Oviposition choices by a pre-dispersal seed predator (Hylemya sp.)

Summary Although the importance of pollinators has most often been examined in the evolution of floral characters, seed predators may also play a role in shaping floral evolution. In this study, I examined the role of interplant distance, plant size, and flower morphology on Ipomopsis aggregatás (Polemoniaceae) attractiveness to a pre-dispersal seed predator, Hylemya sp. (Anthomyiidae) and to hummingbird pollinators. The attractiveness of I. aggregata individuals to Hylemya was nonlinearly related to interplant distance in experimental arrays. Clumped and highly dispersed plants were preyed upon more frequently than those at intermediate distances. I found no relationship between interplant distance and visitation rates by hummingbird pollinators in these experimental arrays. However, in natural populations studied, clumped plants were more frequently approached by hummingbirds than those growing more widely dispersed. Display size was unrelated to visitation by Hylemya on inflorescences I clipped and maintained as large, small and control. Display size was also unrelated to the total number of visits by hummingbird pollinators to each of these experimental plants, however large display plants were more likely to be visited first in any given visitation sequence. Of various morphological measurements, corolla length showed the strongest positive correlation with Hylemya egg presence. To the extent that plant spacing and morphology is correlated with pollinator visits and ultimate seed set, Hylemya could be choosing flowers optimally, and playing a role in the evolution of floral traits.     

Oviposition choices by a pre-dispersal seed predator (Hylemya sp.)

Summary Pre-dispersal seed predators must choose proper hosts and flowers on those hosts that have a high probability of setting seed. In this study, I documented that flowers of Ipomopsis aggregata (Polemoniaceae) chosen by a pre-dispersal seed predator (Anthomyiidae: Hylemya sp.) had a higher probability of seed set than flowers the fly did not choose. I experimentally tested the hypothesis that Hylemya uses a post-fertilization cue in making oviposition choices. I found no evidence for this hypothesis.     

Ovipositional Phenology and Behavior of the Spruce Cone Fly, Strobilomyia neanthracina (Diptera: Anthomyiidae)

The behavior of Strobilomyia neanthracina Michelsen, a phytophage infesting spruce (Picea spp.) seed cones, was observed at a field site in northern Ontario and in cages in a greenhouse to investigate spatiotemporal aspects of mating, host location, and oviposition. In the field, adults emerged from 21 to 24 May 1996, which coincided with bud burst of Picea glauca (Moench) Voss seed cones. For 4 days following emergence, Strobilomyia flies could no longer be seen at a monitored P. glauca tree and may have been on a mating or dispersal flight. Subsequently, females but not males were seen again and the oviposition period of ca. 3 weeks began. Mating was observed only in the greenhouse, mostly (i.e., 65%) at age 5–9 days. Although copulations lasted 11–45 min, these females laid infertile eggs only, beginning at age 4 days. No sperm was found in the spermathecal capsules of females, suggesting that no sperm had been transferred during these copulations. In both the field and the greenhouse, ovipositional sequences that resulted in egg deposition occurred throughout the day but few sequences were observed before 1000, probably because flies were not very active at air temperatures below 14°C (most sequences occurred at 25–27°C). In the greenhouse, the typical ovipositional sequence lasted an average of 7 min and consisted of landing on the cone and examining it with the proboscis and sometimes the ovipositor, egg deposition, and postovipositional behaviors such as tapping (touching the cone surface with the flabellum ca. 5 times s ^–1 ), which possibly represents a host marking behavior. In the field, tapping was seen less frequently than in the greenhouse but occurred significantly (P = 0.014) more often after sequences that resulted in egg deposition than after sequences that did not. Eggs hatched after 4–5 days at 20°C. In the greenhouse, the median longevity of females and males was 24 and 17 days, respectively.     

Taking a chance on moths: oviposition by Delia flavifrons (Diptera: Anthomyiidae) on the flowers of bladder campion, Silene vulgaris (Caryophyllaceae)

Abstract.

