Female sex pheromone-mediated effects on behavior and consequences of male competition in the shore crab (Carcinus maenas)

Exposure to receptive female pheromone elicits guarding behavior in shore crab males (Carcinus maenas), but little is known about the effects of sex pheromone on male competition or if the female plays an active role in mate choice. This study examined whether female pheromone enhanced agonistic behavior between males and what effects visual and chemical cues had on the rules and costs of such contests. We also investigated whether females exhibit a preference for males in terms of size. Under laboratory conditions, solitary male shore crabs engaged males who already had possession of a female. The visual and chemical presence of a receptive female had an impact on contest rules and costs. Fights were costly in terms of duration and of sustaining injury with either one or both crabs incurring injury in 40% of fights. To investigate the metabolic consequences of fighting over a perceived sexual resource (chemical cue only), fights were staged between pairs of size-matched males in the presence of water containing the female sex pheromone, water in which males had been kept, and untreated seawater. The duration and intensity of contests were greater when staged in the presence of the female pheromone compared with the two other treatments. Crabs that fought in the presence of female sex pheromone also had a greater accumulation of L-lactate and a reduction of glycogen stores. Fights were less costly in terms of injury with a single chemical cue compared with enhanced costs with a multiple sexual cue. The importance of female choice was determined by presenting postmolt females with different sized males. Males were kept in a fixed position, and the majority of females approached and performed courtship behavior to the largest males, demonstrating that females may be selective in terms of size.     

Chemically Mediated Mate Location and Evaluation in the Lobster, Homarus americanus

Many aspects of lobster (Homarus americanus) social behavior have been shown to be regulated by chemical signals. Female lobsters can determine the sex of sheltered conspecifics from a distance, and will approach and attempt to enter male-occupied shelters. We investigated (1) if female attraction to male-occupied shelters and entering decisions are based upon chemical signals, particularly male urine signals, and (2) if male dominance status and female molt stage influence these female behaviors. Sheltered dominant males were fitted with urine catheters, which collect urine and prevent its release into the environment. We found that males increased urine output in response to female entering attempts. Blocking male urine release significantly reduced the incidence of female shelter approach and the time spent attempting to enter. Artificial release of male urine in the presence of a catheterized male restored female approach but not time spent attempting to enter, while artificial urine release alone did not induce either behavior. Females approached any single male, but seemed to prefer to enter shelters occupied by more dominant males. Premolt and intermolt females differed in their behaviors: premolts showed more interest in approaching dominant male shelters without much effort to enter, while intermolts appeared less likely to approach but more motivated to attempt to enter shelters. This study suggests that both intermolt and premolt female lobsters use one or more chemical signals from sheltered males for location and evaluation of potential mates. Male urine appears to carry compound(s) important for both location and entering decisions, although nonurine chemical or other sensory signals are also implicated during close-up mate evaluation.     

Factors affecting behavioral phase transition in the desert locust,Schistocerca gregaria (Forskål) (Orthoptera: Acrididae)

The behavior of the desert locust,Schistocerca gregaria (Forskål) (Orthoptera: Acrididae), is adjusted rapidly to population density and is a phase characteristic. We used discriminant analysis to quantify the extent of phase transition from the solitary to the gregarious phase and accurately classify the phase status on the basis of two decisive behavioral parameters: nymphal activity and social interaction. Fecal extracts. examined by olfactometry, attract solitarious nymphs but do not contribute to behavioral phase transition. Neither do visual stimuli alone. Short-range olfaction of airborne volatiles slightly affects behavioral phase transition. Antennectomy abolishes density response. Cuticular lipid extract, containing presumptive contact pheromones, does not attract nymphs, but does significantly affect behavioral phase transition.     

Stereochemistry of Host Plant Monoterpenes as Mate Location Cues for the Gall Wasp Antistrophus rufus

In spring, adult males of the gall wasp Antistrophus rufus L. emerge from inconspicuous galls in stems of their host plant Silphium laciniatum L. and search for sites on stems where females will later emerge. The behavior of males suggests that they use olfaction rather than visual or tactile cues in searching for mates. In an earlier publication, we reported that galls of A. rufus were associated with changes in enantiomeric ratios of alpha- and beta-pinene emitted by plant stems, and hypothesized that monoterpene stereochemistry served as a mate location cue for adult males. Here, we support this hypothesis with bioassays that demonstrate that males can discriminate between galled and ungalled stems, as well as between blends of synthetic monoterpenes with ratios of enantiomers representative of galled and ungalled stems.     

