Sex pheromone of Nearctic Agriotes mancus and its similarity to that of three Palearctic Agriotes invasive in North America

1. The wheat wireworm, Agriotes mancus (Coleoptera: Elateridae), is a predominant elaterid pest species in the Nearctic region, with a life history and morphology similar to those of Agriotes obscurus, Agriotes lineatus and Agriotes sputator, three Palearctic pest elaterids invasive in North America. Here, we report the identification and field testing of the sex pheromone of A. mancus.
2. We collected headspace volatiles from female beetles on Porapak Q, and analysed aliquots of Porapak extract by gas chromatography with electroantennographic detection (GC-EAD) and by GC-mass spectrometry. In GC-EAD recordings, two esters—geranyl butanoate and geranyl hexanoate—elicited antennal responses from A. mancus males. In field experiments, trap lures containing both geranyl butanoate and geranyl hexanoate afforded large captures of A. mancus males, which were—on average—approximately 30-fold higher than captures in traps baited with a single ester.
3. Traps baited with geranyl butanoate as a single-component lure captured a significant number of Palearctic A. sputator, indicating the establishment of A. sputator in its invaded Nearctic range.
4. With the A. mancus sex pheromone now known, it can be included in the development of pheromone-based programmes to monitor and manage native and invasive Agriotes pests in North America.

     

Small‐plot studies comparing pheromone and juice baits for mass‐trapping invasive Synanthedon myopaeformis in Canada

Recent introduction of Synanthedon myopaeformis (Borkhausen) (Lepidoptera: Sesiidae) into organic apple‐growing areas of Canada has stimulated research on semiochemical‐based management of this European pest. Replicated, small‐plot (0.16 ha) experiments were conducted to compare sex pheromone, 3Z,13Z‐octadecadienyl acetate (10 mg), Concord grape juice (300 ml), or their combination, as mass‐trapping lures at trap densities equivalent to 12.5, 25, 50, and 100 traps ha^?1. Total numbers of male and female moths removed from test plots increased significantly with trap density in all juice‐based mass‐trapping experiments. In pheromone mass‐trapping experiments, however, total catches of males did not increase significantly as trap densities were increased and catches appeared to plateau with 25–50 traps ha^?1. With pheromone‐based mass‐trapping, significantly fewer males were caught in pheromone‐baited assessment traps at the centre of each mass‐trapping plot than in identical traps in untreated plots. This reduction is indicative of significant trap interference or trap ‘shut‐down’. Increasing the density of juice‐based mass‐trapping had no effect on catches of male or female moths in juice‐baited assessment traps, indicating a short range of attraction and lack of interference between juice traps. Pheromone‐ and juice‐based mass trapping removed similar numbers of males at each trap density tested, respectively, but summed catches of males and females were greatest with juice baits. Combining pheromone and juice into a single mass‐trapping treatment (50 traps ha^?1) did not significantly increase catches of males or females relative to either treatment alone. If a practical bisexual mass‐trapping system is going to be developed for S. myopaeformis, then identification of volatile kairomones in Concord grape juice may be useful.     

Antennal and behavioural responses of the spruce seed moth,Cydia strobilella,to floral volatiles of Norway spruce,Picea abies,and temporal variation in emission of active compounds

We investigated whether spruce seed moth, Cydia strobilella L. (Lepidoptera: Tortricidae, Grapholitini), one of the most damaging seed predators on Norway spruce, Picea abies (L.) H. Karst (Pinaceae), uses olfactory cues during host search. Analyses with coupled gas chromatography and electroantennography revealed that antennae of both sexes of moths responded consistently to three compounds in the headspace from female spruce flowers, i.e., α‐pinene, β‐pinene, and myrcene, but not to limonene as has been previously reported for this species. The amounts of these active volatiles released from flowers and cones of P. abies were quantified, and their diurnal and seasonal variation was monitored. The total release of the active volatiles correlated well with the diurnal and seasonal flight activity of C. strobilella as revealed by catches of males in pheromone‐baited traps. In field trapping experiments, where baits were loaded with proportions and enantiomeric ratios of α‐pinene, β‐pinene, and myrcene matching those of the female P. abies floral headspace, substantial catches of male C. strobilella were achieved, whereas few females were captured. These surprising results suggest that male C. strobilella make use of host volatiles to aid them in their search for females.     