• 1 Delia flavifrons Zetterstedt (Diptera: Anthomyiidae) visits flowers of Silene vulgaris (Moench) Garcke (Caryophyllaceae), where the adults feed, mate, lay their eggs, and the larvae feed on developing seeds. The objective of the study was to examine how an ovipositing female fly assures a food resource for her progeny.
• 2 Ovipositing females preferred young, non-pollinated flowers over older pollinated ones. The flies did not pollinate the flowers and survival of the larvae depended on the flowers being pollinated by moths.
• 3 Flowers containing fly eggs were pollinated more often than expected from chance, probably as a result of both flies and moths visiting particular flowers.
• 4 Eggs were laid singly, and multiple oviposition occurred randomly. Although most eggs hatched, only about half produced larvae that made their way into the fruits. As a result, the probability of competition arising from multiple oviposition may be reduced to such an extent that selection does not favour females that avoid flowers with conspecific eggs.
• 5 Moth larvae of the noctuid genus Hadena also feed on the seed pods of S. vulgaris and will kill any fly larvae they encounter. However, there was no deviation from random oviposition by the flies in relation to eggs laid by the moths, but the competitively weaker fly usually started to lay eggs towards the end of the moth's egg-laying period.
• 6 The relationship between Delia flavifrons and Silene vulgaris superficially parallels that for known pollinator/predator systems, but floral adaptations to hypothetically pollinating flies seems not to have taken place.

     

Oviposition behavior of the cabbage root fly,Delia radicum (L.), influenced by host plant extracts

An ethanolic extract of cabbage leaves (Brassica oleraceavar. capitata,Golden Acre)and derived fractions were tested on natural and surrogate leaves in order to study the oviposition behavior of the cabbage root fly Delia radicum(Diptera: Anthomyiidae). On surrogate plastic leaves coated with a thin layer of paraffin wax and treated with 0.1 gram leaf equivalent (gle) of an ethanolic raw cabbage extract, the females displayed the same sequence of behavioral patterns as on a natural host plant. The quantified oviposition behavior correlated well with the actual number of eggs laid. The extract-treated surrogate leaves received at least as many eggs as natural leaves with a similar surface area. Previous exposure to surrogate or natural leaves seemed not to induce a specific preference. The three fractions (hexane, butanol, and water) of the raw extract stimulated the oviposition behavior. This was taken as evidence for the presence of a multicomponent mixture or a chemical pattern in the cabbage leaves stimulating oviposition. At the tested concentration each fraction alone could stimulate in some individuals the complete behavior. No effect of volatile components of the raw extract could be detected. This is thought to be due to the extraction procedure, which limits the production of attractive, volatile compounds, such as isothiocyanates.     

Lifetime ovipositional patterns of mated and virgin onion flies,Delia antiqua (Diptera: Anthomyiidae)

Over their 47.2±1.9 (mean±SEM) day lifespan, mated onion flies, Delia antiqua, oviposited more uniformly than did virgins. Mated females began ovipositing at 6.4±0.2 days old and regularly deposited 14.2±0.6 eggs/day for 3-4 weeks. Thereafter, oviposition slowed and stopped at about 1 week before death. Virgin flies began ovipositing 24.7±1.5 days into their 59.0±3.8 day lifespan, and deposited eggs at an increasing rate for 3-4 weeks, generating a mean overall ovipositional rate of 5.8±0.5 eggs/day. The later onset of virgin oviposition (ovipositional activation) and not a shorter ovipositional period largely accounted for the disparity between virgin and mated female lifespans. Mean lifetime egg production of mated females was 475±27 eggs versus 179±30 eggs for virgins. Ovipositional and post-ovipositional periods (34.4±1.8 and 7.2±1.0 days) for mated females were not significantly different from those of virgin females (30.9±3.6 and 6.7±1.2 days). Over 90% of virgin females laid eggs. The distinction between mated and virgin ovipositional patterns may be specifically attributed to differences in: (1) egg maturation, (2) age at ovipositional activation, and (3) egg-laying rate.     

