A second case of genetic host races in Rhagoletis? A population genetic comparison of sympatric host populations in the European cherry fruit fly,Rhagoletis cerasi

Despite an increasing acceptance in the biological community for sympatric speciation as a mode of species formation, well documented examples of sympatrically evolved ‘incipient species’ remain rare. The sympatric host races of apple maggot, Rhagoletis pomonella (Walsh), represent one of the most prominent case studies for sympatric speciation via a host shift. The European cherry fruit fly, R. cerasi (L.), shows strong ecological similarities to R. pomonella: (1) infestation of two different host plants, Lonicera xylosteum L. and Prunus avium L., and (2) divergent phenological and behavioral adaptations of flies on different hosts. The population genetic study presented here addresses whether the host associated populations of R. cerasi also represent genetically differentiated true host races. Out of a total of 29 allozyme loci examined, six were polymorphic and used to analyze six sympatric pairs of R. cerasi populations on Lonicera and Prunus from Switzerland and Germany. A direct comparison of allele frequencies between sympatric sites showed no pattern indicative of host races in R. cerasi. However, the hierarchical F‐statistic for one locus, mannose 6‐phosphate isomerase (Mpi), showed significant population differentiation that was in accordance with host race differentiation. Mpi is one of several loci that are also diagnostic for host race differentiation in R. pomonella. Results from Mpi suggest the formation of sympatric host races in R. cerasi, but additional polymorphic markers are necessary.     

Substantial progress made in the rearing of the European cherry fruit fly, Rhagoletis cerasi

We examined phenyl propionate as an attractant for trapping navel orangeworm, Amyelois transitella (Walker) (Lepidoptera: Pyralidae) adults, with the objective of developing a method of trapping both sexes more effectively than with almond meal. Two initial experiments maximized the total number of adults captured using phenyl propionate released from glass vials with cotton wicks. A third experiment compared the numbers of males and females captured using these glass dispensers in either bucket or sticky traps. The glass vial dispensers captured more adults than 0.1% phenyl propionate in water (as both attractant and killing agent), and far more adults were captured with glass vial phenyl propionate dispensers than with almond meal. On rare occasion, the glass vial dispensers captured as many adults as traps baited with virgin females, but usually phenyl propionate in glass vials captured fewer adults than virgin‐baited traps. Glass vial phenyl propionate dispensers were equally effective in sticky traps or bucket traps. The majority of females captured were mated, and the proportion of males captured increased over time within flights (generations). We conclude that phenyl propionate released from glass vials captured A. transitella adults more effectively than currently available options, and will be useful in research projects where capturing intact adults and comparing mating status are important. Developing a cost‐effective phenyl propionate‐based alternative to the egg traps currently used for commercial monitoring will be more difficult.     

Susceptibility of different life stages of the European cherry fruit fly,Rhagoletis cerasi,to entomopathogenic fungi

The effects of six fungus isolates on the mortality of different life stages of the European cherry fruit fly, Rhagoletis cerasi (Diptera: Tephritidae), were assessed in a series of laboratory experiments to find an isolate suitable for biological control. In a first step, the effects of fungus treatments on mortality, mycosis and fecundity of adult flies at a concentration of 10⁷ conidia/ml were evaluated. All fungus isolates caused mycosis but virulence varied considerably among the isolates. Beauveria bassiana and Isaria fumosorosea caused 90–100% mortality and had the strongest influence on fecundity. Metarhizium anisopliae also induced high rates of mortality, while the pathogenicity of Isaria farinosa was low. The effects of lower conidia concentrations and the influence of the age of flies were assessed in a second step. Higher conidia concentrations generally resulted in a higher mortality. B. bassiana was most efficient at low concentrations. Young flies showed lower mortality rates than older flies but, sub‐lethal effects on eclosion rate of eggs were greater in younger flies. Finally, the effects on L₃ larvae were tested: none of the fungus isolates induced mortality in more than 25% of larvae. As L₃ larvae and pupae are not susceptible to fungus infection, field control of R. cerasi should be focused on adult flies.     