Evolutionary and climatic factors affecting tooth size in the red foxVulpes vulpes in the Holarctic

Research into the geographical pattern of tooth size in the red fox,Vulpes vulpes (Linnaeus, 1758) in the Holarctic was conducted on a sample of 3806 skulls belonging to 41 fox populations. The Nearctic was represented by 948 specimens (249 females, 359 males, 340 specimens of unknown sex) belonging to 13 populations, whereas the Palearctic was represented by 2858 red foxes (1034 females, 1256 males, 568 specimens of unknown sex) from 32 populations. In the Nearctic, the largest foxes live on Kodiak Island (V. v. harrimani) and the Kenai Peninsula (V. v. kenaiensis), while the smallest ones live in California (V. v. necator) and Georgia (V. v. fulvus). In the Palearctic, the largest foxes come from the Far East (V. v. jakutensis, V. v. beringiana, V. v. tobolica), while the smallest are from the southern borders of the Eurasian range (V. v. pusilla, V. v. barbara, V. v. arabica). In both the Palearctic and Nearctic, tooth size in the fox varies depending on the geo-climatic factors. The fox’s tooth size confirms the general basis of Bergmann’s rule. In the Palearctic, specimens with larger teeth occur in cooler habitats with greater seasonality. These are first and foremost Northern and Far Eastern populations. In the Nearctic, tooth size in red foxes depends on the temperature and humidity of their habitat. Competition within the species and between species has important impact on the variation and dimorphism of tooth size in the red fox. Both in the Nearctic and Palearctic, red foxes from regions of sympatric co-occurrence with other closely relatedVulpes species, are more sexually dimorphic in terms of tooth size than red foxes from allopatric regions. Analysis of morphological distance on the basis of the size of dental characteristics shows, that in the Palearctic, the foxes from India (V. v. pusilla), while in the Nearctic, the population from Kodiak Island (V. v. harrimani) are most distant from the remaining populations. Geographic barriers such as the Bering Strait, Parry Channel, Mackenzie River, Kolyma and Omolon River systems have had a critical impact on red fox evolution. The most likely place for the evolution and diversification of the phyletic lineVulpes vulpes seems to be the Middle East region.     

Differentiation of chromosome banding sequence pools and genomic dna in Holarctic natural populations of <Emphasis Type="Italic">Chironomus entis</Emphasis> Shobanov (Diptera,Chironomidae)

Polymorphism and differentiation of the chromosome banding sequence pools and genomic DNA were studied in three natural populations of Chironomus entis from Europe and North America. These populations showed a moderate level of chromosomal polymorphism and high RAPD polymorphism of genomic DNA. The Palearctic and Nearctic populations of this species did not differ significantly in the levels of chromosome and genomic DNA polymorphism. Estimation of the cytogenetic (GD_cg) and genetic (GD_DNA) distances between these C. entis populations showed that their chromosome banding sequence pools and cytogenetic structures are differentiated to a greater extent than genomic DNA. The values of cytogenetic and genetic distances between the Palearctic and Nearctic populations of C. entis are higher than the values of the corresponding distances between the Nearctic populations, but they do not reach the level of divergence between species.     

Sexual differences in the flight performance of the oriental fruit moth, Cydia molesta

Cydia molesta Busck (Lepidoptera: Tortricidae), a major pest of stone fruits and an increasingly important late season pest of apple, is predominately monitored by pheromone trapping of male moths. We investigated flight performance in relation to sex, age, and mating status using computer-linked flight mills, and also examined the relationship between female flight and reproduction. The crepuscular flight pattern of the experimental moths in relation to photointensity was very similar to that reported from field studies. Female moths significantly outperformed males in all measured flight parameters including total distance flown, distance of longest single flight, and velocity. The proportion of long-flying females (categorised as those completing an unbroken flight of greater than 1 km) was three to six times greater than that of males. Female flight performance was not related to mating status, but mated males displayed significantly greater flight than unmated males. The maximal flight period of mated females commenced on the third day after eclosion, following 30% egg deposition. Male flight was not significantly related to age. The data suggest that a limited proportion of the population, in particular females, may have the capacity to make inter-orchard flights. The limitations of monitoring C. molesta populations in apple orchards solely by pheromone trapping of males are discussed.     