Influence of Conspecific Presence, Experience, and Host Quality on Oviposition Behavior and Clutch Size Determination in Anastrepha ludens (Diptera: Tephritidae)

Females of some insect species adjust the number of ovipositions and clutch size adaptively depending on conspecific density and probably experience. In a series of three experiments, we examined the effect of the presence of conspecifics, host quality, and oviposition experience on oviposition behavior and clutch size determination by females of the polyphagous fruit fly Anastrepha ludens (Diptera: Tephritidae). In the first experiment, we determined that grouped (eight females per cage) A. ludens females tended to visit and oviposit in more hosts than did solitary females probably as a result of stimulation by the presence of conspecifics. We also determined that females with previous oviposition experience visited and oviposited in more hosts than inexperienced ones. Importantly, when females were grouped, we observed significantly more landings on unoccupied hosts (i.e., devoid of flies) than on occupied ones (i.e., with at least one fly on it). However, oviposition experience, and not female density, was the most important factor affecting clutch size. Naive females deposited larger egg clutches than experienced ones. In the second experiment, we found that oviposition experience and host quality (i.e., clean fruit or fruit covered with a host marking pheromone [HMP] extract), influenced clutch size and the decision of females to defend or not defend the host. Clutch size and number of fights were greater on clean than on HMP-marked hosts. In the third experiment, we observed that host quality (i.e., size) played a significant role with regard to the number of female fights, host marking behavior, and clutch size. Specifically, females fought and dragged their aculeus longer on small- and medium-sized hosts than on large ones. But this behavior varied according to whether females were kept alone or grouped. Clutch size was greatest in the largest hosts. Considering all the above, we believe that the observed increase in ovipositional activity by grouped A. ludens females can be attributed to competition through mutual interference and not social facilitation as has been reported in other tephritid species.     

Courtship behavior and discrimination between potential mates by maleDelia antiqua (Diptera: Anthomyiidae)

The repertoire of courtship behaviors of male onion maggots,Delia antiqua (Meigen), in a laboratory bioassay chamber, was analyzed by direct observation and by video recordings, in conjunction with a multichannel event recorder. Seven courtship behaviors were categorized: inspection from the substrate, aerial inspection, contact from the substrate, contact from the air, genital alignment, copulation, and male-male interaction. The frequency distribution of copulation bouts was best described by a Poisson distribution; peak mating activity occurred about 1 h into the bioassay. The duration in copulo, however, was extremely variable. On average, males spent 30 sin copulo (n=183); <30% of bouts were >50 s. The ability of males to discriminate between sexes, sexually immature and mature females, and between females ofD. antiqua and the cabbage maggot,Delia radicum (L.), was most pronounced in the elements of genital alignment and attempted copulation. The courtship and mating behavior inD. antiqua is consistent with a sequence that relies initially primarily on indiscriminate visual recognition of a potential mate, followed by species-and sex-specific semiochemical recognition upon contact.     

木瓜实蝇对七种寄主果实的选择性

木瓜实蝇属于我国进境检疫性害虫。为明确该虫对不同寄主果实的选择性,以评价其危害风险,本研究选择橙子、苹果、番石榴、杨桃、木瓜、西红柿、辣椒7种果实作为供试寄主,在室内选择性条件下比较了木瓜实蝇在不同寄主上的产卵量和幼虫发育的适合性。结果表明,以寄主气味诱导产卵器采卵时,雌虫在番石榴、橙子上产卵量明显高于其它所有供试寄主;以果实直接诱导产卵时,则以番石榴饲养出的幼虫数最多,而在橙子、西红柿上的幼虫数为0。在非选择性条件下将幼虫接入果实供试,表明木瓜实蝇幼虫在不同供试寄主上的存活率(化蛹率)均较高,最高为木瓜和杨桃,分别为92.0%和91.3%,最低为苹果,达到66.0%。不同供试果实对木瓜实蝇幼虫的发育历期、存活率(化蛹率)和蛹重有显著影响,但各适合性指标在不同寄主间的变化趋势不一致。上述结果表明,寄主气味及果皮特征对成虫的产卵选择性具有明显的影响,以产卵选择性和幼虫发育为综合指标,认为木瓜实蝇对番石榴的选择性最高。     

