Identification of a Sex Attractant Pheromone for Male Winterform Pear Psylla, <Emphasis Type="Italic">Cacopsylla pyricola</Emphasis>

Pear psylla, Cacopsylla pyricola (Förster) (Hemiptera: Psyllidae), a major economic pest of pears, uses a female-produced sex attractant pheromone. We compared the chemical profiles obtained from cuticular extracts of diapausing and post-diapause winterform males and females to isolate and identify the pheromone. Post-diapause females produced significantly more of the cuticular hydrocarbon, 13-methylheptacosane, than post-diapause males and diapausing females. In olfactometer assays, conspecific males were attracted to synthetic racemic 13-methylheptacosane, whereas females were not, indicating that the behavioral response to this chemical is sex-specific. Furthermore, 13-methylheptacosane was as attractive to males as a cuticular extract of females, suggesting that this chemical was largely responsible for the female attractiveness. A field study showed that males but not females were attracted to 13-methylheptacosane, confirming the olfactometer results. This study provides evidence that 13-methylheptacosane is a sex attractant pheromone for C. pyricola winterform males. This is the first identification of a sex pheromone in the Psylloidea. Our results open the path to developing monitoring tools and possibly new strategies for integrated pest management of this insect.     

Unsaturated Cuticular Hydrocarbons Enhance Responses to Sex Pheromone in Spruce Budworm, <Emphasis Type="Italic">Choristoneura fumiferana</Emphasis>

The primary sex pheromone components of the female spruce budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae), are (E)- and (Z)-11-tetradecenal, produced in 95:5 ratio. However, male flight responses to calling females in a wind tunnel were faster and maintained longer than responses to any synthetic aldehyde blend. Analyses of cuticular extracts from spruce budworm adults revealed series of n-alkanes and n-monoalkenes with predominantly odd numbers of carbon atoms from C₂₃- C₂₉ in both sexes. (Z,Z,Z)-3,6,9-tricosatriene and (Z,Z,Z)-3,6,9-pentacosatriene were identified only in cuticular extracts from females. Pheromonally naïve males showed wing fanning and circling responses to forewing scales from females but not to scales from males. Males also exhibited the same strong responses to scales excised from pharate females, indicating that the pheromone components are produced by females prior to emergence. (Z)-11-hexadecenal and (Z)-5-tricosene enhanced male responses to the primary sex pheromone aldehydes in wind tunnel bioassays, including higher proportions of in-flight and copulatory responses by males and increased time on the source. Addition of (Z,Z,Z)-3,6,9-tricosatriene to the 95/5 blend of (E)- and (Z)-11-tetradecenal released close-range copulatory responses including abdomen curling on treated septa. We propose that the sex pheromone blend of C. fumiferana is composed of the 95/5 blend of (E)- and (Z)-11-tetradecenal as primary components, with (Z)-11-hexadecenal, (Z)-5-tricosene and (Z,Z,Z)-3,6,9-tricosatriene fulfilling secondary roles in orientation and close-range courtship.     

Multiple Roles of a Male-Specific Compound in the Sexual Behavior of the Dried Bean Beetle, <Emphasis Type="Italic">Acanthoscelides Obtectus</Emphasis>