Host marking pheromone of Rhagoletis cerasi: field deployment of synthetic pheromone as a novel cherry fruit fly management strategy

We tested the efficacy of synthetic host marking pheromone (HMP) of the European cherry fruit fly (R. cerasi) as a fruit-infestation-reducing-agent in an experimental cherry orchard. Two different pheromone deployment strategies were compared: covering entire tree canopies with synthetic HMP or treating only one half (top to bottom or lower half) of the tree canopy. Pheromone application caused a tenfold reduction in fruit infestation if the entire tree canopy was covered (0.226 vs 0.021 pupae/fruit in untreated and treated trees, respectively). Results show, nevertheless, that a significant reduction in fruit infestation can be achieved by treating only one half of tree canopies (top to bottom) (0.021 vs 0.048 pupae/fruit when comparing totally vs partially treated trees). We conclude that synthetic cherry fruit fly HMP has potential as a fruit fly management tool, especially in cases where organically grown fruit reaches high market values.     

Wolbachia infections and superinfections in cytoplasmically incompatible populations of the European cherry fruit fly Rhagoletis cerasi (Diptera,Tephritidae)

Wolbachia is an obligately intracellular, maternally inherited bacterium which has been detected in many arthropods. Wolbachia infections disperse in host populations by mechanisms such as cytoplasmic incompatibility (CI). CI leads to embryonic mortality which occurs when infected males mate with uninfected females or females with a different Wolbachia strain. Populations of the European cherry fruit fly Rhagoletis cerasi (Diptera, Tephritidae) were found to be infected by two different Wolbachia strains, wCer1 and wCer2. Superinfections with both strains occurred throughout southern and central Europe and infections with wCer1 were found in northern, western and eastern Europe. Strong unidirectional CI between European populations of R. cerasi were first reported in the 1970s. From the conformity in the recent geographical distribution of the Wolbachia infections and the CI expression patterns found 25 years ago it was deduced that wCer2 potentially causes CI in R. cerasi. The comparison of the geographical distributions indicated that wCer1 + 2 must have spread into wCer1-infected populations in some areas. In other regions, a spread of wCer1 + 2 was probably prevented by dispersal barriers. There, a sharp transition from infected to superinfected populations suggested regional isolation between wCer1 and wCer1 + 2-infected populations.     

Genetic differentiation between populations of the European rose hip fly Rhagoletis alternata

We tested for genetic differentiation between populations of Rhagoletis alternata Fall. (Diptera: Tephritidae) on three different host species. We collected larvae from three rose species of the section Caninae ( Rosa canina L., Rosa corymbifera Borkh . , and Rosa rubiginosa L.) from 15 sites across Germany, where the three roses occurred together. Additionally, we sampled three sites in Switzerland. Roses differ in morphology (e.g. leaf glands) as well as phenology. We were able to score nine allozyme loci (five polymorphic). Populations from the three hosts did not differ in genetic variability. We found significant genetic differentiation between populations from different host species. However, the differentiation was very low (0.9%). Hence, we found no indication for host races. Furthermore, surprisingly little geographical structure of genetic differentiation was found between populations of this fruit fly across central Europe. We offer three mutually non-exclusive explanations for these findings. First, gene flow between populations of Rh. alternata is high. Second, the pattern of genetic differentiation is based on a recent expansion of the distributional range . Third, the ongoing gene flow between roses of the section Caninae acts as a hybrid bridge.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 619–625.     

Hidden Wolbachia diversity in field populations of the European cherry fruit fly, Rhagoletis cerasi (Diptera, Tephritidae)

The European cherry fruit fly Rhagoletis cerasi has been a field model for cytoplasmic incompatibility since the mid 1970s. Two Wolbachia strains were detected in this tephritid species and w Cer2 was described as the CI inducing agent dividing European populations into two unidirectional incompatible groups, i.e. southern females produce viable offspring with northern males, whereas the reciprocal cross results in incompatibility. We detected three new Wolbachia strains by sequencing a multitude of plasmids derived from Wolbachia surface protein gene ( wsp ) polymerase chain reaction (PCR) products. Strain-specific primers were developed allowing individual diagnosis without need for cloning. Hybridization of specific PCR products with a wsp oligonucleotide enhanced the detection limit significantly and revealed the presence of low-titre infections in some strains, in different ontogenetic stages and in adults of different age. We then performed a survey of strain prevalence and infection frequency in eight European regions. w Cer1 was fixed in all populations, whereas w Cer2 was detected only in the South. w Cer3 frequency was the lowest without a clear distribution pattern. The abundance of w Cer4 was homogenous across Europe. Like w Cer2, w Cer5 showed significant differences in spatial distribution. Our new findings of previously undetected and recombinant Wolbachia strains in R. cerasi reveal a major caveat to the research community not to overlook hidden Wolbachia diversity in field populations. Low-titres and geographical variability in Wolbachia diversity are expected to influence the outcome of Wolbachia population dynamics and Wolbachia- based insect population control and may create invasion barriers for expanding and artificially introduced Wolbachia strains.     