Modification of female onion fly,Delia antiqua (Meigen), reproductive behavior by male paragonial gland extracts (Diptera: Anthomyiidae)

Egg depositional rates of onion flies, Delia antiqua(Meigen), injected thoracically with extracts of male paragonial glands were identical (14.5 eggs/female/ day) to those of normally mated females. Moreover, when continuously exposed to males, extract-injected females refused to mate and produced unfertilized eggs for the duration of the > 15- day experiment. For this normally monocoitic dipteran, <1 male equiv of paragonial secretion completely reproduced the ovipositional responses characteristic of normal mating, and this effect required no involvement of the genitalia or genital chamber. We suggest that the receptor for the active chemical (s) (sex peptide?) would be an excellent target for biorational insect control by sterilization. Moreover, these primer sex pheromones might play an important role in insect reproductive isolation and evolution.     

Quantitative analysis of ovipositional behavior: Effects of a host-plant chemical on the onion fly (Diptera: Anthomyiidae)

Behavioral responses of female onion flies, Delia antiqua (Meigen), to hostplant cues were quantified during encounters of individual flies with onion plants and onion foliar surrogates. The behavioral repertoire of such females included sitting, grooming, running up and down foliar surfaces, extension of the proboscis such that the labellum contacted foliar and soil surfaces, movements of the tip of the abdomen over surfaces (surface probing), subsurface probing of soil crevices with the ovipositor, and oviposition. Sequences of behaviors preceding oviposition were probabilistic rather than highly stereotyped but generally followed the order given above. Foliar surrogates were used to determine the effects of n-dipropyl disulfide (Pr₂S₂) on the sequence of behaviors leading up to oviposition. The addition of a Pr₂S₂-treated surrogate to a cage increased the frequency of alighting on that surrogate but also increased alighting on a nearby foliar surrogate without Pr₂S₂. After alighting, females encountering surrogates treated with Pr₂S₂ had shorter latencies to proboscis extension and surface probing, spent less time sitting and grooming, and had runs of shorter duration. These females were also more likely to make the transition from probing of surfaces of foliage and soil to subsurface probing of soil crevices and oviposition. Thus, rather than mediating a particular step in the behavioral sequence, Pr₂S₂ played a role throughout the sequence leading up to oviposition. Collectively, these data and past studies on the onion fly support the hypothesis that egglaying is triggered by a temporal summation of inputs to the central nervous system from various sensory modalities rather than strict behavioral chaining, with each transition effected by some unique cue.     

Oviposition behavior of an ant-parasitizing fly,Neodohrniphora curvinervis (Diptera: Phoridae), and defense behavior by its leaf-cutting ant hostAtta cephalotes (Hymenoptera: Formicidae)

This study examines the oviposition behavior of the phorid parasitoid Neodohrniphora curvinervisand the antiparasitoid defense behavior of its leafcutting ant host Atta cephalotes. N. curvinervisfemales are diurnal sit- and- wait parasitoids that attack only outbound foragers of head width 1.6 mm or greater. Females deposit a single egg through the foramen magnum of each host successfully parasitized. Pursuit of hosts is usually initiated when an outbound forager of acceptable size passes by a parasitoid perch site. Individual foragers defend themselves against pursuing parasitoids by outrunning them along the foraging trail or by standing their ground and fending them off with their legs,antennae, and mandibles. At the colony level, susceptible foragers are protected against parasitism by a shift in the forager size distribution toward smaller unsusceptible sizes during the day when parasitoids are active and toward larger sizes at night when parasitoids are inactive. The frequency of parasitism of susceptible foragers was 15%, which is more than five times the frequency found in another system involving the phorid parasitoid Apocephalus attophilusand the leafcutting ant host Atta colombica.We offer several possible explanations for such differences in the frequency of parasitism and also examine reasons for the high incidence of superparasitism (19%) observed in the system studied.     