Males of Acanthoscelides obtectus (Coleoptera: Chrysomelidae, Bruchinae) emit methyl (E,R)-2,4,5-tetradecatrienoate that attracts females for mating. This study identified further roles for this compound in the sexual behavior of A. obtectus. Earlier observations revealed that males touched females with their antennae while tandem-running with them and initiated mounting and copulation, whereas they showed no such behavior toward other males. A series of subsequent laboratory choice tests were set up to establish if certain cuticular compounds aid contact sex recognition in A. obtectus. Males chose virgin females over other males. The activity toward females could be eliminated by rinsing with hexane, but was regained by application of female extract onto previously rinsed females. Gas chromatographic (GC) comparison of hexane extracts revealed the presence of two male-specific compounds, methyl (E,R)-2,4,5-tetradecatrienoate and octadecanal, which were absent from the behaviorally active female samples. Of the two compounds, methyl (E,R)-2,4,5-tetradecatrienoate was found to be responsible for the inhibition of male sexual behavior, similar to that observed with crude male extracts applied to virgin females. Furthermore, males preferred virgin over mated females. GC analyses revealed the presence of methyl (E,R)-2,4,5-tetradecatrienoate in mated females in amounts sufficient to curtail mating attempts. It appears that methyl (E,R)-2,4,5-tetradecatrienoate, besides being a male-produced sex pheromone, acts as a male-recognition signal in A. obtectus. Males also transfer it onto females during mating, resulting in mated females being avoided by courting males.     

Identification of Host Blends that Attract the African Invasive Fruit Fly,Bactrocera invadens

Bactrocera invadens, an invasive fruit fly species in the Afro-tropical region belonging to the Bactrocera dorsalis complex, causes considerable damage to fruit production and productivity. We sought to find attractants from hosts of B. invadens that could serve as baits in traps for monitoring and management of this pest. The attractiveness of volatiles from four different fruit species (mango, guava, banana and orange) at two stages of ripeness (ripe or unripe) was tested in an olfactometer assay. All fruits were attractive against a clean air control. Using hexane extracts of volatile collections of fruits, we demonstrated that male flies preferred the volatiles of ripe guava and orange over unripe fruit extracts. There was a slight difference in preference between females and males; females preferred orange to guava and mango, whereas males preferred mango and guava to orange. Gas chromatography/electroantennographic detection (GC/EAD) and GC/mass spectrometry (GC/MS) were used to identify compounds to which B. invadens antennae were sensitive. GC/EAD recordings from distal and medio-central parts of the fly antenna showed responses to a number of compounds from each fruit species, with esters dominating the responses. Synthetic blends were made for each fruit species using the shared antennally active compounds in ratios found in the extracts. In the olfactometer, B. invadens was most attracted to the banana and orange blends, followed by the mango and guava blends. The synthetic banana blend was as attractive as the volatile collection of banana, although both were less attractive than the fruit. The results demonstrate that composing attractive blends from GC/EAD-active constituents shared by host fruits can be effective for formulating attractive synthetic host mimics for generalist fruit fly species, such as B. invadens.     

Beyond Cuticular Hydrocarbons: Chemically Mediated Mate Recognition in the Subsocial Burying Beetle <Emphasis Type="Italic">Nicrophorus vespilloides</Emphasis>

Burying beetles have fascinated scientists for centuries due to their elaborate form of biparental care that includes the burial and defense of a vertebrate carcass, as well as the subsequent feeding of the larvae. However, besides extensive research on burying beetles, one fundamental question has yet to be answered: what cues do males use to discriminate between the sexes? Here, we show in the burying beetle Nicrophorus vespilloides that cuticular lipids trigger male mating behavior. Previous chemical analyses have revealed sex differences in cuticular hydrocarbon (CHC) composition; however, in the current study, fractionated-guided bioassay showed that cuticular lipids, other than CHCs, elicit copulation. Chemical analyses of the behaviorally active fraction revealed 17 compounds, mainly aldehydes and fatty acid esters, with small quantitative but no qualitative differences between the sexes. Supplementation of males with hexadecanal, the compound contributing most to the statistical separation of the chemical profiles of males and females, did not trigger copulation attempts by males. Therefore, a possible explanation is that the whole profile of polar lipids mediates sex recognition in N. vespilloides.     

