Anne L. Averill

Associative learning in egglaying site selection by apple maggot flies.

Evidence is presented demonstrating that associative learning during oviposition in Crataegus or apple hosts can significantly influence the propensity of apple maggot flies to accept or reject these hosts in future encounters. The data suggest that within resource patches of a given host type there may be an enhancement of foraging efficiency.

Intraspecific competition in the tephritid fruit fly Rhagoletis pomonella

At densities found in nature, competition among larvae of the frugivorous fruit fly Rhagoletis pomonella (Diptera: Tephritidae) was pronounced. In fruits of Crataegus mollis, a small native host of the fly, survivorship dropped sharply when > 1 larva developed per fruit. Multiple infestation also caused decreased size of pupae, which resulted in pro- longed maturation of females and lower daily fecundity. Our findings suggest that inter- ference competition occurs among rival R. pomonella larvae within the same fruit and that older larvae may be competitively dominant. In most instances, when 2 d separated the introduction of two larvae into unpicked fruits of Crataegus oxyacantha, a very small ornamental hawthorn, the first larva pupated while the subsequent larva failed to complete development. Rhagoletis pomonella marks its oviposition site with a contact pheromone. The amount of pheromone deposited following a single egg-laying in C. mollis fruit was sufficient to deter most females from laying more eggs. By avoiding marked fruits in which a larva is already developing, a female may conserve eggs and add measurably to her fitness. Because most C. mollis fruits can support more larvae to puparium formation than they actually do, it appears that the strategy of host marking by females and monopolization of fruit by larvae may leave some fruit resources unexploited.

Specificity of olfactory responses in the tephritid fruit fly, Rhagoletis pomonella

Electroantennograms (EAG) were recorded from, and behavior observed of female apple maggot flies, Rhagoletis pomonella (Walsh) (Diptera: Tephritidae), in response to over 60 individual esters. For acetates through decanoates, 2 methylbutyrates, and isobutyrates, we tested homologous series of systematically altered chain lengths. Most of the compounds had been isolated from behaviorally active fractions derived from extracts of volatiles produced by host fruits of R. pomonella. For the acetates through hexanoates, maximum EAG amplitudes were elicited by esters with chain lengths of 9 carbons and for the heptanoates through nonanoates, by 10 carbon esters. Recovery time, or the rate at which the EAG trace returned to the baseline following maximum depolarization, was slowest for straight chain esters that were 9–11 carbons long. Branching of the chain by addition of a methyl group to either side of the ester resulted in a decline in amplitude and a faster recovery time. Compared to EAG results, only 5 esters (butyl and pentyl hexanoate; propyl and butyl heptanoate; propyl octanoate) were highly active in wind tunnel bioassays, demonstrating (1) the hazard of assigning significance to EAG‐active compounds without accompanying behavioral data, but more importantly; (2) a high degree of olfactory specificity. Maximum behavioral response was contingent upon the following rules regarding size and structure of the molecule. The ester must (1) be a straight chain; (2) be 10–11 carbons in length; (3) have an acid portion of 6–8 carbons and an alcohol portion of 3–5 carbons. One of the active esters, butyl hexanoate, appears in significant concentrations in the headspace of host fruit and, because this ester elicits such a pronounced behavioral response, our results suggest that R. pomonella is adapted to perception of a compound that is typical of its hosts.

Trapping Phyllophaga spp. (Coleoptera: Scarabaeidae: Melolonthinae) in the United States and Canada using sex attractants.

Abstract The sex pheromone of the scarab beetle, Phyllophaga anxia, is a blend of the methyl esters of two amino acids, L-valine and L-isoleucine. A field trapping study was conducted, deploying different blends of the two compounds at 59 locations in the United States and Canada. More than 57,000 males of 61 Phyllophaga species (Coleoptera: Scarabaeidae: Melolonthinae) were captured and identified. Three major findings included: (1) widespread use of the two compounds [of the 147Phyllophaga (sensu stricto) species found in the United States and Canada, males of nearly 40% were captured]; (2) in most species intraspecific male response to the pheromone blends was stable between years and over geography; and (3) an unusual pheromone polymorphism was described from P. anxia. Populations at some locations were captured with L-valine methyl ester alone, whereas populations at other locations were captured with L-isoleucine methyl ester alone. At additional locations, the L-valine methyl ester-responding populations and the L-isoleucine methyl ester-responding populations were both present, producing a bimodal capture curve. In southeastern Massachusetts and in Rhode Island, in the United States, P. anxia males were captured with blends of L-valine methyl ester and L-isoleucine methyl ester. Resumen La feromona sexual del escarabajo, Phyllophaga anxia, es una mezcla de los ésteres metílicos de dos aminoácidos, L-valina y L-isoleucina. Se condujo un estudio de campo usando diferentes mezclas de los dos componentes en 59 sitios de Estados Unidos y Canada. Más de 57,000 machos de 61 especies dePhyllophaga fueron capturados e identificados. Tres de los resultados más importantes incluyen: (1) el extenso uso de los dos componentes [de las 147 especies de Phyllophaga (sensu stricto), en Estados Unidos y Canada, fueron capturados machos de cerca del 40% de ellas.]; (2) para la mayoría de las especies, la respuesta intraespecífica de los machos a las combinaciones de los dos aminoácidos fue consistente entre años diferentes, y en todos los sitios geográficos; y (3) un inusual polymorfismo de la feromona fue descrito para P. anxia. Poblaciones de algunos sitios fueron atrapados sólo con valina, mientras que poblaciones de otros sitios fueron atrapados sólo con isoleucina. También se encontraron sitios donde las poblaciones responden a ambos componentes, valina e isoleucina, produciendo una curva de captura bimodal. En el sureste del estado de Massachusetts y en Rhode Island, en Estados Unidos, machos de P. anxia fueron atrapados en trampas con mezclas de valina e isoleucina.

