Sex pheromone of the large aspen tortrix, Choristoneura conflictana(Lepidoptera: Tortricidae)

Summary. Three components that elicited antennal response from male Choristoneura conflictana were found from female gland extracts analyzed using a coupled gas chromatographic-electroantennographic detector system. The main component in gland extracts was (Z)-11-tetradecenal (Z11-14:Ald). Two minor components also elicited antennal response: (E)11-tetradecenal (E11-14:Ald) and (Z)-11- tetradecen-1-ol (Z11-14:OH). Analysis of effluvia indicated that calling virgin females release mostly Z11-14:Ald and trace amounts of Z11-14:OH. Field and wind tunnel behavioral studies showed that Z11-14:Ald alone attracted male moths in a dose response pattern. Tests comparing male response to blends of components detected in gland extracts showed that addition of 1.8% of E11-14:Ald to Z11-14:Ald did not influence male moths in the wind tunnel, but resulted in significantly lower trap captures in the field. The threecomponent blend [Z11-14:Ald (100), E11-14:Ald (1.8), Z11-14:OH (11)], was less attractive than Z11-14:Ald alone in both field and wind tunnel studies. Traps baited with two virgin female moths were equally attractive to males as traps baited with the three-component synthetic blend but less attractive than traps baited with Z11-14:Ald alone. Field tests of various blends of the two components (Z11-14:Ald, Z11-14:OH) detected in the females’ effluvia showed that the addition of 1–10% Z11-14:OH to Z11-14:Ald did not affect the males’ response to Z11-14:Ald. Our data demonstrate that female C. conflictana release sex pheromone components in a different ratio than they are stored in the pheromone gland. The sex pheromone is comprised of a single component, Z11-14:Ald, that can be used to monitor mated and virgin male C. conflictana throughout their flight period.     

Lipid biochemistry and reproductive biology in two species of Gammaridae (Crustacea: Amphipoda)

Gammarus oceanicus Segerstråle, 1947 and Echinogammarus marinus (Leach, 1815) were sampled during the breeding season from Oslofjord in 1984, and their lipid composition examined in relation to reproductive condition. In G. oceanicus, female lipid content increased as the ovary matured. Both the amount of lipid stored and the rate of accumulation were greater in spring than in winter. Spring eggs contained 12.4 g lipid, of which 63% was triacylglycerol and 27% phospholipid. Both fractions decreased steadily during embryonic development. Winter eggs contained 19.2g lipid, of which 52% was triacylglycerol and 43% phospholipid. During the early stages of embryonic development the amount of phospholipid decreased sharply, whereas that of triacylglycerol increased, suggesting that some of the fatty acid released from phospholipid was sequestered temporarily as triacylglycerol. When newly spawned, both winter and spring eggs were richer in monoenoic fatty acids than adult amphipods and these acids were the major fuel consumed during development. 6 fatty acids were utilised more slowly than 3 acids, and egg carotenoid pigment content remained constant. Female E. marinus increased in lipid content as the ovary matured. Spring eggs contained 14.7 g lipid when newly spawned and this increased to 16.6 g during the early stages of development. This increase was entirely triacylglycerol, which declined in later stages; the source of the extra lipid was unclear. Eggs contained very little phospholipid or sterol, and both of these components remained at a steady low level during development. E. marinus eggs were not significantly rich in thonoenoic acids compared with adults, and saturated, monoenoic and polyenoic acids were utilised about equally during development. Both adults and eggs were rich in 20.46, which was utilised at a slower rate than the 3 polyunsaturated acids during embryonic development; again, egg carotenoid pigment content remained constant. In both species there was a decrease in the size of the egg (and as a result, of the newly hatched juvenile), but an increase in total reproductive output (i.e., the total weight of the egg clutch) per female as the breeding season proceeded. The reproductive output of an individual female is probably related to food availability during the period of ovarian maturation, whereas the size of an individual egg is dictated largely by feeding conditions for the juveniles once they are independent of the female. The different patterns of lipid utilisation during development found in this study emphasize the flexibility of response in the reproductive biology of gammarid amphipods. It is not yet possible, however, to relate the differing patterns in a simple way either to egg size or total female reproductive output. Two outstanding problems are the source of extra triacylglycerol during the early stages of development of E. marinus and the metabolic cost of brooding eggs.     