The diapause response of Rhagoletis pomonella to varying environmental conditions and its significance for geographic and host plant‐related adaptation

The recent shift of Rhagoletis pomonella Walsh (Diptera: Tephritidae) from its ancestral host hawthorn to apple is a model for incipient sympatric speciation in action. Previous studies have shown that changes in the over‐wintering pupal diapause are critical for differentially adapting R. pomonella flies to a difference in the fruiting times of apples vs. hawthorns, generating ecologically based reproductive isolation. Here, we exposed pupae of the hawthorn race to various combinations of pre‐ and over‐wintering rearing conditions and analyzed their effects on eclosion time and genetics. We report certain unexpected results in regards to a combination of brief pre‐winter and over‐wintering periods indicative of gene*environment interactions requiring a reassessment of our current understanding of R. pomonella diapause. We present a hypothesis that involves physiological factors related to stored energy reserves in pupae that influences the depth and duration of Rhagoletis diapause. This ‘pupal energy reserve’ hypothesis can account for our findings and help clarify the role host plant‐related life history adaptation plays in phytophage biodiversity.     

Microsatellite markers for testing host plant‐related population differentiation in the moth genus Yponomeuta

Nine microsatellite primers were developed for Yponomeuta padellus (Lepidoptera: Yponomeutidae) and tested for their applicability in analysing genetic population structure. Eight of the nine loci were highly polymorphic with on average 11.4 alleles. Cross‐species amplification of the nine primer pairs was tested in five other moth species. Primer pairs amplified in Y. cagnagellus, Y. malinellus, Y. evonymellus, and Y. rorellus but not in Y. sedellus and Plutella xylostella.     

Standing geographic variation in eclosion time and the genomics of host race formation in Rhagoletis pomonella fruit flies

Taxa harboring high levels of standing variation may be more likely to adapt to rapid environmental shifts and experience ecological speciation. Here, we characterize geographic and host‐related differentiation for 10,241 single nucleotide polymorphisms in Rhagoletis pomonella fruit flies to infer whether standing genetic variation in adult eclosion time in the ancestral hawthorn (Crataegus spp.)‐infesting host race, as opposed to new mutations, contributed substantially to its recent shift to earlier fruiting apple (Malus domestica). Allele frequency differences associated with early vs. late eclosion time within each host race were significantly related to geographic genetic variation and host race differentiation across four sites, arrayed from north to south along a 430‐km transect, where the host races co‐occur in sympatry in the Midwest United States. Host fruiting phenology is clinal, with both apple and hawthorn trees fruiting earlier in the North and later in the South. Thus, we expected alleles associated with earlier eclosion to be at higher frequencies in northern populations. This pattern was observed in the hawthorn race across all four populations; however, allele frequency patterns in the apple race were more complex. Despite the generally earlier eclosion timing of apple flies and corresponding apple fruiting phenology, alleles on chromosomes 2 and 3 associated with earlier emergence were paradoxically at lower frequency in the apple than hawthorn host race across all four sympatric sites. However, loci on chromosome 1 did show higher frequencies of early eclosion‐associated alleles in the apple than hawthorn host race at the two southern sites, potentially accounting for their earlier eclosion phenotype. Thus, although extensive clinal genetic variation in the ancestral hawthorn race exists and contributed to the host shift to apple, further study is needed to resolve details of how this standing variation was selected to generate earlier eclosing apple fly populations in the North.     

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.