The Desert Locust, <Emphasis Type="Italic">Schistocerca gregaria</Emphasis>, Detoxifies the Glucosinolates of <Emphasis Type="Italic">Schouwia purpurea</Emphasis> by Desulfation

Desert locusts (Schistocerca gregaria) occasionally feed on Schouwia purpurea, a plant that contains tenfold higher levels of glucosinolates than most other Brassicaceae. Whereas this unusually high level of glucosinolates is expected to be toxic and/or deterrent to most insects, locusts feed on the plant with no apparent ill effects. In this paper, we demonstrate that the desert locust, like larvae of the diamondback moth (Plutella xylostella), possesses a glucosinolate sulfatase in the gut that hydrolyzes glucosinolates to their corresponding desulfonated forms. These are no longer susceptible to cleavage by myrosinase, thus eliminating the formation of toxic glucosinolate hydrolysis products. Sulfatase is found throughout the desert locust gut and can catalyze the hydrolysis of all of the glucosinolates present in S. purpurea. The enzyme was detected in all larval stages of locusts as well as in both male and female adults feeding on this plant species. Glucosinolate sulfatase activity is induced tenfold when locusts are fed S. purpurea after being maintained on a glucosinolate-free diet, and activity declines when glucosinolates are removed from the locust diet. A detoxification system that is sensitive to the dietary levels of a plant toxin may minimize the physiological costs of toxin processing, especially for a generalist insect herbivore that encounters large variations in plant defense metabolites while feeding on different species.     

Airborne Chemical Communication in the Wolf Spider Pardosa milvina

Most studies involving chemical communication in spiders focus on contact pheromones attached to spider silk. Here we tested if males of the wolf spider Pardosa milvina use airborne pheromones to identify, locate, and follow females. Using a two-choice olfactometer, we tested the response of adult male P. milvina to a number of potential chemical cues while controlling for concomitant visual and vibratory stimuli. An airborne chemical cue from adult virgin female P. milvina elicited a positive taxis response from the male. We also tested adult male responses to penultimate instar female P. milvina, one adult male P. milvina, and two adult males together. In each case, test males showed no attraction to the stimuli. Additional experiments were run with pitfall traps baited with adult virgin female P. milvina as attractants. Again, we controlled for visual and vibratory cues from females. Pitfall traps containing virgin females captured significantly more males than control traps. Collectively, these experiments demonstrate evidence of an airborne sex pheromone in P. milvina.     

Mate location strategies of gypsy moths in dense populations

In high-density populations, the male gypsy moth (Lymantria dispar) appetitive flight is primarily vertical and within 1–50 cm of tree boles. Preceding location of a female, males land on trees or occasionally tree foliage and walk while wing-fanning. In high-density populations females may mate before calling, often prior to wing expansion or hardening. Additional matings may occur before any or full deposition of egg masses. Virgin females are not coy, whether their wings are unexpanded, or hardened and held rooflike, or whether they are calling or not. They generally mate with the first willing male and do not exercise sexual selection by an acceptance or rejection reaction. The mating structure in dense populations would seem to reduce selective pressure for female emission of and direct male anemotactic flight to attractant pheromone. A low proportion of males search appropriate objects, especially tree trunks, in the walking-wing-fanning state. Such males often are successful in locating virgin females before calling commences. This male strategy presumably would not be successful in low population densities.Michigan Agricultural Experiment Station Journal Article No. 10253.     

Cuticular Surface Hydrocarbons of Desert Locust Nymphs, Schistocerca gregaria, and Their Effect on Phase Behavior

The quantity of cuticular hydrocarbons is higher in solitarious nymphs of the desert locust, Schistocerca gregaria, compared to gregarious nymphs, but the total hydrocarbon fraction of solitarious nymphs does not significantly divert behavioral transition of isolated nymphs to the gregarious phase, while gregarious hydrocarbon extracts do. This suggests that qualitative differences in composition are responsible for the biological effect. The profile of cuticular hydrocarbon components is similar in the two phases, but some peak ratios differ. Crowding of solitarious nymphs leads to rapid changes in the profile of the hydrocarbon fraction, suggesting that specific hydrocarbons are produced and secreted as a consequence. Isolating previously crowded nymphs has an opposite effect. The composition of cuticular hydrocarbons from the migratory locust, Locusta migratoria, which differs considerably from that of S. gregaria, does not induce the gregarious behavioral phase in solitarious nymphs of the latter.     