Residual activity of oviposition-deterring pheromone inRhagoletis pomonella (Diptera: Tephritidae) and female response to infested fruit.

Under dry conditions, oviposition-deterring pheromone (ODP) of the apple maggot fly (Rhagoletis pomonella Walsh) on host fruit proved deterrent to ovipositing females for at least three weeks, with a half-life of ca. 11 days. There was no difference in decline of residual activity under lab vs. field conditions or between fly-deposited ODP vs. an application of a water extract of ODP. A decline in pheromone activity resulted from exposure to both natural and simulated rainfall. For natural rainfall, greatest losses (50– 61%) in activity resulted from high-intensity rains, with substantially less activity lost (13–35%) following light or moderate intensity rains.R. pomonella females discriminated against fruit without ODP but with conspecific larvae. Discrimination against infested fruit was manifested within fewer days following infestation of small fruit (9 mm diam.) as compared to larger fruit (15 mm diam.).

Site of production of an oviposition-deterring pheromone component in Rhagoletis pomonella flies

Abstract Using behavioural and electrophysiological assay techniques, we identified the posterior half of the midgut as being a principal site of production of a major component of the oviposition-deterring, fruit-marking pheromone of female Rhagoletis pomonella flies. Following secretion into, and accumulation in, the gut lumen, this component is released, together with other gut contents, in the marking trail deposited during dragging of the ovipositor on the fruit surface after egg-laying, as well as in the faeces. Other components of the pheromone may be produced elsewhere.

Costs and Benefits of Food Foraging for a Braconid Parasitoid

The costs and benefits of food foraging for the insect parasitoid, Phanerotoma franklini Gahan (Hymenoptera: Braconidae), were examined. Feeding benefited P. franklini by increasing longevity, but did not influence egg maturation. Costs of feeding are represented by the time required to locate and consume food. Field observations of starved wasps revealed that wasps spent approximately 25% of their time grazing on substances on the surface of cranberry foliage. However, our initial laboratory tests to determine the value of these substances were inconclusive. We also examined how grazing behavior integrated with host foraging by comparing the attack rate of fed and unfed wasps held under different host densities. While feeding status did not affect the mean attack rate of wasps, starvation enhanced the probability that a wasp will engage in movement over the host plant foliage. We conclude that feeding benefits P. franklini by increasing longevity, that travel time to food sources is likely to be low, and that the observed food-foraging behavior does not appear to influence host-foraging efficiency.

Factors influencing release of hostmarking pheromone byRhagoletis pomonella flies.

The effect of fly or fruit treatments on quality and/or quantity of host-marking pheromone (HMP) trail substance released by apple maggot flies (Rhagoletis pomonella) following oviposition was evaluated. Among flies, considerable variation existed in the amount of HMP substance deposited, but overall, the amount of substance released on successively offered fruit (over a day or a week) did not change appreciably. Fly diet did not influence pheromone activity. Older flies (28 days) or smaller flies released less or less active HMP trail substance than younger flies (14 days) or larger flies. Females deposited a similar amount of trail substance on large (18–19 mm diam.) or HMP-marked fruit as on small (12–13 mm) or unmarked fruit. Starvation reduced the amount of measurable trail substance deposited but resulted in a more active HMP deposition. Discrepancy between trail measurement and behavioral bioassay results for the starvation treatment indicated that trail measurement results may be misleading under conditions that reduce gut contents of the fly.

Flooding in cranberry to minimize insecticide and fungicide inputs

A traditional flooding practice showed promise for reducing pesticide use in Massachusetts cranberry beds. The 4-week long spring flood, termed “late water,” checked pressure of two of the most prevalent and severe management problems, cranberry fruitworm (CFW) and fungal finit rots (CFR). Fruit rot generally was equal or lower and infestation of fruitworm was consistently lower at the late water beds compared with unflooded beds, even though no fungicides were used for CFR control (standard practice is two or three applications per season) and insecticide applications for CFW were cut by almost 70%. Although there were no significant reductions in yield in two of the years studied, a significant decline (about 25%) occurred on late water beds in one year for unknown reasons.

Mating Disruption of Oriental Beetle (Coleoptera: Scarabaeidae) in Cranberry Using Retrievable, Point-Source Dispensers of Sex Pheromone

Abstract We evaluated the potential management of oriental beetle, Anomala orientalis (Waterhouse), in cranberry by disruption of communication between the sexes through application of the female sex pheromone deployed from wax disks in 2003 and 2004. Mean weekly catch of males in pheromone traps on each untreated bog minus that on each pheromone-treated bog was significantly greater than zero throughout the male flight period for both years of the experiment. Captures of males in traps placed on the edge of pheromone-treated bogs were intermediate between those on control and treated sites; thus, disruption may be limited a short distance from pheromone-treated plots, and treatment of surrounding areas that harbor beetles may be necessary. We deployed a limited number of tethered virgin females in the field for two nights in 2003; when returned to the laboratory, only females on control sites laid fertile eggs. In 2004, we assessed whether delayed mating occurred in the field by deploying tethered females within soil-filled pots and estimated mating rates over five nights by the presence or absence of fertile eggs within soil samples. Only one female among the treated sites laid fertile eggs, whereas the cumulative percentage of females laying fertile eggs in control sites rose steadily over five nights to 96%. The results of this study suggest the strong potential of mating disruption of oriental beetle in cranberry using retrievable, high-dose, point-source dispensers of pheromone.