Reproductive isolation and reproductive output in two sympatric mussel species (Mytilus edulis, M. trossulus) and their hybrids from Newfoundland

The mussels Mytilus edulis L. and M. trossulus Gould are found sympatrically in most areas of Newfoundland, with a low frequency of hybrids. To assess the potential for reproductive isolation, we sampled mussels from three sites in an eastern Newfoundland Bay from May–October 1996 to determine if there were differences in the reproductive cycles of the two species and their natural hybrids. In mussels with sheil lengths of 38–42 mm, males and females with mature gametes were dominant in June for M. edulis and hybrids, while M. trossulus showed a lower frequency of individuals with mature gametes. M. trossulus and hybrids spawned over a prolonged period (from late spring to early autumn) compared with most M. edulis individuals that spawned over a period of 2–3 weeks in July. This asynchrony in spawning activity between the two species may partially explain the low frequency of hybrids found in previous studies of these mussel populations. Female and male hybrids between M. edulis and M. trossulus showed normal gonad development, ripening and spawning, providing an opportunity for the introgression of genes between the two species. M. trossulus had a higher reproductive output than M. edulis of similar shell length, while hybrids showed intermediate values of reproductive output. M. trossulus females produced smaller eggs than either M. edulis or hybrids. Differences in reproductive traits may partially explain the maintenance of the mussel hybrid zone in Newfoundland. Published online: 13 August 2002     

Sex pheromones and their potential role in the evolution of reproductive isolation in small ermine moths (Yponomeutidae)

Summary Sex pheromone communication in the nine European species of small ermine moths (Yponomeuta) is reviewed in regard to the potential role of pheromones in the speciation process. Six of the nine species studied (viz.,Y. evonymellus, Y. cagnagellus, Y. padellus, Y. irrorellus, Y. plumbellus, andY. vigintipunctatus) use a mixture of (E)-11-and (Z)-11-tetradecenyl acetate in different ratios as primary pheromone components, with combinations of tetradecyl acetate, (Z)-9-tetradecenyl acetate, (Z)-11-hexadecenyl acetate and the corresponding alcohols of the acetates as additional pheromone components. Analysis of (Z)- to (E)-11-tetradecenyl acetate ratios produced by individual females of these species demonstrated significant variation among females of all species. However, the ranges of ratios produced byY. cagnagellus, Y. irrorellus, andY. plumbellus, sharing the same host-plant species, spindle tree, did not overlap. Niche separation of all six species mentioned required consideration of at least one additional pheromone component or of temporal aspects. The remaining three species,i.e. Y. malinellus, Y. mahalebellus andY. rorellus, have pheromones that differ qualitatively.Biosynthetic routes to the pheromone components identified are proposed on the basis of fatty acid pheromone precursors found in the pheromone glands. A phylogenetic tree for the genus is constructed based on allozyme frequency data and changes in pheromone composition are superimposed on this tree. We suggest that the ancestral ermine moth pheromone is a mixture of (Z)-11- and (E)-11-tetradecenyl acetate and the corresponding alcohols, and a scenario of how present-day patterns evolved is outlined. The pheromone differences among the three species using spindle tree as their host-plant might have evolved throughreproductive character displacement upon secondary contact between populations that had already diverged genetically in allopatry. Pheromone differences within the so-calledpadellus-complex (includingY. cagnagellus, Y. mahalebellus, Y. malinellus, Y. padellus, andY. rorellus) in which species might have originated sympatrically, may have evolved byreinforcing selection as these species still hybridise and produce viable offspring when confined in cages. The role of pheromones in reproductive isolation amongYponomeuta species is emphasised by (1) the function of pheromone components of some of the species as behavioural antagonists to other species, (2) the cross-attraction under experimental conditions between allochronic species with similar pheromones, and (3) the formation of hybrids in the laboratory between species that are isolated in nature by pheromone differences.     