Identification and field testing of sex pheromone components of a Korean population of the allium leafminer,Acrolepiopsis sapporensis

The allium leafminer, Acrolepiopsis sapporensis Matsumura (Lepidoptera: Acrolepiidae), is a pest of Allium species (Liliaceae) in Asia and Hawaii, USA. We identified candidate sex pheromone components in pheromone gland extracts of female moths and field tested the response of male moths to blends with different components and ratios. Gas chromatographic comparison of abdominal tip extracts from both sexes showed three female‐specific components: (Z)‐11‐hexadecenal (Z11‐16:Ald), (Z)‐11‐hexadecenyl acetate (Z11‐16:OAc), and (Z)‐11‐hexadecen‐1‐ol (Z11‐16:OH). These compounds were identified by mass spectral analysis of natural pheromone components and dimethyldisulfide adducts, and retention index comparisons with synthetic standards. The average ratio of three components, Z11‐16:Ald, Z11‐16:OAc, and Z11‐16:OH, in female extract was 33:100:14. Field trapping experiments indicated that all three components were essential for maximal attraction of male moths. Traps baited with a ternary blend mimicking the blend found in the pheromone gland extracts caught significantly more males than traps baited with caged live females. Increasing doses of the pheromone blend in the lures from 0.01 to 1.0 mg increased catches of male A. sapporensis.     

Comparison of mating disruption and mass trapping with Pyralidae and Sesiidae moths

Mating disruption and mass trapping for control of lepidopteran pests use synthetic sex pheromone to prevent males from finding and mating with females. Here, we identify the behavioral mechanism underlying mating disruption and mass trapping of American plum borer, Euzophera semifuneralis (Walker) (Lepidoptera: Pyralidae), peachtree borer, Synanthedon exitiosa Say, and lesser peachtree borer, Synanthedon pictipes (Groeten) (Lepidoptera: Sesiidae). In addition, we derive relative dispenser activity (Relative D_a) from the competitive attraction equation to compare the disruptive activity of the devices used in mating disruption and mass trapping. Dispensers and traps were deployed in replicated 0.14‐ha cherry or peach plots with E. semifuneralis or the Synanthedon moths, respectively. Dispenser densities were 0, 10, 20, 59, 185, and 371 per ha, whereas trap densities were 0, 10, 20, 40, 79, and 158 per ha. Moth catch in a centrally placed, pheromone‐baited monitoring trap in each plot was used to evaluate the treatments. The profile of moth captures in mating disruption and mass trapping with the three species indicates that competitive attraction is the behavioral mechanism responsible for trap disruption. Relative D_a is 0.27, 0.23, and 0.53 with American plum borer, peachtree borer, and lesser peachtree borer, respectively, which indicates that the traps are 1.9–4.4 times more effective in reducing moth catch than the dispensers. Relative D_a can be used to compare devices for pheromone‐based behavioral manipulation of these and other species that are competitively attracted to artificial pheromone sources. When the same type of trap is employed for monitoring and mass trapping, Relative D_a is the same as dispenser activity D_a.     

褛裳夜蛾的交配行为及雄蛾对性腺提取物的反应节律

详细观察了褛裳夜蛾成虫的交配行为,利用风洞、触角电位技术研究了雄蛾对性腺提取物的反应节律,通过林间诱蛾试验进行了验证,旨在为褛裳夜蛾性信息素的精确提取及性信息素组份分离、鉴定提供依据。研究结果表明:成虫的求偶、交配行为均发生在暗期,有一定的节律性:雌蛾在羽化3d以后开始求偶,1周左右表现最为强烈,3-4日龄雌蛾在暗期4-6h开始求偶,最大求偶率在暗期6-8h;5-9日龄在进入暗期就开始求偶,最大求偶率出现在暗期4-6h。风洞试验表明,3-7日龄的雄蛾对性腺提取物均有明显的性行为反应,5日龄雄蛾在暗期4-6h对性腺提取物的反应最为强烈。3-6日龄的成虫在暗期4-6h开始交配,而7-8日龄的成虫于暗期2-4h开始交配,6日龄的成虫交配率最高,交配高峰在暗期4-8h。雄蛾对性腺提取物的触角电位反应也有一定的节律性:雄蛾对4日龄雌蛾性腺提取物开始有电生理反应,对6-7日龄暗期5h性腺提取物反应最为强烈。林间诱蛾试验测定了性腺提取物的引诱活性,7日龄雌蛾性腺提取物林间诱蛾量最高,引诱高峰在暗期4-6h,该结果也验证了褛裳夜蛾雄蛾对性腺提取物的反应节律。     