Volatile and Contact Chemical Cues Associated with Host and Mate Recognition Behavior of <Emphasis Type="Italic">Sphenophorus venatus</Emphasis> and <Emphasis Type="Italic">Sphenophorus parvulus</Emphasis> (Coleoptera: Dryophthoridae)

Beetles in the genus Sphenophorus Schönherr, or billbugs, potentially utilize both volatile and non-volatile behavior-modifying chemical signals. These insects are widely distributed across North America, often occurring in multi-species assemblages in grasses. However, details about their host- and mate-finding behavior are poorly understood. This study tested the hypothesis that volatile organic compounds from host-plants and conspecifics direct the dispersal behavior of hunting billbug S. venatus Say. Further, we characterized the cuticular hydrocarbon profiles of two widespread pest species, S. venatus and bluegrass billbug S. parvulus Gyllenhaal, to assess the potential role of contact pheromones in mate-recognition. In Y-tube olfactometer bioassays, S. venatus males were attracted to a combination of conspecifics and Cynodon dactylon host-plant material, as well as C. dactylon plant material alone. S. venatus females were attracted to a combination of male conspecifics and host-plants but were also attracted to male conspecifics alone. Field evaluation of a putative male-produced aggregation pheromone, 2-methyl-4-octanol, identified from two congeners, S. levis Vaurie and S. incurrens Gyllenhaal, did not support the hypothesis that S. venatus and S. parvulus were also attracted to this compound. Gas chromatography-mass spectrometry analysis of S. venatus and S. parvulus whole-body cuticular extracts indicated a series of hydrocarbons with qualitative and quantitative interspecific variation in addition to intraspecific quantitative variation between males and females. This study provides the first evidence that S. venatus orients toward host- and insect-derived volatile organic compounds and substantiates the presence of species-specific cuticular hydrocarbons that could serve as contact pheromones for sympatric Sphenophorus species.     

Stingless Bees: Chemical Differences and Potential Functions in <Emphasis Type="Italic">Nannotrigona testaceicornis</Emphasis> and <Emphasis Type="Italic">Plebeia droryana</Emphasis> Males and Workers

Cuticular wax, abdominal and cephalic extracts of foraging workers and males of Nannotrigona testaceicornis and Plebeia droryana, from the “Aretuzina” farm in São Simão, SP, Brazil, were analyzed by GC-MS. The principal constituents were hydrocarbons, terpenes, aldehydes, esters, steroids, alcohols, and fatty acids. Interspecific differences for both cuticular wax and cephalic extracts were found. The composition of cuticular wax and cephalic extracts was similar at the intraspecific level, with minor component differences between males and workers. Abdominal extracts differentiated sexes (male and worker) at the intraspecific and interspecific levels. The main chemical components in abdominal extracts of N. testaceicornis workers and males were geranylgeranyl acetate and (Z)-9-nonacosene, respectively. The principal components of abdominal extracts from P. droryana workers and males were tetradecanal and unsaturated fatty acids (linoleic and linolenic acids), respectively. A secondary alcohol, (S)-2-nonanol, was detected in Plebeia droryana males only, but not in workers. Preliminary field experiments showed that (S)-(+)-2-heptanol and (S)-(+)-2-heptanol/ (S)-(+)-2-nonanol (1:1) attracted workers of P. droryana, N. testaceicornis, and Frieseomelitta silvestrii. However, males did not respond suggesting that these compounds do not function as alarm or recruitment pheromones . In addition, racemic mixtures were inactive.     

Experience-Induced Habituation and Preference Towards Non-Host Plant Odors in Ovipositing Females of a Moth

In phytophagous insects, experience can increase positive responses towards non-host plant extracts or induce oviposition on non-host plants, but the underlying chemical and behavioral mechanisms are poorly understood. By using the diamondback moth, Plutella xylostella, its host plant Chinese cabbage, and a non-host plant Chrysanthemum morifolium, as a model system, we observed the experience-altered olfactory responses of ovipositing females towards volatiles of the non-host plant, volatiles of pure chemicals (p-cymene and α-terpinene) found in the non-host plant, and volatiles of host plants treated with these chemicals. We assessed the experience-altered oviposition preference towards host plants treated with p-cymene. Naive females showed aversion to the odors of the non-host plant, the pure chemicals, and the pure chemical-treated host plants. In contrast, experienced females either became attracted by these non-host odors or were no longer repelled by these odors. Similarly, naive females laid a significantly lower proportion of eggs on pure chemical-treated host plants than on untreated host plants, but experienced females laid a similar or higher proportion of eggs on pure chemical-treated host plants compared to untreated host plants. Chemical analysis indicated that application of the non-host pure chemicals on Chinese cabbage induced emissions of volatiles by this host plant. We conclude that induced preference for previously repellent compounds is a major mechanism that leads to behavioral changes of this moth towards non-host plants or their extracts.     