Sex pheromone components of male <Emphasis Type="Italic">Tirathaba mundella</Emphasis> (Lepidoptera: Pyralidae)

Summary. During peak calling activity by male oil palm bunch moths, Tirathaba mundella Walker (Lepidoptera: Pyralidae), their hairpencils, wings or entire body were extracted in hexane. Gas chromatographic-electroantennographic detection (GC-EAD) analyses of hair pencil extracts revealed four compounds that consistently elicited responses from female antennae. The NMR spectrum of isolated compound 1, and mass spectra and retention indices of compounds 1–4 suggested that they were (3S,6S)-2,2,6-trimethyl-6-vinyl-tetrahydro-pyran-3-ol (1), 4-hydroxy-3-methoxy-benzaldehyde (2, vanillin), 6,10,14-trimethyl-2-pentadecanone (3), and 6,10,14-trimethyl-2-pentadecanol (4). Comparative GC and GC-MS analyses of hair pencils extract and synthetic standards confirmed these structural assignments. Moreover, comparative chromatography of synthetic and hairpencil-isolated 1 on a Cyclodex-B column (which separated the four stereoisomers with baseline resolution) revealed that male T. mundella produce the SS-stereoisomer (SS-1). In field cage bioassay experiments in Palembang, Indonesia, synthetic SS-1 and vanillin in combination, but not singly, attracted female T. mundella. SS-1 plus vanillin were as effective as male T. mundella in attracting females. Compounds 3 and 4 did not enhance the blend's attractiveness. Received October 11 2002; accepted March 14, 2003. R1D=" Correspondence to: Gerhard Gries, email: gries@sfu.ca     

Oviposition-stimulating activity of (E)-capsaicin identified in Capsicum annuum fruit and related compounds towards Helicoverpa assulta (Lepidoptera: Noctuidae)

Summary. The oviposition-stimulating activity of (E)-capsaicin identified in the fruits of red pepper, Capsicum annuum L. (Solanaceae), towards the oriental tobacco budworm, Helicoverpa assulta (Guenée), was examined in the laboratory by using choice and no-choice bioassays. Results were compared with those of the commercially available capsanthin and dihydrocapsaicin, another known constituents of C. annuum fruit, and (Z)-capsaicin. In no-choice oviposition bioassays at 10.5 μg/cm², (E)-capsaicin was the most active oviposition stimulant. Weak stimulatory activity was observed with capsanthin, (Z)-capsaicin and dihydrocapsaicin. In choice oviposition bioassays at 2.6 μg/cm², there were significant differences in oviposition-stimulating activity between (E)-capsaicin and each of the other three alkaloids. Structure-activity relationships indicate that the presence of double bond and its trans configuration appeared to play a crucial role in the oviposition-stimulating activity. On the basis of our results, (E)-capsaicin acts as a contact oviposition stimulant of female H. assulta.     

Sex pheromones and attractants in the Eucosmini and Grapholitini (Lepidoptera,Tortricidae)

Summary The geometric isomers (E,E)-, (E,Z)-, (Z,E)-, and (Z,Z)-8,10-dodecadien-1-yl acetate were identified as sex pheromone components or sex attractants in the tribes Eucosmini and Grapholitini of the tortricid subfamily Olethreutinae. Species belonging to the more ancestral Tortricinae were not attracted. Each one isomer was behaviourally active in males ofCydia andGrapholita (Grapholitini), either as main pheromone compound, attraction synergist or attraction inhibitor. Their reciprocal attractive/antagonistic activity in a number of species enables specific communication with these four compounds.Pammene, as well as otherGrapholita andCydia responded to the monoenic 8- or 10-dodecen-1-yl acetates. Of the tribes Olethreutini and Eucosmini,Hedya, Epiblema, Eucosma, andNotocelia trimaculana were also attracted to 8,10-dodecadien-1-yl acetates, but several otherNotocelia to 10,12-tetradecadien-1-yl acetates. The female sex pheromones ofC. fagiglandana, C. pyrivora, C. splendana, Epiblema foenella andNotocelia roborana were identified. (E,E)- and (E,Z)-8,10-dodecadien-1-yl acetate are producedvia a commonE9 desaturation pathway inC. splendana. CallingC. nigricana andC. fagiglandana females are attracted to wingfanning males.     

