the viscera ofDesmognathus salamanders, and (3) denatured or deactivated by heating. These results also suggest that an avoidance response to chemical cues from damaged conspecifics has adaptive value in predator avoidance in terrestrial as well as aquatic vertebrates.     

Evidence for Use of Chemical Cues by Male Horseshoe Crabs When Locating Nesting Females (Limulus polyphemus)

Horseshoe crabs come ashore in attached pairs during spring high tides to mate and nest on beaches of the Atlantic and Gulf coasts. Unattached males also come ashore and crowd around the nesting pairs as satellites and engage in sperm competition with the attached male. Females with no satellites and females with large numbers of satellites nest next to one another on the same tide. When females are removed and replaced by a cement model, satellite males continue to be attracted to the same location. Models over sites where females with many satellites had nested are more attractive to males than sites from which a female with no satellites had been removed or a site where no crab had been nesting recently. A second experiment demonstrated that males are responding to chemical cues. A sponge filled with seawater taken from below a female with many satellites and placed under a model female was more attractive to males than a sponge filled with seawater. This is the first demonstration that horseshoe crabs use chemical cues, in addition to visual cues, to locate mates.     

Detection and Discrimination of Conspecific Scents by the Anguid Slow-Worm Anguis fragilis

We tested the ability of male slow-worms, Anguis fragilis, a limbless anguid lizard with secretive, semifossorial habits, to detect chemical associated with conspecifics by using a T-maze in the laboratory. Male slow-worms discriminated conspecific male and female scent deposits. Males selected the arm with female scent, suggesting that scent deposits may be used to locate potential mates. Also, male slow-worms did not avoid the chemicals of other males, suggesting that they are not territorial. However, males discriminated their own scent from those of other males, and spent more time exploring the arm with other male scent, which suggests that scent marks may bear information that could be used in future intrasexual social contexts. We conclude that discrimination of conspecifics based on scents may be more widespread than previously expected among lizards inhabiting visually restricted environments.     

Chemical communication in adult schistosomes

Lipids released bySchistosoma mansoni adult males attract females in vitro. Lipid release is modulated by the presence of other worms. AlthoughS. mansoni males release lipid when paired with females, the release is enhanced when they are separated.S. japonicum adults release more free sterols when incubated individually than when incubated together. Similarly, individually incubatedS. haematobium males release more free sterols than when incubated in groups. However,S. haematobium females incubated in groups release more free fatty acids than do equal numbers of males or pairs incubated in groups. There is evidence thatS. mansoni adult females concomitantly accumulate and release cholesterol in the absence of an exogenous supply, although de novo synthesis of cholesterol in schistosomes has not yet been demonstrated. Schistosomula and adult schistosomes incorporate exogenous lipids. Lipids are incorporated chiefly through the tegument. Cholesterol is transferred between males and females.     

Wolf Spiders Show Graded Antipredator Behavior in the Presence of Chemical Cues from Different Sized Predators

The wolf spider, Pardosa milvina, displays effective antipredator behavior (reduced activity) in the presence of silk and excreta cues from adults of another cooccurring wolf spider, Hogna helluo. However, Pardosa and Hogna engage in size-structured intraguild predation, where Pardosa may be either the prey or predator of Hogna. We tested the ability of adult female Pardosa to vary antipredator responses toward kairomones produced by Hogna that vary in size. Hogna were maintained on filter paper for 24 hr. We then presented the paper to adult female Pardosa simultaneously paired with a blank sheet of paper. One treatment had two sheets of blank paper to serve as a control. The Hogna stimulus treatments were as follows (N = 15/treatment): (1) 1 Hogna half the mass of Pardosa; (2) 1 Hogna of equal mass of a Pardosa; (3) 1 adult Hogna, 30 times the mass of Pardosa; and (4) 8 Hogna each 0.25 the mass of Pardosa. Pardosa decreased activity in the presence of kairomones from Hogna of equal or larger size, but showed no change in activity in the presence of a blank control or from a single Hogna smaller than itself. Pardosa showed a reduction in activity in the presence of cues from eight small Hogna. Pardosa avoided substrates with adult Hogna cues, but showed no avoidance response to any other treatment. These results suggest that Pardosa is showing graded antipredator behavior relative to the quantity of predator kairomones present rather than directly discriminating among the different sizes of the predator.     