Identification of Sex Pheromones and Sex Pheromone Mimics for Two North American Click Beetle Species (Coleoptera: Elateridae) in the Genus <Emphasis Type="Italic">Cardiophorus</Emphasis> Esch.

To date, all known or suspected pheromones of click beetles (Coleoptera: Elateridae) have been identified solely from species native to Europe and Asia; reports of identifications from North American species dating from the 1970s have since proven to be incorrect. While conducting bioassays of pheromones of a longhorned beetle (Coleoptera: Cerambycidae), we serendipitously discovered that males of Cardiophorus tenebrosus L. and Cardiophorus edwardsi Horn were specifically attracted to the cerambycid pheromone fuscumol acetate, (E)-6,10-dimethylundeca-5,9-dien-2-yl acetate, suggesting that this compound might also be a sex pheromone for the two Cardiophorus species. Further field bioassays and electrophysiological assays with the enantiomers of fuscumol acetate determined that males were specifically attracted by the (R)-enantiomer. However, subsequent analyses of extracts of volatiles from female C. tenebrosus and C. edwardsi showed that the females actually produced a different compound, which was identified as (3R,6E)-3,7,11-trimethyl-6,10-dodecadienoic acid methyl ester (methyl (3R,6E)-2,3-dihydrofarnesoate). In field trials, both the racemate and the (R)-enantiomer of the pheromone attracted similar numbers of male beetles, suggesting that the (S)-enantiomer was not interfering with responses to the insect-produced (R)-enantiomer. This report constitutes the first conclusive identification of sex pheromones for any North American click beetle species. Possible reasons for the strong and specific attraction of males to fuscumol acetate, which is markedly different in structure to the actual pheromone, are discussed.     

Abstinons

Glossina morsitans morsitans Westwood andMusca domestica Linne have an efficient strategy for preventing wasteful homosexual activity in males: hexane-soluble substances on the cuticle of the male flies terminate courting by other males on contact. The inhibitory activity of these materials is demonstrated by the abstention of males from mating with females treated with male extract.     

Influence of Host-Plant Surface Chemicals on the Oviposition of the Cereal Stemborer Busseola Fusca

The chemical composition of plant surfaces plays a role in selection of host plants by herbivorous insects. Once the insect reaches the plant, these cues determine host acceptance. Laboratory studies have shown that the stem borer Busseola fusca (Lepidoptera: Noctuidae), an important pest of sorghum and maize in sub-Saharan Africa, is able to differentiate between host and non-host plant species. However, no information is available on the cues used by this insect to seek and accept the host plant. Thus, the role of surface phytochemical stimuli on host selection and oviposition by B. fusca was studied in the laboratory using two host plants, sorghum, Sorghum bicolor, and maize, Zea mays, and one non-host plant, Napier grass, Pennisetum purpureum. The numbers of eggs and egg masses deposited on the three plant species were compared first under no-choice and choice conditions. In both cases, more eggs and egg masses were laid on maize and sorghum than on the non-host. Artificial surrogate stems treated with a water or chloroform surface extract of each plant were then compared with surrogate stems treated with, respectively, water or chloroform as controls, under similar conditions. Surrogate stems treated with plant water extracts did not show an increase in oviposition when compared to controls, indicating that the major compounds in these extracts, i.e., simple sugars and free amino acids, are not significantly responsible for the oviposition preference. By contrast, a chloroform extract of sorghum enhanced oviposition on the surrogate stems compared to the control, while those of maize and Napier grass showed no significant effects. Analysis of the chloroform extract of sorghum showed higher amounts of α-amyrin, ß-amyrin, and n-nonacosane compared to those of maize and Napier grass. A blend of the three chemicals significantly increased oviposition compared to the chloroform-treated control, indicating that these compounds are part of the surface chemical signature of the plant responsible for host recognition and oviposition by B. fusca.     

