Monitoring codling moth (Lepidoptera: Tortricidae) with a four‐component volatile blend compared to a sex pheromone‐based blend

Monitoring adult codling moth, Cydia pomonella (L.), is a crucial component in implementing effective integrated management programmes in apple, Malus domestica Borkhausen. Use of sex pheromone lures to track male populations has been the traditional approach, but their use in orchards treated with sex pheromone for mating disruption (MD) has been problematic. Development of kairomone and kairomone–pheromone combination lures has allowed the catch of female moths and has benefited several aspects of codling moth management through improved spray timings and action thresholds. Recently, a new four‐component volatile blend (4‐K) comprised of pear ester, (E,Z)‐2,4‐ethyl decadienoate (PE), (E)‐11 4,8‐dimethyl‐1,3,7‐nonatriene, all isomers of pyranoid linalool oxide and acetic acid (AA) has been characterized that has increased female moth catch threefold versus any previous blend. Field trapping studies were conducted to compare moth catches in traps baited with 4‐K versus the use of sex pheromone, (E,E)‐8,10‐dodecadien‐1‐ol (PH) in combination with PE and AA. Trials were conducted in orchards left either untreated, or treated with PH or PH + PE. Traps baited with 4‐K and 4‐K + PH lures caught significantly more females than traps baited with PH + PE + AA lures. Traps baited with 4‐K + PH lures caught significantly more total moths than traps baited with PH + PE + AA lures in all three orchards. Adding a PH lure to traps with the 4‐K lure did not affect female catch, but significantly increased male and total moth catches. These studies demonstrate that codling moth can be trapped effectively in apple under MD without the use of sex pheromone lures. The significant increase in female codling moth catch with the 4‐K lure suggests that efforts to improve spray timings and action threshold determinations as well as mass trapping might be enhanced with this new lure.     

Sex pheromone of the alfalfa plant bug,Adelphocoris lineolatus

In northern China, due to the large‐scale adoption of transgenic Bacillus thuringiensis Berliner (Bt) cotton, the number of Adelphocoris lineolatus (Goeze) (Hemiptera: Miridae) has increased quickly, causing significant loss in cotton and alfalfa production. One of the environmentally safe strategies without use of pesticides is the application of insect pheromone for mating disruption. In our study, we aim to identify the active components in sexually mature virgin female A. lineolatus, and activity and optimal ratio of these components. By using gas chromatography–mass spectrometry (GC‐MS) and gas chromatography–electroantennographic detection (GC‐EAD), we identified three active compounds: hexyl butyrate (HB), (E)‐2‐hexenyl butyrate (E2HB), and (E)‐4‐oxo‐2‐hexenal (4‐OHE). We examined the release rate of septum and tube dispensers in a wind tunnel. In a field experiment, deletion of 4‐OHE or E2HB resulted in significant suppression of male trap catches, indicating that E2HB or 4‐OHE may be the active pheromone components. Traps baited with a blend of 4‐OHE and E2HB at 2:5, 3:4, and 4:3 caught significantly more males, suggesting that the optimal ratio of 4‐OHE and E2HB could be in the range of 1:1 to 1:2. The addition of a large amount of HB could strongly reduce the attractive activity of both virgin females and artificial lures. We also compared the attractiveness of septum lures and tube lures in field experiments. The septum lures attracted males in the first 3 days. The tube lures caught more males after 3 days and the attractive effects existed up to 5 weeks, suggesting them as a tool for long‐term monitoring and control of plant bugs.     

Mating disruption of Lasioderma serricorne (Coleoptera: Anobiidae) in stored product habitats using the synthetic pheromone serricornin