稻纵卷叶螟对性信息素和花香的嗅觉行为反应及在其种群监测中的应用

[目的]明确稻纵卷叶螟Cnaphalocrocis medinalis生理状态对性信息素和花香气味的嗅觉行为反应的影响,以及在田间性信息素和花香气味诱蛾量的动态.[方法]于2020年在广西、湖南、浙江、四川5个试验点,通过田间网捕、花香气味和性信息素诱捕稻纵卷叶螟成虫,结合解剖卵巢和精巢,比较性信息素和花香诱捕的基本迁...     

Intercontinental karyotypic differentiation of Chironomus entis Shobanov, a Holarctic member of the C. plumosus group (Diptera, Chironomidae).

Analysis of banding sequences of polytene chromosomes in Palearctic (Russian) and Nearctic (North American) Chironomus entis shows strong karyotype divergence between populations on the two continents. Four out of seven chromosomal arms in the North American C. entis karyotype are characterized by sequences found only in the Nearctic. In total, 44 banding sequences are now known for this species across the Holarctic, including 22 exclusively Palearctic, 6 Holarctic, and 16 exclusively Nearctic sequences. The degree of cytogenetic differentiation between Palearctic and Nearctic C. entis populations is an order of magnitude greater than differentiation among populations within either continent, but is only one third as great as the cytogenetic distance between the sibling species C. entis and C. plumosus. C. entis is the only sibling species of C. plumosus uncovered during cytological identification of Chironomus species from more than 50 North American lakes, indicating that the plumosus sibling-species group is much smaller in the Nearctic than in the Palearctic, where a dozen sibling species are known. Cytogenetic distance values calculated between Nearctic and Palearctic representatives of both C. entis and its sibling species C. plumosus are similar, but result from different patterns of karyotype divergence. New World C. entis is distinguished from Old World populations by the 16 uniquely Nearctic sequences, four of which occur in the homozygous state. In contrast, North American C. plumosus has fewer uniquely Nearctic sequences, and only one that occurs as a homozygote. However, four chromosomal arms in C. plumosus that are polymorphic in the Palearctic show fixation, or near fixation, of Holarctic sequences in the Nearctic C. plumosus karyotype. Thus, both the fixation of Holarctic sequences, and the occurrence or fixation of distinctly Nearctic sequences, contribute significantly to karyotype divergence. Patterns of karyotype divergence in Palearctic and Nearctic populations of different Holarctic chironomid species are discussed relative to intercontinental cytogenetic differentiation in other dipterans.     

Sex pheromone of the cloaked pug moth,Eupithecia abietaria (Lepidoptera: Geometridae), a pest of spruce cones

The sex pheromone of the cloaked pug moth, Eupithecia abietaria Götze, an important cone‐feeding pest in spruce seed orchards in Europe, was investigated. Chemical and electrophysiological analyses of pheromone gland extracts of female moths and analogous analyses of synthetic hydrocarbons and epoxides of chain length C₁₉ and C₂₁ revealed (3Z,6Z,9Z)‐3,6,9‐nonadecatriene (3Z,6Z,9Z‐19:H) and 3Z,6Z‐cis‐9,10‐epoxynonadecadiene (3Z,6Z‐cis‐9,10‐epoxy‐19:H) as candidate pheromone components, which were found in a gland extract in a ratio of 95 : 5. In field trapping experiments, conspecific males were only attracted to a combination of 3Z,6Z,9Z‐19:H and the (9S,10R)‐enantiomer of 3Z,6Z‐cis‐9,10‐epoxy‐19:H. The (9R,10S)‐enantiomer was not attractive, which is in agreement with studies on other Eupithecia species, for which males have only been attracted by the (9S,10R)‐enantiomer of epoxides. Subsequent experiments showed that E. abietaria males were attracted to a wide range of ratios of the two active compounds and that trap catches increased with increasing dose of the binary blend. A two‐component bait containing 300 μg 3Z,6Z,9Z‐19:H and 33 μg of the (9S,10R)‐enantiomer of 3Z,6Z‐cis‐9,10‐epoxy‐19:H was efficient for monitoring E. abietaria in spruce seed orchards in southern Sweden, where this species has probably been overlooked as an important pest in the past. With sex pheromones recently identified for two other moths that are major pests on spruce cones, the spruce seed moth, Cydia strobilella L., and the spruce coneworm, Dioryctria abietella Denis & Schiffermüller, pheromone‐based monitoring can now be achieved for the whole guild of cone‐feeding moths in European spruce seed orchards.     