Modulation of Reproductive Behaviors by Non-Host Volatiles in the Polyphagous Egyptian Cotton Leafworm, Spodoptera littoralis

In order to locate mates, food, and oviposition sites, insects mainly rely on volatile cues released by their sexual partners, food sources, and host and non-host plants. Calling, mating, and oviposition behaviors, as well as fecundity and longevity, of newly emerged Spodoptera littoralis (Bois.) moths were recorded in the presence of volatiles from leaves of a host plant, Gossypium hirsutum (cotton) and two non-host plants, Adhatoda vasica (Av) or Picea abies (spruce), either alone or in host/non-host combinations. Females exposed to cotton volatiles started calling earlier than females exposed to non-host plant volatiles (NHV), or the blank control. Likewise, moth pairs exposed to cotton volatiles started mating earlier than the other treatments. The period of calling in females alone was longer than females kept with males, having the opportunity to mate. However, the callings, as well as mating durations in the moth pairs, in different treatments were not different. Longevity was decreased either in the absence of cotton or the presence of Av, and spruce leaves. Fecundity was reduced in moths exposed to a combination of spruce and cotton. The effect of NHV on attraction of 2-3–day-old male moths towards a pheromone (Ph) source was studied in a wind tunnel. In the no-choice assay, more males arrived at close approach and landed on the Ph source when the host plant, cotton, was offered in the background as compared to the non-hosts. In the dual-choice assay, more males landed on the Ph source in front of the host plant compared to the Ph source in front of non-hosts. Gas chromatography-electroantennographic detection on female S. littoralis revealed five antennally active compounds in headspace collections of spruce and three compounds in Av.     

Olfaction in the boll weevil,Anthonomus grandis Boh. (Coleoptera: Curculionidae): Electroantennogram studies

Electroantennogram (EAG) techniques were utilized to measure the antennal olfactory responsiveness of adult boll weevils,Anthonomus grandis Boh. (Coleoptera: Curculionidae), to 38 odorants, including both insect and host plant (Gossypium hirsutum L.) volatiles. EAGs of both sexes were indicative of at least two receptor populations: one receptor population primarily responsive to pheromone components and related compounds, the other receptor population primarily responsive to plant odors. Similar responses to male aggregation pheromone components (i.e., compounds I, II, and III + IV) were obtained from both sexes, but females were slightly more sensitive to I. Both sexes were highly responsive to components of the “green leaf volatile complex,” especially the six-carbon saturated and monounsaturated primary alcohols. Heptanal was the most active aldehyde tested. More acceptors responded to oxygenated monoterpenes than to monoterpene hydrocarbons. β-Bisabolol, the major volatile of cotton, was the most active sesquiterpene. In general, males, which are responsible for host selection and pheromone production, were more sensitive to plant odors than were females. In fact, males were as sensitive to β-bisabolol and heptanal as to aggregation pheromone components. Electrophysiological data are discussed with regard to the role of insect and host plant volatiles in host selection and aggregation behavior of the boll weevil.     

Biochemical adaptations of the bruchid beetle,Caryedes brasiliensis tol-canavanine,a higher plant allelochemical

This communication reviews recent biochemical studies ofl-canavanine that have provided significant understanding of the interaction between the seed ofDioclea megacarpa (Leguminosae) and the bruchid beetleCaryedes brasiliensis (Bruchidae). The principal biochemical bases are proposed for: canavanine toxicity, the ability of the beetle larvae to adapt to its presence, the metabolic sequestration and detoxification of ammonia, and the potential amplification by microbial symbionts of the beetle's abilities to adapt to toxic components of the host.     