Cigarette beetles, Lasioderma serricorne (F.) (Coleoptera: Anobiidae), cause significant damage to the multibillion dollar food and tobacco industries worldwide each year. A non‐insecticidal alternative to manage L. serricorne is the application of mating disruption, in which high levels of synthetic sex pheromone are released to create an atmosphere that results in males failing to mate females, thereby causing population suppression or extinction. The reported work used synthetic serricornin, the predominant sex pheromone of L. serricorne, in mating disruption trials conducted in selected food‐ and feed‐processing facilities in South Carolina during 2010 and 2011. Mills subjected to mating disruption trials were monitored using oviposition cups filled with larval food and pheromone traps for males that contained monitoring lures. Immediately after deployment of mating disruption dispensers, trap captures declined significantly and indicated a reduction in population levels, that is, there was ‘trap shutdown’. A significant reduction was observed in numbers of adult beetles caught in the traps 8 weeks before and 8 weeks after treatment in both years. Beetle numbers from pheromone traps in untreated buildings remained at similar levels or increased after the time of mating disruption deployment in treated buildings. The numbers of adults that emerged from oviposition food cups were generally low and varied irregularly in treated and untreated buildings and were determined to be of little value for assessing treatment effects on reproduction. These initial field studies in the USA suggest that release of the synthetic sex pheromone of L. serricorne for mating disruption can significantly inhibit proper orientation behaviour of male L. serricorne to females and may lead to pest population decline from mating disruption.     

SPLAT-OrB reveals competitive attraction as a mechanism of mating disruption in oriental beetle (Coleoptera: Scarabaeidae)

This study compared the efficacy of SPLAT-OrB, a new pheromone formulation for oriental beetle mating disruption that can be mechanically applied, with hand-applied plastic dispensers in commercial blueberry fields. Both formulations were tested at 2.5 and 5 g of the major sex pheromone component (Z)-7-tetradecen-2-one per hectare, and evaluated by measuring trap shut-down, mating success of caged females, and the number of grubs in sentinel blueberry pots baited with tethered females. All pheromone-treated plots had fewer male captures in traps and lower mating success of caged females compared with untreated plots. SPLAT-OrB, and plastic dispensers at the higher rate, also reduced the number of grubs in sentinel pots. To understand the mechanism of mating disruption in oriental beetle, males were observed approaching the pheromone sources in disrupted plots. In addition, male oriental beetle captures were quantified in plots treated with varying SPLAT-OrB dollop densities per ha. Consistent with predictions for competitive attraction, field observations revealed males approaching the pheromone source and male captures decreasing concavely with increasing dollop density. In a mark-release-recapture study, male oriental beetles responded to SPLAT-OrB dollops and plastic dispensers at least 60 m from the source. Additionally, SPLAT-OrB emitted pheromone that was attractive to male oriental beetles for >5 wk; however, emission rates and attraction dropped rapidly during the first 3-4 wk. This study demonstrates the feasibility of using SPLAT-OrB for oriental beetle mating disruption as an alternative to hand-applied plastic dispensers, and conclusively reveals that a principal mechanism is the competitive attraction of males.     

Attractiveness and competitiveness of irradiated light brown apple moths

The sterile insect technique (SIT) potentially provides a socially acceptable approach for insect eradication of new pest incursions. The light brown apple moth, Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae), was discovered in Berkeley (CA, USA) in 2006, leading to an incursion response that included this technology. In this study, we assessed factors affecting mating success from a bisex release of irradiated moths: effects of radiation dose on male multiple mating, male flight competition, female sex pheromone titre and attractiveness of irradiated females to males, and identification of successful mating in vineyards of either irradiated or wild males (identified by isotope analysis of spermatophores from sentinel females). There was a significant negative relationship between male radiation dose and mating frequency. In head‐to‐head flights of irradiated males against non‐irradiated males to a pheromone lure in a wind tunnel, irradiated males reached the lure first only 31% of the time. With increasing radiation dose, the production of the major sex pheromone component in females, (E)‐11‐tetradecenyl acetate, dropped, from 0.7 ± 0.1 ng per female in non‐irradiated females to 0.2 ± 0.07 ng per female when irradiated at 300 Gy. Male catch was reduced to 11% of control females in traps containing females irradiated at 300 Gy. Isotope analysis of spermatophores found in the bursa copulatrix of females indicated that mating success of irradiated males inside the live (entry‐only) traps containing virgin females was lower (13.1 ± 3.3%) than suggested by male catch (21.2 ± 3.8%) in pheromone traps, the current standard for assessing field competitiveness. Impacts of irradiation on male and female moth fitness should be taken into account to improve estimates of irradiated to wild male E. postvittana overflooding ratios needed for population suppression.     