A sex pheromone component novel to Ostrinia identified from Ostrinia latipennis (Lepidoptera: Crambidae)

Summary. Extracts from the sex pheromone gland of Ostrinia latipennis (Lepidoptera: Crambidae) were analyzed by gas chromatography-electroantennographic detection (GC-EAD) and GC-mass spectrometry. Only an EAD-active compound was detected in the extract, and it was identified as (E)-11-tetradecenol (E11-14:OH). In a wind-tunnel bioassay, E11-14:OH elicited a series of mate finding behaviors from males, although it was far less active than virgin females and crude extract of the pheromone gland. The attractiveness of E11-14:OH to O. latipennis males was confirmed by field trapping experiments. Based on these findings, we concluded that E11-14:OH, which is novel to the genus Ostrinia, is a major component of the sex pheromone in O. latipennis. The significance of the use of alcohol in place of the usual acetates in Ostrinia is discussed in relation to the pheromone biosynthesis system. Received 9 December 1999; accepted 14 March 2000     

Sex and reproductive aspects in Serranus cabrilla (Osteichthyes: Serranidae): macroscopic and histological approaches

The gonads of Serranus cabrilla (Linnaeus, 1758) from the Canary Islands were studied between September 1992 and July 1993. Macroscopic classification of the maturity stages was compared with histolog ical classification of gonad characteristics, and the effect␣of the classification method on the determination of the spawning season and size at maturity was investigated. Analysis of gonad organization and development confirmed that this species is synchronously hermaphroditic. Ovarian-tissue development is asynchronous, and testicular tissue consists of continuous spermatogonia. Six stages of oogenesis and five of spermatogenesis are described, based on differences in staining and in size and on differences in the nucleus and cytoplasm structure as viewed through a light microscope. Two types of atretic degeneration of eggs are also described. Agreement between macroscopic and histological staging was low, although both methods provided similar results in spawning-season determination and size at maturity. The spawning season of S. cabrilla is from February to July, with a peak in May. Fish size at first maturity (size at which 50% of all fish sampled are in relevant maturity stage) is 152 mm standard length (SL), and size at mass maturity (size at which 95% of all fish sampled are in relevant maturity stage) is 167 mm SL. Received: 21 May 1996 / Accepted: 9 August 1996     

A synomone imparting distinct sex pheromone communication channels for Choristoneura rosaceana (Harris) and Pandemis limitata (Robinson) (Lepidoptera: Tortricidae)

Summary. Male obliquebanded leafrollers, Choristoneura rosaceana (Harris), were induced to respond to a pheromone source tainted with a behavioural antagonist, Z9-tetradecenyl acetate, when a source releasing the antagonist was placed 10 cm upwind of the tainted source in a wind tunnel. However, placement of the antagonist upwind of an attractive pheromone source did not interrupt pheromone-mediated responses. Placement of a source releasing Z9-tetradecenyl acetate, a minor pheromone component of the sympatric species, the threelined leafroller, Pandemis limitata (Robinson), upwind of a calling P. limitata female, reduced conspecific male pheromone-mediated response but resulted in upwind flight by male C. rosaceana and contact with heterospecific females. Male P. limitata locked on and flew upwind to but did not contact heterospecific females when a source releasing Z9-tetradecenyl acetate was positioned upwind of a calling C. rosaceana female. In the field, adaptation or habituation to Z9-tetradecenyl acetate caused by atmospheric treatment with this compound apparently resulted in reciprocal heterospecific pheromone responses. More C. rosaceana males were captured in traps baited with their pheromone and the behavioural antagonist in small field plots treated atmospherically with Z9-tetradecenyl acetate than in nontreated control plots. Fewer male P. limitata were captured in traps baited with their own pheromone, or with C. rosaceana pheromone tainted with Z9-tetradecenyl acetate in plots treated atmospherically with Z9-tetradecenyl acetate than in nontreated control plots. We argue that Z9-tetradecenyl acetate is an important synomone which assists in partitioning the sexual chemical communication channels of C. rosaceana and P. limitata. Received 9 February 1999; accepted 22 March 1999.     