Behavioral Evidence for Olfactory-Based Location of Honeybee Colonies by the Scarab <Emphasis Type="Italic">Oplostomus haroldi</Emphasis>

The Afro-tropical scarab Oplostomus haroldi (Witte) is a pest of honeybees in East Africa with little information available on its chemical ecology. Recently, we identified a female-produced contact sex pheromone, (Z)-9-pentacosene, from the cuticular lipids that attracted males. Here, we investigated the kairomonal basis of host location in O. haroldi. We used coupled gas chromatography/electroantennographic detection (GC/EAD) and GC/mass spectrometry to identify antennally-active compounds from volatiles collected from honeybee colonies. Antennae of both sexes of the beetle consistently detected seven components, which were identified as 3-hydroxy-2-butanone, 2,3-butanediol, butyl acetate, isopentyl acetate, butyl butyrate, hexyl acetate, and methyl benzoate. In olfactometer bioassays, both sexes responded to the full seven-component synthetic blend over solvent controls, but chose honeybee colony odors over the blend. These findings suggest that the seven compounds are components of a kairomone from honeybee colonies used by O. haroldi.     

Selected factors influencing neurohormonal regulation of sex pheromone production inHeliothis species

Sex pheromone production and release in females ofHeliothis species exhibit a diel periodicity. Phermone production is controlled by a hormone, the pheromone biosynthesis activating neuropeptide (PBAN). Release of PBAN to activate pheromone production follows a circadian rhythm. InH. zea females, mating terminates pheromone production. An unidentified hemolymph-borne factor is transferred from the male to the female during mating. It is speculated that this factor interacts with the release mechanism of PBAN to prevent further production of the pheromone following mating. Wild females ofH. phloxiphaga (reared from larvae collected in the field) did not produce or release the sex pheromone unless kept in association with the host plant. Pheromone production could be induced in these females by the injection of PBAN. It is suggested that a signal from the host plant is essential to trigger the release of PBAN to induce pheromone production.     

Characterization of a Female-Produced Courtship Pheromone in the Parasitoid Nasonia vitripennis

Males of the parasitoid Nasonia vitripennis (Walker) (Hymenoptera: Pteromalidae) show a characteristic courtship behavior. We demonstrate that male arrestment and key behavioral elements of the courtship sequence are mediated by a female-derived contact sex pheromone. Males were arrested on paper disks treated with female extracts but not on those treated with male extracts. Male responsiveness was influenced by the surface to which female extracts were applied. Extracts applied to an extracted beetle elytron arrested males more strongly than those applied to filter paper of comparable size. However, more complex behavioral elements, such as head nodding and copulation attempts, were shown only when extracts were applied to extracted male cadavers, suggesting that tactile or visual cues synergize the male response. The chemicals involved are stable, of low volatility, and nonpolar. Dead females arrested males and elicited courtship behavior for at least 8 d. Males showed no sign of attraction to live females at a distance of 3 cm in an olfactometer. Fractionation of female extracts demonstrated that the activity was exclusively located in the nonpolar fraction. Analysis of the active fraction by gas chromatography–mass spectrometry revealed that cuticular hydrocarbons with chain lengths between 25 and 37 carbon units were present. Comparison of hydrocarbon profiles from males and females showed qualitative and quantitative differences. These results suggest that sex-specific cuticular hydrocarbons are the key signals mediating the male courtship behavior in N. vitripennis.     