Pheromone release rate determines whether sexual communication of Oriental fruit moth is disrupted competitively vs. non‐competitively

Communicational disruption mechanisms for Oriental fruit moth, Grapholita molesta (Busck) (Lepidoptera: Tortricidae), were determined using a suite of mathematical tools and graphical operations that enable differentiation between competitive attraction and non‐competitive mechanisms of disruption. Research was conducted in 20 field cages, each covering 12 mature apple trees. Commercial monitoring lures releasing Oriental fruit moth pheromone at a rate of 0.04 μg h^?1 and distributed at densities of 0, 1, 2, 4, 8, and 17 per cage were used to evaluate the effect of low‐releasing dispensers on the disruption of sexual communication. Graphical analyses revealed that near‐female‐equivalent pheromone dispensers disrupted Oriental fruit moth competitively. Commercial mating disruption dispensers releasing Oriental fruit moth pheromone at 60 μg h^?1 and deployed at 0, 4, 6, 10, 15, 20, and 30 per cage were used to evaluate the effect of high‐releasing dispensers on the disruption of sexual communication. Oriental fruit moth disruption shifted to a non‐competitive mechanism for high‐releasing dispensers. This is the first time such a shift in disruption mechanism has been demonstrated against a background of otherwise identical experimental conditions. Near‐female‐equivalent pheromone dispensers were also used to quantify the additive effect of an attract‐and‐remove control strategy compared with competitive mating disruption. We report a five‐fold reduction in Oriental fruit moth captures under attract‐and‐remove compared to mating disruption using near‐female‐equivalent dispensers. Surprisingly, release of female Oriental fruit moths into these large‐cage disruption studies had no measurable impact on male trapping.     

Silk‐ and volatile‐based male mate choice in the genital plug‐producing spider

Female mating history can have a strong effect on male fertilization success. Although males often prefer to mate with virgin females, they often also engage with mated females. As the intensity of sperm competition can differ among mated females, males are expected to evolve means to identify their status. In spiders, males often use female silk to gather information about female quality. Males of many spider species deposit mating plugs into female genitalia to hinder further copulations. We tested whether males of the foliage‐dwelling, plug‐producing spider Philodromus cespitum, which is an important natural enemy of pests, discriminate between females of different mating status and whether they can determine the extent of genital plugging in mated females solely on the basis of cues gained from deposited female silk. We presented males with draglines of females that varied in either mating status (virgin vs. mated), the extent of plugging (small vs. big plug), or the age of the plug (fresh vs. old plug) and examined their mate preferences. Additionally, we tested whether males were attracted to volatile cues produced by female bodies. Our experiments revealed that males preferred draglines of virgin females to those of mated females, and mated females with small plugs to those with large plugs. They were also attracted to female volatile cues. This study suggests that males are able to extract fine‐scale information on mating status from female draglines.     

Context‐ and State‐Dependent Male Mate Choice in a Sexually Cannibalistic Spider

High male mating investment may favor selection on male mate choice particularly if females vary in quality. Terminal investment strategies constitute a maximal mating effort and have evolved independently in the absence of paternal investment in several spider taxa including the genus Argiope. To test for male mate preferences in the above context, we used the sexually cannibalistic spider A. bruennichi. We varied male state (mating status and post‐maturation age) as well as the competitive context and quantified male mate choice decisions between females of different states and developmental stages in binary choice tests. We found an overall adaptive preference for the virgin against the mated female regardless of male mating state. Furthermore, we demonstrated that older males paid more attention to female fecundity‐related traits than to mating status. In a second set of experiments, we offered males a choice between a virgin and a subadult female and varied the competitive context which had no effect on male decisions. Curiously, a preference for the virgin adult female was only apparent after exclusion of females that matured <3 d prior to the test. Repeated tests of males supported the hypothesis that males do not distinguish between a freshly matured virgin female and a subadult female. Our results show that male spiders execute mate choice based on information collected from female silk strands and that they integrate their own state into mating decisions.     

Genotypes and their interaction effects on reproduction and mating‐induced immune activation in Drosophila melanogaster

Mating causes considerable alterations in female physiology and behaviour, and immune gene expression, partly due to proteins transferred from males to females during copulation. The magnitude of these phenotypic changes could be driven by the genotypes of males and females, as well as their interaction. To test this, we carried out a series of genotype‐by‐genotype (G × G) experiments using Drosophila melanogaster populations from two distant geographical locations. We expected lines to have diverged in male reproductive traits and females to differ in their responses to these traits. We examined female physiological and behavioural post‐mating responses to male mating traits, that is behaviour and ejaculate composition, in the short to mid‐term (48 hr) following mating. We then explored whether a sexually transferred molecule, sex peptide (SP), is the mechanism behind our observed female post‐mating responses. Our results show that the genotypes of both sexes as well as the interaction between male and female genotypes affect mating and post‐mating reproductive traits. Immune gene expression of three candidate genes increased in response to mating and was genotype‐dependent but did not show a G × G signature. Males showed genotype‐dependent SP expression in the 7 days following eclosion, but female genotypes showed no differential sensitivity to the receipt of SP. The two genotypes demonstrated clear divergence in physiological traits in short‐ to mid‐term responses to mating, but the longer‐term consequences of these initial dynamics remain to be uncovered.     