Ovarian development of Agrilus planipennis: effects of age and mating status and influence on attraction to host volatiles

Emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), is a major pest of ash trees, Fraxinus spp. (Oleaceae), in North America. This study investigated the timing of reproductive development in female beetles and the influence of female reproductive maturity on attraction to host volatiles. Based on dissections of females of increasing age, females with access to males for mating, and thus presumed mated, developed mature eggs only after 18–24 days. In contrast, female beetles reared without access to males, and thus unmated, did not develop mature eggs at any age. Chemical analysis of cuticular hydrocarbons detected the contact sex pheromone, 9‐methyl‐pentacosane, in cohorts of females which were 8–9 days old and older, supporting previous research that this compound signals sexual maturity to males. Results from field‐trapping bioassays demonstrated that stage of female reproductive maturity influenced their attraction to host volatiles: females caught on traps baited with foliar volatiles contained eggs and ovarioles that were significantly less developed than those on traps baited with bark sesquiterpenes. However, our results revealed that females with immature stages of ovarioles and undeveloped eggs, such as those observed in unmated females, were rarely ever caught on traps baited with either of the host volatile lures. Further research on host compounds attractive to immature females is critical for early detection and possible control of A. planipennis populations during the extended pre‐oviposition period.     

Effect of sex pheromone emission on the attraction of Lobesia botrana

Since the discovery of Lobesia botrana Denis & Schiffermüller (Lepidoptera: Tortricidae) sex pheromone, it has played an important role in the control and detection of this pest, for example, through the use of pheromone‐baited traps and mating disruption techniques. Rubber septa are the most common pheromone dispensers used in monitoring traps, but often dispenser performance is not optimized. The key to improve methods based on pheromones as attractants (monitoring, mass trapping, or ‘attract and kill’) is to know the optimum emission interval, because release rates can strongly affect the attraction. In this work, five levels of pheromone load with different release rates were compared in traps using mesoporous pheromone dispensers to investigate the optimum release rate maximizing L. botrana catches. Residual pheromone loads of the dispensers were extracted and quantified by gas chromatography, to study release profiles and to estimate the various emission levels. The efficacy of pheromone emission was measured in field trials as number of moths caught. A quadratic model was fitted to relate the numbers caught vs. the daily emission rates. The resulting quadratic term was statistically significant, confirming the existence of a relative maximum for L. botrana catches. Taking into account that the trial was carried out only in one location, an optimum emission value of ca. 400 μg per day could be considered to enhance the attraction of L. botrana under West‐Mediterranean weather conditions.     

n-Butyl (E)-7,9-decadienoate: sex pheromone component of the nettle caterpillar, Darna pallivitta

The nettle caterpillar, Darna pallivitta (Moore) (Lepidoptera: Limacodidae), is an invasive pest on the island of Hawai’i, causing defoliation of ornamental nursery stock and posing a human health hazard due to their urticating hairs that can cause painful stings. Wind tunnel and field tests with 2–3‐day‐old moths revealed behavioral responses of males to caged females, which is indicative of a female‐released sex pheromone. Coupled gas chromatography‐electroantennogram detection (GC‐EAD) analysis of female abdominal tip extracts revealed two male electroantennographically active compounds produced by female D. pallivitta. Mass spectral analysis and subsequent synthesis identified the active compounds as n‐butyl (E)‐7,9‐decadienoate (major component) and ethyl (E)‐7,9‐decadienoate (minor component), both structurally similar to sex pheromone components previously reported from related Darna spp. Additionally, methyl (E)‐7,9‐decadienoate was identified from female abdominal tip extracts and a strong EAD response was elicited by the synthetic compound. n‐Butyl (E)‐7,9‐decadienoate was the only component detected by solid phase microextraction (SPME) collections from single calling female moths, however, the apparent absence of minor components may be a result of their lower abundance. Field trials showed significant attraction of male moths to all lures containing the n‐butyl ester, while the methyl and ethyl esters did not increase trap captures at the levels and ratios tested. Synthetic pheromone lures (2.5 mg) outperformed virgin moths as attractant baits and could be used for monitoring D. pallivitta populations of the island of Hawai’i and detection on other Hawai’ian islands and at ports and nurseries that receive plants from Hawai’i (e.g., California and Florida).     