Multifunctional communication inRiptortus clavatus (Heteroptera: Alydidae): Conspecific nymphs and egg parasitoidOoencyrtus nezarae use the same adult attractant pheromone as chemical cue

The bean bug,Riptortus clavatus lays scattered eggs (as opposed to the egg masses of pentatomids) on host as well as nonhost plants. Therefore, the first feeding stage (second-instar) nymphs emerging from eggs laid on nonhost plants need a signal that enables them to locate a food source at the lowest energy cost. Male-released (E)-2-hexenyl (E)-2-hexenoate, (E)-2-hexenyl (Z)-3-hexenoate, and myristyl isobutyrate play the double role of attractant pheromone for adults as well as aggregation pheromone, which enables the second-instar nymphs to find the host food plant. These male-specific semiochemicals are released only when foodstuff is available. On the other hand, females ofOoencyrtus nezarae, the most effective parasitoid of the host in Kumamoto, Japan (where the field experiments were conducted), utilize these semiochemicals as kairomones in order to locate the potential host community. Field experiments revealed that the synthetic pheromone rivaled 10 live males in the attraction of adults and second-instar nymphs. Captures of the egg parasitoidO. nezarae females in cylindrical sticky traps were significantly higher in traps baited with the synthetic semiochemicals than in control traps. The number of females captured was significantly higher than the number of males, although the captures in the sticky suction trap system revealed that the populations of male and female were not significantly different.     

Chemical communication in the mating behavior ofTrogoderma glabrum (Herbst) (Coleoptera: Dermestidae)

Male mating behavior of the stored product pest beetleTrogoderma glabrum (Herbst) was analyzed into three major phases: arousal/searching, preliminary recognition, and genital (copulatory). Airborne pheromone released by calling females elicits male arousal via antennal sensillae. Contact chemoreception via male mouthpart sensillae appears to be necessary for copulation to occur. A procedure was developed for quantitative bioassay of presumptive pheromone compounds in both airborne and contact chemoreception. (E)-14-methyl-8-hexadecenal, present in airborne pheromone, but not detectable in extracts of whole females, elicits both arousal and attempted copulation. The activity of (E)-14-methyl-8-hexadecenal is equal to that of the total airborne pheromone. Activity of additional possible pheromone component candidates [including (E)-14-methyl-8-hexadecen-1-ol, -caprolactone,n-hexanoic acid, and methyl (Z)-7-hexadecenoate] was investigated. Evidence for a behavioral role forn-hexanoic acid is presented.     

Chemical communication in the dacetine antDaceton armigerum (Hymenoptera: Formicidae)

Contrary to previous assumptions,Daceton armigerum, the largest ant in the myrmicine tribe Dacetini, employs trail communication. We identified two anatomical sources of trail pheromones: Trails drawn with poison gland contents can last for more than seven days. Trails drawn with the newly discovered sternal glands (in the VIth and VIIth abdominal sternites) are effective but relatively short-lived. In addition, our bioassays revealed that the contents of the mandibular glands elicit alarm behavior, and secretions from the pygidial gland release attraction. When tested with artificial poison gland trails from seven other myrmicine species,Daceton did not exhibit trail following behavior. We confirmed, however, previous findings thatAtta respond toDaceton poison gland trails andSolenopsis followDaceton Dufour's gland trails.     

Chemical cues associated with visually elaborate aggressive displays of three-spine sticklebacks

A two-choice chamber was used to determine if three-spine sticklebacks could distinguish between water conditioned by a displaying male and water conditioned by a male that was active but not displaying. Both nonterritorial males and gravid females appeared to recognize which male was displaying; i.e., males and females were more likely to bite or bump the substrate in front of a chamber receiving water from a displaying male. Nonterritorial males were more likely to enter the chamber receiving water from a male that was not displaying; gravid females entered the chamber that received water from the displaying male more often than males but not more often than expected by chance. The aggressive displays of male sticklebacks may have both visual and chemical components; alternatively, the fish may have been capable of detecting chemical by-products associated with different rates of physical activity.     