A female‐produced short‐range sex pheromone in the egg parasitoid Trissolcus brochymenae

The potential for short‐range sex pheromone communication by the egg parasitoid wasp Trissolcus brochymenae (Hymenoptera: Platygastridae) was investigated in closed arena bioassays. Males of this parasitoid showed more antennal drumming and more frequent mounting behaviour on 1‐ to 2‐d‐old virgin females compared with 8‐d‐old virgin females. Male copulation attempts were fewer with previously mated females than with virgin females. Males courted and made copulation attempts with 1‐ to 2‐d‐old female cadavers, but not with male cadavers or with female cadavers rinsed in organic solvents of different polarities. Male attraction to female cadavers was re‐established by treating cadavers with acetone extracts of females, but not with ether or hexane extracts. In experiments using female cadavers dissected into head, mesosoma, and gaster, and then reassembled using one unwashed body section and two body sections washed in acetone, males were attracted only to the reassembled cadavers with an unwashed mesosoma. These findings suggest that (1) courtship behaviour in males of T. brochymenae is triggered by a short‐range sex pheromone produced by females; (2) the age and the physiological condition of females (virgin/mated) influence pheromone release or production; (3) the female's mesosoma is the source of the sex pheromone; and (4) polar components of the sex pheromone play a major role in influencing male behaviour. Our results suggest that quasi‐gregarious egg parasitoids are selected for short‐range rather than long‐range sex pheromones.     

Mating disruption of the ambrosia beetle Megaplatypus mutatus in poplar and hazelnut plantations using reservoir systems for pheromones

Megaplatypus mutatus (Chapuis) (Coleoptera: Curculionidae: Platypodinae) is an ambrosia beetle native to South America, but it has recently been introduced into Italy and represents a serious problem in commercial poplar and fruit tree plantations. Male M. mutatus emit a sex pheromone composed of (+)‐6‐methyl‐5‐hepten‐2‐ol [(+)‐sulcatol], 6‐methyl‐5‐hepten‐2‐one (sulcatone), and 3‐pentanol. We performed three field trials of mating disruption of M. mutatus in hazelnut and poplar plantations in Argentina and Italy. To perform these tests, we made plastic pheromone reservoir‐type dispensers for sulcatol, sulcatone, and 3‐pentanol with zero‐order kinetics that were deployed in the field during the female flight period. The number of galleries where mating took place was significantly higher in control than in treated areas, indicating that pheromone application had interfered with female behavior and male localization. Because damage reduction was >56% in both countries, these results demonstrate the potential for the strategy of pheromone‐mediated mating disruption of M. mutatus in commercial poplar and hazelnut plantations. Also, our study provides the first evidence for successful pheromone‐mediated mating disruption in a forest beetle.     

Allyl‐2,6‐dimethoxyphenol,a female‐biased compound,is robustly attractive to conspecific males of Bactrocera dorsalis at close range

Insect cuticular hydrocarbons (CHCs) play important roles in mate recognition and chemical communication. To explore the cues regulating courtship and mating behaviour in the oriental fruit fly, Bactrocera dorsalis (Hendel) (Diptera: Tephritidae), CHCs were extracted from mature virgin female and male oriental fruit flies using n‐hexane. Nine compounds – i.e., 4‐allyl‐2,6‐dimethoxyphenol (designated ‘compound 3’) and eight ester compounds – had significantly greater abundance in female samples than in males. Of these nine compounds, six (1, 2, 4, 5, 6, and 8) elicited electrophysiological responses from the antennae of male flies. Compound 3 did not elicit a detectable male antennal response, but in Y‐tube olfactometer bioassays, it exhibited robust attractiveness to conspecific males at close range. We speculate that compound 3 is a close‐range sex pheromone in B. dorsalis. A mating competition experiment revealed that compound 3 significantly increased the competitiveness of males, which implied that compound 3 might be used in control programs of B. dorsalis.     