Cuticular Hydrocarbons of <Emphasis Type="Italic">Tribolium confusum</Emphasis> Larvae Mediate Trail Following and Host Recognition in the Ectoparasitoid <Emphasis Type="Italic">Holepyris sylvanidis</Emphasis>

Parasitic wasps which attack insects infesting processed stored food need to locate their hosts hidden inside these products. Their host search is well-known to be guided by host kairomones, perceived via olfaction or contact. Among contact kairomones, host cuticular hydrocarbons (CHCs) may provide reliable information for a parasitoid. However, the chemistry of CHC profiles of hosts living in processed stored food products is largely unknown. Here we showed that the ectoparasitoid Holepyris sylvanidis uses CHCs of its host Tribolium confusum, a worldwide stored product pest, as kairomones for host location and recognition at short range. Chemical analysis of T. confusum larval extracts by gas chromatography coupled with mass spectrometry revealed a rich blend of long-chain (C25-C30) hydrocarbons, including n-alkanes, mono-, and dimethylalkanes. We further studied whether host larvae leave sufficient CHCs on a substrate where they walk along, thus allowing parasitoids to perceive a CHC trail and follow it to their host larvae. We detected 18 CHCs on a substrate that had been exposed to host larvae. These compounds were also found in crude extracts of host larvae and made up about a fifth of the CHC amount extracted. Behavioral assays showed that trails of host CHCs were followed by the parasitoids and reduced their searching time until successful host recognition. Host CHC trails deposited on different substrates were persistent for about a day. Hence, the parasitoid H. sylvanidis exploits CHCs of T. confusum larvae for host finding by following host CHC trails and for host recognition by direct contact with host larvae.     

Chemical Communication in the Honey Bee Scarab Pest Oplostomus haroldi: Role of (Z)-9-Pentacosene

Oplostomus haroldi Witte belongs to a unique genus of afro-tropical scarabs that have associations with honey bee colonies, from which they derive vital nutrients. Although the attributes of the honey bee nest impose barriers to communication among nest invaders, this beetle still is able to detect conspecific mates for reproduction. Here, we show, through behavioral studies, that cuticular lipids serve as mate discrimination cues in this beetle. We observed five steps during mating: arrestment, alignment, mounting, and copulation, and a post-copulatory stage, lasting ~40–70 % of the total mating duration, that suggested mate guarding. Chemical analysis identified the same nine straight-chain alkanes (C₂₃–C₃₁), six methyl-branched alkanes (6), and five mono-unsaturated alkenes in the cuticular lipids of both sexes. Methyl alkanes constituted the major component (46 %) of male cuticular lipids, while mono-unsaturated alkenes were most abundant (53 %) in females. (Z)-9-Pentacosene was twice as abundant in females than in males, and ~20 fold more concentrated in beetles than in worker bees. In mating assays, (Z)-9-pentacosene elicited arrestment, alignment, and mounting, but not copulation, by male beetles. These results represent the first evidence of a contact sex pheromone in a scarab beetle. Such contact pheromones may be an essential, cryptic mechanism for arthropods associated with eusocial insects.     

Mites as Matchmakers: Semiochemicals from Host-associated Mites Attract Both Sexes of the Parasitoid Lariophagus distinguendus

The role of volatile chemicals used for mate finding was studied for males of Lariophagus distinguendus (Först.), a parasitoid of the granary weevil Sitophilus granarius (L.). In bioassays that used a static four-chamber olfactometer, males were attracted by host feces, hexane extracts from host feces, and volatile extracts of the feces obtained by closed-loop stripping (CLS). On the other hand, volatiles emitted by unmated females did not elicit any preferences in males. Both sexes of L. distinguendus responded to a synthetic mixture of neral, geranial, neryl formate, and tridecane occurring in the investigated extracts. All compounds are common constituents of astigmatid mites that are often associated with possible hosts of L. distinguendus. In the system investigated, all main compounds found in CLS extracts from larval feces of S. granarius are due to the mold mite Tyrophagus putrescentiae (Schrank) that uses neral, geranial, and neryl formate as an alarm pheromone. The possible role of host-associated astigmatid mites in mate and host finding of L. distinguendus is discussed.