A Multi-Component Species Identifying Pheromone in the Goldfish

Although it has been established that sexually-immature goldfish and their relatives recognize members of their own species by using chemicals that they release, the identity of this cue(s) and whether it might be produced and used by other life stages is not yet known. To address this question, this study tested the behavioral responses of sexually immature and mature goldfish to each other’s body washings, their sensitivity to this cue, the role of the olfactory sense in detecting it, and whether it is comprised of either polar and/or non-polar compounds. Tests that used two-choice mazes discovered that juvenile, immature, mature male, and mature female goldfish all release and respond to a common chemical cue(s). Dilution studies next demonstrated that this cue is active when diluted over 10 times and thus capable of functioning as a short range attractant/identifier. Olfactory occlusion demonstrated that it is detected by the olfactory sense. Finally, chemical fractionation demonstrated that it is comprised of both polar and non-polar components but likely does not include bile acids. Together, these results suggest that all life stages of goldfish use a complex multicomponent pheromonal odor to discern species identity, and that this odor has the potential to function with hormonal metabolites to identify sexual condition in behaviorally active fish of many species.     

The annual cycle of Flehmen in black-tailed deer (Odocoileus hemionus columbianus)

Flehmen is a stereotyped response of ungulates and other mammals to urine. In black-tailed deer, Flehmen occurs typically in males (rarely in females), and its frequency shows an annual cycle. Flehmen in response to female urine was most frequent in November, and to male urine in January. The response minimum to both types of urine occurred in May. Fifty-five to 100% of initial responses to female urine resulted in Flehmen. The deer responded more often to female urine, and the difference between the responses to female and male urine increased from May to November. A male's own urine released Flehmen more often than did urine of other males. The response intensity varied with the spatial orientation of the male to the urinating female, and responses were usually limited to distances of 15 m or less.     

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.     

Attraction of male beetles to grubs: Evidence for evolution of a sex pheromone from larval odor

Females of the scarabaeid beetleCyclocephala lurida produce a volatile sex pheromone which attracts conspecific males. Field experiments demonstrated that larvae of both sexes also emit volatile chemicals that stimulate similar responses in adult males, including attempts by the attracted males to mate with the nonreproductive immature stage. Significantly more adult males were caught in traps baited with conspecific male or female larvae or adult females than in blank control traps. Hexane extracts of both male and female grubs were at least as effective as live larvae in trapping male adults, demonstrating that the behavioral responses are mediated by volatile chemicals. Sensory and behavioral responses of males to sex pheromones emitted by adult females are part of the functional communication system. However, their response to grubs is not functional, because grubs are normally temporally and spatially inaccessible to mate-seeking males. In theory, the evolution of a communication system is problematic because it requires the development of a signal in one sex and the sensory and behavioral attributes to respond to that signal in the other sex. The ontogeny of sex pheromone communication inC. lurida suggests a partial solution to this evolutionary problem. We propose that this sex pheromone communication system is probably derived from noncommunicative volatile chemicals that are lost in adult males and retained by adult females.     

Sexual communication and associative learning in the parasitic waspItoplectis conquisitor (Say)

Extracts of newly emerged virgin females and newly emerged virgin males of the parasitic waspItoplectis conquisitor were highly active in eliciting sexual behavior in older males. Newly emerged males did not respond to the extracts. A reliable biological assay was devised in which the responses of males to extract could be evaluated under varying conditions. Optimum male responses to female extract were obtained at 27 °C and at 1.0 female equivalents (the amount of pheromone extractable from the body surface of a female). A decrease in sexual response was observed when males were repeatedly exposed to pheromone. Males also learned to associate bioassay conditions with the presence of pheromone.     