Effect of anti‐hail nets on Cydia pomonella behavior in apple orchards

Over the last 10 years, the use of nets to protect pome fruit from hailstorms has increased. In this study, we investigated the effect of these nets on the behavior of the codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), a major pest of apple, Malus domestica Borkh. (Rosaceae). Experiments were carried out in net‐covered and uncovered apple orchards treated with conventional insecticides. The number of codling moth males caught in pheromone‐ and virgin female‐baited traps was significantly reduced in net‐covered compared to uncovered plots. In addition, inhibition of mating by the net was demonstrated by significant reductions in mating of tethered virgin females. Fruit injury was consistently lower in net‐covered plots vs. uncovered plots. Because of the reductions in male trap catch, the reduced female mating frequency, and the lower level of damage, we conclude that flat anti‐hail nets have a disruptive effect on the reproductive behavior of the codling moth.     

Potential of high‐density pheromone‐releasing microtraps for control of codling moth Cydia pomonella and obliquebanded leafroller Choristoneura rosaceana

Recent large‐cage studies with codling moth Cydia pomonella (L.) reveal that the removal of moths from an apple orchard using pheromone‐releasing traps is more effective at reducing capture in a central monitoring trap than is a mating disruption protocol without kill/capture. The present study uses open orchard 0.2‐ha plots comparing a high‐density trapping scenario with mating disruption to confirm those results. Two tortricid moth pests of tree fruit are studied: codling moth and obliquebanded leafroller Choristoneura rosaceana (Harris). Codling moth treatments include Isomate CM FLEX (ShinEtsu Ltd, Japan), nonsticky traps baited with Trécé CM lures (Trécé, Inc., Adair, Oklahoma), and sticky traps baited with Trécé CM lures, all at equal application rates of 500 dispensers ha^?1, as well as a no pheromone control. These microtraps are of a novel design, small and easy to apply, and potentially inexpensive to produce. Mating disruption using Isomate CM FLEX and nonsticky traps reduces codling moth capture in standard monitoring traps by 58% and 71%, respectively. The attract‐and‐remove treatment with sticky traps reduces capture by 92%. Obliquebanded leafroller treatments include Isomate OBLR/PLR Plus and Pherocon IIB microtraps baited with Trécé OBLR lures, both applied at 500 dispensers ha^?1, as well as a no pheromone control. Mating disruption reduces capture in monitoring traps by 69%. The attract‐and‐remove treatment reduces capture by 85%. Both studies suggest that an attract‐and‐remove approach has the potential to provide superior control of moth populations compared with that achieved by mating disruption operating by competitive attraction.     

Evidence for a female‐produced,long range pheromone of Anoplophora glabripennis (Coleoptera: Cerambycidae)

Abstract Volatiles from female Asian longhorned beetle (ALB), Anoplophora glabripennis, were evaluated as candidate sex pheromone components. Previous studies on ALB have revealed several antennally active compounds from virgin females; however the origins and activity of these compounds were not apparent and require further investigation. We tested the hypothesis that one or more of the ALB contact sex pheromones is a precursor that undergoes abiotic oxidation to yield volatile pheromone components, and evaluated the activity of these compounds using laboratory and field bioassays. Gas chromatography coupled electroantennography detection (GC‐EAD) analysis indicated the presence of three antennally active aldehydes (heptanal, nonanal, and hexadecanal) in female cuticular extracts exposed to ozone or UV and visible light. In laboratory bioassays using a Y‐tube olfactometer, males were preferentially attracted to ozonized female body washes over crude body washes. Similarly, synthetic formulations of these compounds were preferred over controls in the olfactometer. Field trapping experiments conducted from 2006 to 2008 in Ningxia, China showed that synthetic lures of the three aldehydes formulated in a ratio simulating that of virgin females attracted more beetles compared to controls, and that combinations of these aldehydes, linalool oxide, and host kairomones captured more beetles than controls, and captured significantly more males.     