Quantitative and qualitative variation in male pheromones ofPhragmatobia fuliginosa andPyrrharctia isabella (Lepidoptera: Arctiidae)

The dihydropyrrolizine pheromones, hydroxydanaidal and danaidal, were identified from the scent organs of malePhragmatobia fuliginosa (L.) andPyrrharctia isabella (J.E. Smith). Qualitative and quantitative GLC analyses were conducted on ca. 80 field-collected males of each species. The total pheromone titer was distributed bimodally in each species with most males having either a small amount (< 10 ng) of pheromone or a large amount (1–10 g inPyrrharctia and 0.3–3 g inPhragmatobia).Pyrrharctia males in the 1- to 10-g range had a predominance of hydroxydanaidal, with little if any danaidal. MostPhragmatobia males in the 0.3- to 3-g range had danaidal with little if any hydroxydanaidal. These compounds elicited a courtship response in sexually receptive females of both species. A bioassay based on this response was used to measure the thresholds of female response to these compounds.Pyrrharctia females were more sensitive to (R)-(–)-hydroxydanaidal than to danaidal.Phragmatobia females were more sensitive to danaidal then to (R)-(–)-hydroxydanaidal.     

Olfactory Sensitivity of the Gilthead Seabream (Sparus auratus L) to Conspecific Body Fluids

The potential for intraspecific chemical communication in the gilthead seabream (a marine perciform) was investigated by assessing the olfactory sensitivity to conspecific body-fluids (water occupied by conspecifics, intestinal fluid, urine, semen, egg fluid) by multiunit electrophysiological recording from the olfactory nerve. The olfactory system was responsive to water previously occupied by conspecifics, and the active compound(s) could be extracted by solid-phase extraction. The olfactory system was extremely sensitive to body fluids of sexually mature conspecifics: thresholds of detection were 1:10^7.4 (intestinal fluid), 1:10^6.1 (gametes), and 1:10^4.2 (urine). The olfactory system was also sensitive to amino acids with thresholds of detection from 10^–8.1 M (l-leucine) to 10^–6.1 M (l-phenylalanine). However, a range of other known fish odorants (steroids, bile acids, and prostaglandins) failed to evoke significant responses. Given the high olfactory sensitivity to intestinal fluid and the low urine release rates of marine compared with freshwater fish, we suggest that chemical communication is likely to be mediated via compounds present in the intestinal fluid rather than urine. Furthermore, the types of chemicals involved are likely to be different from those of freshwater fish. Their exact chemical identity and biological roles remain to be established.     

Mammalian Exocrine Secretions. XVII: Chemical Characterization of Preorbital Secretion of Male Suni, Neotragus moschatus

Gas chromatographic and gas chromatographic–mass spectrometric techniques were employed to identify 83 compounds, including alkanes, alkenes, aldehydes, 2-methylalkanes, carboxylic acids, 1-alkyl formates and alken-1-yl formates, benzoic acid, and cholesterol, in the preorbital secretion of the male suni, Neotragus moschatus. Dimethyl disulfide derivatization and lithium aluminum hydride reduction were used to determine the position of double bonds and to confirm the identity of the functional groups in some of the constituents of the secretion.     

Chemical defense secretions of some species of Malaysian rhinotermitidae (Isoptera,Rhinotermitidae)

The defense secretions of the soldiers of the Malaysian rhinotermitid,Parrhinotermes aequalis (Havilandi) andP. pygmaeus (John),Termitogeton planus (Havilandi) andSchedorhinotermes malaccensis (Holmgren) consist mainly of vinyl ketones, whereas that ofProrhinotermes flavus (Bugnion & Popoff) gives (E)-1-nitropentadecene. The chemistry of the defense secretions ofParrhinotermes andTermitogeton is documented and based on their chemical relationships;Termitogeton shows a closer affinity to the Rhinotermitinae than Heterotermitinae.     

Chemical detection of “self” and conspecifics by crayfish

Stimulus waters were used to examine behaviorally chemical communication in female crayfishProcambarus clarkii. Animals detected conspecifics. Response to stimulus water drawn from the animal's own tank, self water, mimicked either response to distilled water or response to water drawn from the tank of another animal. The response to self water depended on the relative concentration of substance(s) in self water stimuli to that of the same substance(s) in the test animal's tank.