Pheromone Trailing Behavior of the Brown Tree Snake, Boiga irregularis

The ability of snakes to follow pheromone trails has significant consequences for survival and reproduction. Of particular importance is the ability of snakes to locate conspecifics during the breeding season via the detection of pheromone trails. In this study, the ability of male brown tree snakes (Boiga irregularis), a tropical, rear-fanged colubrid, to follow pheromone trails produced by reproductively active conspecifics was tested in the laboratory by using a Y maze. Males displayed a trailing response to both female and male pheromone trails over blank controls. As males of this species display ritualized combat behavior, these responses likely represent both direct and indirect mechanisms, respectively, for the location of potential mates in the wild. Males did not, however, discriminate between male and female trails when given a choice on the Y maze.     

Changing Oviposition Preferences of the Desert Locust, Schistocerca gregaria, Suggest a Strong Species Predisposition for Gregarization

Field surveys at five sites within desert locust breeding habitats around Port Sudan during three successive seasons indicated that early in the rainy season the incoming solitary females oviposited predominantly in the vicinity of Heliotropium spp. (66%) and millet (32%) seedlings. Solitary nymphs also preferred to feed on these plants. Follow-up cage experiments were conducted in the field in which solitary and gregarious female locusts were presented with choices of selected desert plants and egg pods. When presented with bulrush millet, Heliotropium spp., Zygophyllum simplex, and untreated moistened sand, solitary females oviposited adjacent to the first two plants (40% and 60%, respectively). However, when offered a choice of either or both of these plants together with egg pods derived from gregarious and/or solitary insects, solitary females showed a significantly higher preference for ovipositing near gregaria egg pods than near the plants, with solitaria egg pods eliciting the least response. In contrast with solitary females and in the absence of gregaria egg pods, gregarious females preferred to oviposit in untreated moist (control) sand (74–77%) away from the plants (6–14%) or solitaria egg pods (4%). However, when present, gregaria egg pods elicited significantly more oviposition. These and previous results indicate a hierarchy of phase-dependent oviposition preferences in the desert locust and are interpreted in terms of a strong propensity of the species to exploit opportunities under appropriate conditions to facilitate congregation and the gregarization of the progeny.     

Constituents of wing gland and abdominal hair pencil secretions of male African sugarcane borer,Eldana saccharina walker (Lepidoptera: Pyralidae)

In addition totrans-3,7-dimethyl-6-octen-4-olide (eldanolide), vanillin, and 4-hydroxybenzaldehyde, identified by French workers in the wing gland and abdominal hair pencil secretions of the male African sugarcane borer,Eldana saccharina, we have, in an earlier note, reported the presence of several other terpenoid, aromatic, and unbranched-chain compounds such as, (Z)-3,7-dimethylocta-2,6-dienoic acid, 6,10,14-trimethyl-2-pentadecanol, 4-hydroxy-3-methoxybenzyl alcohol, 1-octadecane thiol, 16-hexadecanolide, and 18-octadecanolide in these secretions. In the present paper experimental details and spectral evidence supporting the identification of these compounds, as well as the identification of (Z)-9-hexadecenal and cw-3,7-di-methyl-6-octen-4-olide (cis-eldanolide), are reported. Using electroantennography it was found that male and female antennae reacted approximately equally strongly to both secretions. This result was confirmed in analyses of the secretions using coupled gas chromatography-electroantennography and it was found that male as well as female antennae responded to eldanolide. Vanillin, substituted phenols related to vanillin, and some oxygenated monoterpenes elicited weak responses in male and female antennae. In some analyses 6,10,14-trimethyl-2-pentadecanol, present in the secretions of the insect, gave a strong antennal response. The results obtained in dynamic and static headspace determinations showed that several of the organic compounds present in the glandular secretions are released in detectable quantities and are present in widely varying quantitative ratios in the effluvia of individual calling male moths.     

Conspecific scent trailing by garter snakes (Thamnophis sirtalis) during autumn Further evidence for use of pheromones in den location

Adult garter snakes (Thamnophis sirtalis), collected in October near a traditional, communal hibernaculum in central Wisconsin, were acclimated to autumnal conditions and subjected to laboratory tests to determine whether they could follow scent trails of a conspecific. Graded responses were obtained, but 75% of the sample showed at least some inclination to follow scent trails. The results suggest that pheromone cues may be used by male and female garter snakes to locate traditional dens during autumnal migrations. Evidence from this and other studies suggests that pheromones are probably used in conjunction with other homing mechanisms and that the role of pheromones in den location may be more important in younger snakes and in populations inhabiting northern latitudes.     