Variability in the efficacy of sex pheromone lures for monitoring oriental fruit moth (Lepidoptera: Tortricidae)

Studies were conducted in Chile and the United States to compare the attractiveness of various commercial sex pheromone lures and two experimental lures for oriental fruit moth, Grapholita molesta (Busck), in peach orchards treated with or without sex pheromone dispensers. The experimental lures contained the three‐component sex pheromone blend of G. molesta: Z‐8‐dodecenyl acetate, E‐8‐dodecenyl acetate and Z‐8‐dodecenol (Z8‐12:OH), and the sex pheromone of codling moth, Cydia pomonella (L.), (E,E)‐8,10‐dodecadien‐1‐ol, (codlemone). Commercial lures varied in their substrate, initial loading and blend ratio of components. Significant differences in male catches were found among commercial lures in orchards treated with or without sex pheromone dispensers. Experimental lures with the addition of codlemone significantly increased the catches of G. molesta using lures loaded with 0%, 1% or 5% Z8‐12:OH in the G. molesta blend compared with the same ratio of components in just the G. molesta blend. The experimental lures were significantly more attractive than all commercial lures in the untreated orchard. However, moth catch with the experimental lures in the sex pheromone‐treated orchard was only intermediate among all of the lures tested. These findings highlight the need to develop more effective and standardized lures that can be used in trap‐based monitoring programme for this important pest.     

Development of 2‐phenylethanol plus acetic acid lures to monitor obliquebanded leafroller (Lepidoptera: Tortricidae) under mating disruption

We evaluated the effectiveness of 2‐phenylethanol (PET) in combination with acetic acid (AA) as a binary lure for monitoring male and female obliquebanded leafroller, Choristoneura rosaceana (Harris). Studies were conducted in apple, Malus domestica Borkhausen, orchards treated with or without sex pheromone dispensers for mating disruption (MD). Open polypropylene vials, closed membrane cups, and rubber septa loaded with AA and/or PET in varying amounts were first evaluated in a series of trapping experiments. Membrane cups loaded with 800 mg of PET were as effective as 10‐mg septa, but longer lasting, and were comparable to the open vials. A membrane cup AA lure was effective in tests, but further work is needed to increase its release rate and extend its activity. Catches of codling moth, Cydia pomonella (L.), and C. rosaceana were unaffected by combining PET with (E,E)‐8,10‐dodecadien‐1‐ol, the sex pheromone of codling moth, pear ester, (E,Z)‐2,4‐ethyl‐decadienoate and AA lures. Adding (E)‐4,8‐dimethyl‐1,3,7‐nonatriene to this blend to enhance codling moth catch significantly reduced catches of C. rosaceana. PET + AA was a more attractive binary lure than AA plus phenylacetonitrile (PAN) for C. rosaceana. The addition of PET or PAN to traps already baited with the sex pheromone of C. rosaceana significantly reduced male catches. Traps baited with PET + AA placed in blocks not treated with MD caught significantly fewer C. rosaceana than traps baited with sex pheromone. In comparison, sex pheromone‐baited traps in MD blocks caught ≤1 male moth per season which was significantly lower than total moth (>10) or female moth (≥3) catch in these blocks with PET + AA. A high proportion (>70%) of trapped females were mated in both untreated and MD‐treated orchards. Further refinement of this binary, bisexual lure using membrane cup technology may allow the establishment of action thresholds and improve management timings for C. rosaceana.     

Old‐male paternity advantage is a function of accumulating sperm and last‐male precedence in a butterfly

Old‐male mating advantage has been convincingly demonstrated in Bicyclus anynana butterflies. This intriguing pattern may be explained by two alternative hypotheses: (i) an increased aggressiveness and persistence of older males during courtship, being caused by the older males' low residual reproductive value; and (ii) an active preference of females towards older males what reflects a good genes hypothesis. Against this background, we here investigate postcopulatory sexual selection by double‐mating Bicyclus anynana females to older and younger males, thus allowing for sperm competition and cryptic mate choice, and by genotyping the resulting offspring. Virgin females were mated with a younger virgin (2–3 days old) and afterwards an older virgin male (12–13 days old) or vice versa. Older males had a higher paternity success than younger ones, but only when being the second (=last) mating partner, while paternity success was equal among older and younger males when older males were the first mating partner. Older males produced larger spermatophores with much higher numbers of fertile sperm than younger males. Thus, we found no evidence for cryptic female mate choice. Rather, the findings reported here seem to result from a combination of last‐male precedence and the number of sperm transferred upon mating, both increasing paternity success.