Preoviposition drinking byCulex restuans (Diptera: Culicidae)

In the field 92.9% of female Culex restuansdrank from the oviposition site before beginning to oviposit. Individuals that drank did so an average of 6.4 times, spending an average of 114.5 s in this behavior. Individuals prevented from drinking by sealing their proboscises either failed to lay or laid abnormal egg rafts. Drinking does not appear to be related to evaluation of the ovisite. Instead, it may be necessary for emptying the entire clutch of eggs from the ovarioles and may also assist in the construction of a normal egg raft.     

Relationship between oviposition preference and offspring performance in Australian Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

Abstract We investigated the oviposition preference and larval performance of Helicoverpa armigera under laboratory conditions to determine if the oviposition preference of individual females on maize, cowpea and cotton correlates with offspring performance on the leaves of the same host plants. The host-plant preference hierarchy of females did not correlate with their offspring performance. Female moths chose host plants that contributed less to their offspring fitness. Plant effects accounted for the largest amount of variation in offspring performance, while the effects of female (family) was low. The offspring of most females (80%, n  = 10) were broadly similar, but 20% (two out of 10), showed marked difference in their offspring performance across the host-plant species. Similarly, there was no relation between larval feeding preference and performance. However, like most laboratory experiments, our experi-mental design does not allow the evaluation of ecological factors (for example, natural enemies, host abundance, etc.) that can play an important role in larval performance in the field. Overall, the results highlight the importance of carrying out preference performance analysis on the individual or family level, rather than pooling individuals to obtain average population data.     

Conspecific Sharing of Breeding Sites by Anopheline Female Mosquitoes (Diptera: Culicidae) Inferred from Microsatellite Markers

The number of Anopheles gambiae and Anopheles arabiensis females that used each of the 33 sampled breeding sites in west Kenya was estimated by microsatellite markers and related statistics to test the hypothesis that conspecific females share aquatic sites. Totally, 166 An. gambiae and 168 An. arabiensis larvae were identified and were genotyped. The mean number of larvae per breeding site was 8.3 for An. gambiae and 8.4 for An. arabiensis. The likelihood method estimated that, for An. gambiae, the mean number of females that would have laid eggs per breeding site was 5.2 and ranged from 2 to 9, and for An. arabiensis, the mean was 5.0 with a range of 2–10. The clustering method estimated that the mean number of females laying eggs per breeding site was 6.8 for An. gambiae. The results provide molecular evidence that females of one or both species share breeding sites.     

Oviposition behavior of Edovum puttleri, reared on two hosts, Leptinotarsa decemlineata and L. texana

Female Edovum puttleri Grissell [Hymenoptera: Eulophidae], reared from eggs of Leptinotarsa decemlineata (Say) or Leptinotarsa texana Schaeffer [Coleoptera: Chrysomelidae], were videotaped as they attacked egg masses of L. decemlineata containing 20 host eggs. We identified 15 components of ovipositional behavior. Parasitoids reared on L. texana attacked and oviposited in significantly more host eggs than did females reared on L. decemlineata. Ethometric analyses of behavioral transitions and a clustering analysis of 34 behavioral parameters showed that females reared on L. texana attacked the host egg mass in a different manner than those reared from L. decemlineata. It was concluded that differences were associated with the host species upon which they were reared. Contrary to previous reports, mortality of unparasitized hosts was caused by an ovipositor probe of short duration, which was not related to host-feeding.     