Sex pheromone of the tea aphid,Toxoptera aurantii (Boyer de Fonscolombe) (Hemiptera: Aphididae)

The tea aphid, Toxoptera aurantii, also called the “black citrus aphid”, is one of the most destructive insect pests in commercial tea plantations and gardens in southern China. In autumn, declining day length triggers production of winged T. aurantii sexuparae, which produce both winged males and wingless oviparae. Oviparous females then release sex pheromone that attracts potential mates. GC–MS analysis of volatile headspace extracts of T. aurantii oviparae revealed that they emit (4aS,7S,7aR)-nepetalactone (I) and (1R,4aS,7S,7aR)-nepetalactol (II) in a ratio of 4.3–4.9:1. Field-trapping experiments with synthetic I and II singly or as two-component blends of different doses and ratios showed significant attraction of T. aurantii males, as well as weak attraction of sexuparae. Identification of the T. aurantii sex pheromone provides a new opportunity for developing a pheromone-based monitoring and management strategy for the sexual phase of tea aphids and, possibly, the alate sexparous generation in late summer and fall.     

Sex ratio adjustment and maternal condition in two aphid species

We analyzed how offspring sex ratio varies with maternal condition in order to obtain evidence on the population structure in two aphid species with different life cycles. When fitness returns per unit investment differ for the production of daughters and sons, selection will favor an increasing investment into the sex with the higher returns. Therefore, the offspring sex ratios of individual mothers should become more biased towards the sex with the higher fitness returns as their condition or fecundity improves. The pattern of sex ratio adjustment we found in Uroleucon cirsii indicates local mate competition among males, while the pattern we found in Rhopalosiphum padi suggests local resource competition among sexual females. This might be the first evidence for local resource competition among females in an invertebrate species. Local mate competition means that fitness returns are limited by the availability of females as mates within local breeding groups, whereas local resource competition means that fitness returns are limited by the availability of resources for females competing within local groups. We discuss how the life cycles of both species fit to these hypotheses.

Short run aspects of a genetic control program for Heliothis virescens (Lepidoptera: Noctuidae)

The possibility that suppressing the tobacco budworm, Heliothis virescens (F.), by means of genetic control will lead to short run crop yield sacrifices is examined. A simulation model of budworm/backcross population dynamics is used to investigate the effects of backcross releases which are too small or improperly timed. Apart from its potential for ultimate long-term suppression of the tobacco budworm, adverse short-term effects on crop yields are not likely to be a serious problem with a genetic control program. Furthermore, backcross release appears to have some potential as a substitute for chemical pesticides during the period prior to budworm elimination by genetic control.     

Modified pheromone communication associated with insecticidal resistance in the obliquebanded leafroller, Choristoneura rosaceana (Lepidoptera: Tortricidae)

Summary. This study reports on the impact of insecticidal resistance on the diel periodicity of the calling behaviour and pheromone production of different-aged virgin females of the obliquebanded leafroller (OBL), Choristoneura rosaceana. While both resistant (R) and susceptible (S) females initiated calling on the first night following emergence, the periodicity of the calling behaviour, as determined by the mean onset time of calling (MOTC) and the mean time spent calling (MTSC) over the first six nights of calling, differed between the two strains. R females started calling significantly later in the night. However, as the MOTC of R females advanced with age but did not do so in S individuals, the difference between strains was more pronounced in younger than older females. Furthermore, R females spent less time calling than S individuals. However, the MTSC increased as a function of age in both R and S females, so the difference between strains remained fairly constant for each night of calling. The major component of OBL sex pheromone, the Z11-14: Ac, determined at peak calling activity, significantly declined with female age. Overall, pheromone production was lower in R females than in S females, with the difference being more pronounced in younger than in older individuals. Thus, resistant females may have a lower mating success. The mating success of both R and S strain males did not vary with the number of previous matings acquired. With regard to males, although there was a significant decline in spermatophore size with successive matings, there was no significant difference between strains. However, R males are smaller and may be disadvantaged through female choice and/or may respond differently to pheromone source compared with S individuals. If the reproductive success of both sexes is affected, this may have a profound influence on the dynamics of insecticidal resistance in the presence or absence of selection in OBL populations. Received 4 July 2001; accepted 19 October 2001.     