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Résumé Les réactions de R. pomonella à plus de 60 esters différents ont été enregistreées par électroantennogrammes (EAG) et par ovservation du comportement. Pour les acétates, avec les décanoates, les 2-méthylbutyrates et les iso butyrates, nous avons examiné des séries homologues de chaînes aux longueurs systématiquement altérées. La plupart de ces composés ont été isolés à partir des fractions actives sur le comportement, issues des extraits des substances volatiles des fruits des plantes hôtes de cette téphritidae. Pour les acétates, avec les hexanoates, les EAG aux plus grandes amplitudes ont été obtenus avec des esters dont la longueur des chaînes est de 9 carbones, et avec les heptanoates et les nonanoates pour les esters à 10 carbones. Le temps de récupération ou temps mis par l'EAG pour revenir à l'ordonnée de départ après dépolarisation maximum, a été plus lent pour les esters à chaînes droites avec 9 à 11 carbones. La ramification de la chaîne par addition d'un groupe méthyl de chaque côté de l'ester a provoqué une réduction de l'amplitude et une accélération de la récupération. 5 esters seuls provoquent une forte réaction dans le tunnel à vent (hexanoates butilique et pentylique, heptanoates propylique et butylique, octanoate propylique). Ceci montre (1) le risque qu'il y a à attribuer une signification aux substances provoquant une réaction en EAG, lorsqu'il n'y a pas d'observations comportementales parallèles, et, surtout; (2) le degré élevé de spécificité olfactive.La résponse comportementale optimale obéit aux règles suivantes concernant la taille et la structure de la molécule. L'ester doit avoir: (1) une chaîne linéaire; (2) 10–11 carbone de longueur; (3) une portion acide de 6 à 8 C et une portion alcool de 3 à 5 C. Le butyl hexanoate, l'un des esters actifs, présent en concentration suffisante près de fruits utilisés, provoque pour cette raison une réponse comportementale tellement marquée, que nos résultats suggèrent que R. pomonella est adapté à la perception des substances caractéristiques de leurs hôtes.

     

Variability in response specificity of apple, hawthorn, and flowering dogwood-infesting Rhagoletis flies to host fruit volatile blends: implications for sympatric host shifts

Rhagoletis pomonella Walsh (Diptera: Tephritidae) originating from domesticated apple (Malus pumila), hawthorn (Crataegus mollis) (Rosaceae), and flowering dogwood (Cornus florida) (Cornaceae) were tested sequentially in flight‐tunnel assays to volatile blends previously identified from the three fruit types. The majority of flies flew to odor sources containing their natal blend (68–83%). Some flies from each fruit type also flew to non‐natal fruit blends (11–39%), but of these non‐natal responders the vast majority were flies that responded to their natal blend as well. The results indicate that individual flies within R. pomonella populations infesting different host types have different degrees of specificity with respect to discriminating among fruit volatile blends, and that a moderate proportion of apple, hawthorn, and dogwood flies (10–30%) are broad responders, with the capacity to recognize and orient to more than one blend. The observed variability in response specificity could facilitate sympatric shifts to new host plants.     

Tarsal contact chemoreceptor for the host marking pheromone of the cherry fruit fly, Rhagoletis cerasi: responses to natural and synthetic compounds

Abstract. The European cherry fruit fly (Rhagoletis cerasi L.; Diptera, Tephritidae) marks cherries (Prunus avium L.) after oviposition with a host marking pheromone (HMP). The marking trail prevents additional oviposition by the same or other females into the same fruit. On the ventral side of the tarsi of both sexes, contact-chemoreceptor sensilla were identified which contain a receptor cell selectively sensitive to HMP. The HMP receptors of males were slightly more sensitive than those of females, suggesting that the more general term ‘host-marking pheromone’ is more appropriate than the previously used ‘oviposition deterring pheromone (ODP)’. The four structural isomers of the HMP, N(15R, S(β-glucopyranosyl)-oxy-8RS-hydroxypalmitoyl)-taurine, and various derivatives were synthesized and tested in an electrophysiological bioassay. Both the 8R,15R and the 8S,15RS isomers of the HMP were equally active with a threshold of about 2 times 10^-10M, and were shown to be present in the female faeces in similar proportions. The two 15S HMP isomers were about 13 times less active. Testing synthetic derivatives of the HMP molecule revealed that the presence of the four moieties of the molecule are important for the activity: taurine, palmitic acid, C₍₈₎ hydroxyl group, and glucose (C₍₁₅₎). The chain length of the fatty acid, the hydroxyl group at C₍₈₎ and the position of glucose at C₍₁₅₎ also influenced the activity. Only minor loss of activity (factor 2) relative to the natural molecule was observed when the methyl group in the C₍₁₅₎ position was removed. The removal of the β-glycosidically linked glucose (replaced by a hydroxyl group) resulted in about a 4-fold loss of activity. The cation of the HMP molecule seemed to have no effect on its activity, whereas both low and high pH reduced it significantly. Based on these results, field experiments have been initiated to control oviposition by cherry fruit flies on cherries applying the 15-desmethyl-HMP derivative.     

Enzymatic and protein patterns at different developmental stages in the cherry fruit fly Rhagoletis cerasi L.