Identity and bioactivity of oviposition deterrents in pine oil for the onion maggot, Delia antiqua

The oviposition deterrent properties of pine oil (Norpine 65, Northwest Petrochemicals, Anacortes, Washington) for the onion maggot, Delia antiqua (Meigen), were verified using a two-choice bioassay with onion oil as an attractive control. The principal deterrent property of this pine oil was found to reside in three monoterpenes, 3-carene, limonene, and p-cymene, which were the primary constituents identified in the most deterrent of two fractions made by preparative gas chromatography of steam-distilled pine oil. At release rates of 220, 320 and 320 g per 24 h in two-choice bioassays these monoterpenes respectively caused 73.2, 65.4 and 56.3% deterrency of oviposition, while the ternary mixture released at 320 g per 24 h caused 88.6% deterrency. The ternary mixture also caused 62.5% deterrency in a no-choice bioassay. Of eight other monoterpenes tested, myrcene, -phellandrene, -terpinene, -phellandrene, -terpinene, terpinolene, and -pinene were significantly deterrent in declining order, while -pinene was inactive. The ternary mixture was released from glass capillary tubes or flexible plastic cylinders in further bioassays that challenged caged females to oviposit around the base of 35 potted onion seedlings with release devices placed on the soil surface. The most effective deterrency (85.3%) was achieved at a release rate of 280 g per 24 h per pot if plastic cylinder devices were deployed 24 h before the treated pot was exposed to D. antiqua females. If female D. antiqua were given only a treated pot, deterrency of oviposition on potted onion seedlings was significant, but low (11.7–63.2%). Because of incomplete efficacy, a monoterpene-based deterrent formulation would be best used operationally if combined with other deterrents, or if it were integrated with some other tactic.     

Oviposition Site Selection by the Regal Fritillary, Speyeria idalia, as Affected by Proximity of Violet Host Plants

Selection of oviposition sites by female regal fritillary butterflies, Speyeria idalia (Drury), in relation to the location and abundance of their larval food plant, Viola pedatifida G. Don, was studied in Kansas tallgrass prairie. To identify potential cues that females use to select oviposition microsites, we compared plots in which eggs were laid with paired control plots in terms of violet abundance, distance from plot center to the nearest violet plant, plant species composition and relative abundance, plant diversity (Shannon's index), and average plant canopy height. No significant differences occurred between control and oviposition sites for any of these variables (P > 0.15). Violets were present in only 30% of the 1-m-diameter oviposition plots, compared with 24% of the corresponding controls. Furthermore, eggs were never laid directly on violets even when they were present nearby. These data imply that S. idalia females do not preferentially orient to or oviposit on their larval food plant. Nevertheless, female S. idalia did not oviposit randomly, for they invariably chose microsites near the ground that were shaded by the grass/forb overstory and usually deposited eggs on the underside of dead vegetation. We speculate that environmental stresses such as high temperatures and intense solar insolation during the egg stage or harsh overwintering conditions experienced by first-instar larvae impact survival and influence choice of oviposition sites. High fecundity (more than 2000 eggs in some individuals) may allow this species to use a conditional, sweepstakes strategy, i.e., producing and spreading many eggs over prairie that harbors host plants to compensate for low egg and larval survivorship.     

Oviposition behavior in spruce budwormChoristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae)

Epicuticular waxes of balsam fir, Abies balsamea,act as oviposition stimulants for fertilized females of the eastern spruce budworm, Choristoneura fumiferana(Clem.). Those same waxes do not stimulate unfertilized females, nor do the latter respond to nonchemical cues such as leaf texture, shape, etc. Ablation experiments suggest that the chemoreceptors responsible for the perception of waxes are located on the proboscis.     

椰子木蛾的产卵节律及其对寄主植物的产卵选择性

【背景】椰子木蛾是近年来新入侵我国棕榈科植物的害虫,研究其产卵习性可为监测和防治该虫提供参考。【方法】在室内条件下,观察、记录了椰子木蛾雌成虫的产卵节律及其对不同寄主植物的产卵选择性。【结果】椰子木蛾最高日产卵量可达34.4粒·头-1,且主要集中在羽化后4 d内产卵,占总产卵量的54.1%;产卵活动主要发生在夜间23:00到次日8:00;在椰子、蒲葵、大王棕、槟榔和散尾葵等寄主植物上的产卵量无显著差异,为89.3~147.7粒·头-1,但产卵位置存在差别。【结论与意义】椰子木蛾雌成虫具有较强的繁殖能力和产卵节律性,且在不同寄主植物上的产卵量一致。