Sex-identifying urine and molt signals in lobster (Homarus americanus)

During courtship, premolt female lobsters,Homarus americanus, choose a male and initiate a pair bond by repeated approaches to his shelter. The male allows such a female to share his shelter for about one week. This knowledge formed the basis to search for quantitative evidence for lobster sex pheromone(s) used in courtship: male cues to allow premolt females to identify a preferred male, and female cues to allow males to identify a premolt mature female. In each of four 1500-liter naturalistic aquaria, the behavioral responses of one female and two male lobsters to male and female lobster urine (0.5 ml) and body odor (20 ml) stimuli were observed. These stimuli were injected once or twice per day into a continuously flowing delivery tube attached to lobster shelters. Habituation to stimulus introduction—a serious problem in earlier experiments—was apparently avoided in the more natural social and physical environment we employed in these experiments. We demonstrated that male and female molt body odors contain different chemical substances: females responded to male molt body odor and males responded to female molt body odor but not vice versa. In general, male and female intermolt urine caused strong responses; however, females responded only weakly to male urine. This suggests that male and female urine are chemically different. Female urine and molt body odor caused a typically male high-on-legs response. These results show that molt body odors and intermolt urine contain sex-specific substances, which may be used in lobster courtship as well as other social interactions.     

Behavioral responses of the parsnip webworm to host plant volatiles

Sixth instars of the parsnip webworm, Depressaria pastinacella, orient by olfaction to bud, male flowers, and female flowers of their primary host plant, Pastinaca sativa. Because octyl acetate and octyl butyrate are characteristic of tissues consumed by the sixth instar, we investigated the influence of these esters on webworm feeding behavior and chemo-orientation. Although octyl acetate and octyl butyrate are feeding deterrents, and octyl butyrate is an olfactory repellent, octyl acetate serves as an olfactory attractant. In olfactometers, webworms do not show a preference when given a choice between octyl acetate and host plant tissues. These findings suggest that octyl acetate is a sufficient cue for olfactory orientation. Such behavior may explain differences in the relative abundance of these esters observed among populations of wild parsnip under differential selection pressure from these insects.     

The Egg Parasitoid <Emphasis Type="Italic">Trissolcus basalis</Emphasis> uses <Emphasis Type="Italic">n</Emphasis>-nonadecane,a Cuticular Hydrocarbon from its Stink Bug Host <Emphasis Type="Italic">Nezara viridula</Emphasis>, to Discriminate Between Female and Male Hosts

Contact kairomones from adult southern green stink bugs, Nezara viridula (L.) (Heteroptera: Pentatomidae) that elicit foraging behavior of the egg parasitoid Trissolcus basalis (Wollaston) were investigated in laboratory experiments. Chemical residues from tarsi and scutella of N. viridula induced foraging by gravid female T. basalis. Residues from body parts of female N. viridula elicited stronger responses than those from the corresponding body parts of males. Deproteinized tarsi still elicited searching responses from wasps, indicating that the kairomone was not proteinaceous. Hexane extracts of host cuticular lipids induced searching responses from T. basalis, with a strong preference for extracts from female hosts. Extracts consisted primarily of linear alkanes from nC₁₉ to nC₃₄, with quantitative and qualitative differences between the sexes. Extracts of female N. viridula contained more nC₂₃, nC₂₄, and nC₂₅ than the corresponding extracts from males, whereas nC₁₉ was detected only in extracts from males. Direct-contact solid phase microextraction (DC-SPME) of N. viridula cuticle and of residues left by adult bugs walking on a glass plate confirmed gender-specific differences in nC₁₉. Trissolcus basalis females responded weakly to a reconstructed blend of the straight-chain hydrocarbons, suggesting that minor components other than linear alkanes must be part of the kairomone. Addition of nC₁₉ to hexane extracts of female N. viridula significantly reduced the wasps’ arrestment responses, similar to wasps’ responses to hexane extracts of male hosts. Overall, our results suggest that a contact kairomone that elicits foraging by T. basalis females is present in the cuticular lipids of N. viridula, and that the presence or absence of nC₁₉ allows T. basalis females to distinguish between residues left by male or female hosts. The ecological significance of these results in the host location behavior of scelionid egg parasitoids is discussed.