Geographic differentiation of genomic DNA of <Emphasis Type="Italic">Chironomus plumosus</Emphasis> (Diptera,Chironomidae) in natural holarctic populations

Using RAPD markers, polymorphism and differentiation of genomic DNA was examined in seven natural populations of Chironomus plumosus from Europe, Siberia, and North America. All these populations showed high polymorphism of genomic DNA. The Palearctic and Nearctic populations of this species were not statistically significantly different in the genomic DNA polymorphism level. The genetic distance (GD), which characterizes the extent of intraspecific differentiation of population genetic structure, was determined among the natural populations of C. plumosus. The genetic distance was on average 0.245. It was demonstrated that genetic structures of the Palearctic and Nearctic populations of C. plumosus was differentiated to a higher extent than in Palearctic. However, the genetic distances between the populations from different zoogeographical zones (0.313) did not exceed the level characteristic of the among-population differences, which do not disturb the species genetic integrity.     

Sampling procedures and adult sex ratios in spruce budworm

Unbiased estimates of sex ratios that reflect local abundance of adult insects are practically difficult to obtain because many gender‐specific behavioural adaptations differentially influence the catchability of males and females in commonly applied sampling procedures. Historic data on outbreak populations of spruce budworm, Choristoneura fumiferana Clemens (Lepidoptera: Tortricidae), the major pest of conifers in Nearctic boreal forests, include dozens of sex ratio observations for 10 different sampling procedures; these data illustrate the importance of understanding the reproductive ecology of adults to contextualize sex ratio assessments. Sex ratios of resident adults (assessed by rearing field‐collected pupae to adulthood or fogging host trees with insecticide) were not different from 1:1. Sex ratios of in‐flight adults collected using Malaise traps or light traps deployed in tree canopies were consistently male‐biased, which presumably reflects the higher level of flight activity for males relative to females. Sex ratios of moths captured outside the forest canopy (presumed migrants), in contrast, were consistently female‐biased, a trend which is expected because females seeking oviposition sites are more likely to undergo migration than are males. The sex ratio among adults that died from natural causes (collected on drop trays) was not distinguishable from 1:1. In pre‐outbreak (endemic) populations, sex ratios estimated by light trapping were much more strongly male‐biased than in outbreak populations. This surprising result should, however, be interpreted with caution because little is known of reproductive ecology in endemic budworm populations.     

Determination of Agriotes obscurus (Coleoptera: Elateridae) sex pheromone attraction range using target male behavioural responses

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A sex pheromone component novel to the Geometridae identified fromEpirrita autumnata

(Z,Z,Z)-1,3,6,9-Heneicosatetraene was identified as a sex pheromone component ofEpirrita autumnata (Borkhausen), (Lepidoptera: Geometridae). Gas chromatography coupled with electroantennographic detection revealed one active peak from female pheromone gland extracts. The chemical identification of the compound was based on gas chromatography-mass spectrometry, chemical micro-reactions and unambiguous synthesis. In a field test, a rubber septum loaded with 3000 μg of this compound showed highest attraction of maleE. autumnata moths compared to lower doses and the control. The attraction of males to a 300 μg bait was equivalent to that obtained using live virgin females.     

Bionomics of Prays citri (Lepidoptera: Yponomeutidae) and their use in a model of control by male mass trapping*

Abstract. 1. The paper describes the population dynamics of Prays citri (Milliere), relevant to the control of this pest by synthetic pheromone, as well as infestation of lemon trees in various areas and seasons. 2. The responses of males to natural and synthetic sex pheromones are compared, and ratios for the degree of attraction of traps compared with the degree of attraction of competing wild females are calculated. 3. The ratios obtained with various population densities over two seasons of the pest's activity are presented. 4. The data are used in a model simulating control of Prays citri by male mass trapping.