Identification and field bioassays of the sex pheromone of the yellow-legged clearwing Synanthedon vespiformis (Lepidoptera: Sesiidae)

The yellow-legged clearwing (YLC) Synanthedon vespiformis (Lepidoptera: Sesiidae) occurs in the Mediterranean and central Europe. It is polyphagous, boring into the woody parts of broadleaf species including forest trees as well as various Rosaceae species. S. vespiformis has been reported as an economically important pest causing severe injury to stone fruit plantations. Many attractants for sesiid species were discovered by random field screening using 2,13- and 3,13-octadecadienyl alcohols, acetates and aldehydes, including one for S. vespiformis; and about 20 sex pheromones of sesiids have been identified so far. In the present study we identified the natural composition of the sex pheromone of YLC laboratory reared females as a blend of E3,Z13- and Z3,Z13-octadecadienyl acetates, at a ratio of 4:1. We developed an efficient lure for monitoring the pest. Pheromone funnel traps with rubber septa, impregnated with 1 mg pheromone blend, efficiently captured males for 10 weeks. Suspension of Shin-Etsu^? ropes containing a 2:1 blend of E3,Z13-18:Ac and Z3,Z13-18:Ac at 13.74 mg/ha/h, resulted in shutdown of trap catches in the treated plots and closely situated neighboring plots indicating that mating disruption is feasible.     

Comparative efficacy of certain plant extracts alone and combination with profenofos against Spodoptera littoralis (Boisd.) (Lepidoptera: Noctuidae)

Ethyl alcohol, acetone, and petroleum ether extracts of three plant species belonging to three different botanical families [Strychnos nux-vomica (Loganiaceae), Euphorbia lathyrus (Euphorbiaceae), and Datura stramonlum (Solanaceae)], a chemical insecticide; profenofos and their combinations were tested against second and fourth instars of Spodoptera littoralis under lab conditions. Results revealed that the ethanol extract of S. nux-vomica was the most effective among all plant extracts, where the corrected mortality% were 92, 81, 58, and 27% to 2nd instar and 89, 74, 34, and 11% to 4th instar at concentrations 0.5, 0.25, 0.125, and 0.0625%, respectively. Calculated LC₅₀'s were 0.11, 0.22, and 0.34% to 2nd instar and 0.17, 0.37, and 0.52% to 4th instar for ethanol, petroleum ether, and acetone extracts, respectively. Acetone extracts of all plants were of lower effect. The chemical insecticide profenofos displayed higher efficacy than plant extracts (LC₅₀ = 0.002 and 0.003% for S. littoralis 2nd and 4th larval instars, respectively). The co-toxicity factor reached 76 and 60 when mixing S. nux-vomica + profenofos and D. stramonlum + profenofos at ratio 1:1 against S. littoralis 2nd instar larvae, thus indicating a potentiating effect. While treatment of the 4th instar larvae by the same mixtures resulted in a co-toxicity factor below 20 at all mixing ratios indicating, only, an additive effect against this instar.     

Green leaf volatiles enhance sex attractant pheromone of the tobacco budworm,Heliothis virescens (Lep.: Noctuidae)

Summary Components of the green leaf volatile complex (Z-3-hexenyl acetate andE-2-hexenyl acetate) were shown to enhance responses of tobacco budworm,Heliothis virescens, males to the sex attractant pheromone of conspecific females in the field. The results are discussed with regard to green leaf volatiles which enhance the attractant pheromone of a cohabiting species, and serve as attractants of a parasitoid of conspecific larvae.