Protein patterns and two isoenzyme systems (leucine aminopeptidase, LAP, and alanine aminopeptidase, AAP) were examined in the cherry fruit fly Rhagoletis cerasi L. at different stages of ontogeny LAP activity was quantitatively determined.Differences were observed in protein patterns for puparia in diapause, post-diapause and at adult emergence. Some LAP and AAP isoenzymes appear in post-diapause which had not been observed during diapause. It is suggested that there is a direct link between variations in protein patterns and proteolytic activity.     

The effect of temperature on the post-diapause development of four geographical populations of the European cherry fruit fly (Rhagoletis cerasi)

Pupae of Rhagoletis cerasi (L.) from Austria, Czechoslovakia, Switzerland and Italy were stored at 4° for 155 days and then transferred to eleven constant and four cycling temperature regimes until the flies emerged. No flies emerged in 324 days at 8° constant but mean emergence was 84% for the remaining regimes. Female flies emerged slightly earlier than males and flies from Italy were the last to start emerging. Reduced to a common mean base temperature of 6.8°, temperature sums in day degrees C to 10% emergence of female flies were 331 (Austria), 319 (Czechoslovakia), 359 (Italy) and 321 (Switzerland). The equivalent sums are also given for first, 50% and 90% emergence. When tested with soil temperature data from Switzerland 321 day degrees above 6.8° and 430 day degrees above 5° gave effectively the same estimates for practical forecasting purposes.

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Zusammenfassung Puppen von Rhagoletis cerasi L. aus üOsterreich, der Slovakei, der Schweiz und aus Italien, welche aus infizierten Kirschen gewonnen worden waren, wurden wüahrend 155 Tagen bei 4\dg gelagert und anschliessend füur die Weiterentwicklung in 11 konstanten und 4 fluktuierenden Temperaturregimen untersucht. Keine Fliegen schlüupften nach 324 Tagen Inkubationsdauer bei 8\dg, wüahrend die durchschnittliche Schlüupfrate bei den üubrigen Temperaturbedingungen 84% betrug. Die beobachteten und berechneten Werte füur die Entwicklungsdauer bis zur Erreichung der Schlüupfraten von 10% und 50% sind nach Geschlechtern und Herküunften getrennt in Tabellen aufgefüuhrt. Lineare Regressionsgleichungen wurden berechnet füur die Kurven der Entwicklungsraten und verwendet füur die Berechnung der Entwicklungsnullpunkte und effektiven Temperatursummen. Weibliche Fliegen schlüupften früuher als Müannchen und Puppen aus Italien entwickelten sich von allen vier Herküunften am langsamsten. Berechnet auf der Basis eines durchschnittlichen Entwicklungsnullpunktes von 6.8\dg als Schwellenwert benüotigten die Weibchen der verschiedenen Herküunfte bis zum Erreichen der Schlüupfrate von 10% folgende Temperatursummen in Gradtagen: üOsterreich 331\dg, Slovakei 319\dg, Italien 359\dg und die Schweiz 321.Die entsprechenden Werte für das Erreichen des Zeitpunktes von Schlüpfbeginn, 50% und 90% Schlüpfrate sind ebenfalls in Tabellenform dargestellt. Der normalerweise füur Prognosezwecke verwendete Schwellenwert von 5\dg wurde zu Vergleichszwecken ebenfalls in die Analyse der experimentellen Daten einbezogen, doch zeigte es sich, dass er zu tief liegt. Die üUberprüufung der Prognosemethode in der Schweiz mittels Bodentemperaturen in 5 cm Bodentiefe unter Anwendung des neuen Schwellenwertes von 6.8\dg und der effektiven Temperatursumme von 321 Gradtagen füuhrte jedoch zu den gleichen Schlussfolgerungen wie die Verwendung des traditionellen Schwellenwertes von 5\dg und der Summe von 430 Gradtagen. Da die Berechnungen auf der Basis von 5\dg einfacher sind und gute üUbereinstimmung mit den Fangresultaten im Freiland zeigen, besteht keine Veranlassung füur die praktische Anwendung den alten Schwellenwert zu üandern.

     

Relative air humidity influences the function of the tarsal chemoreceptor cells of the cherry fruit fly (Rhagoletis cerasi)

ABSTRACT. Electrophysiological recordings from the tarsal D-sensilla of mature, 5–7-day-old female European cherry fruit fly ( Rhagoletis cerasi L.) revealed that a large proportion of sensilla showed no sensory activity in response to stimulation with the oviposition deterring pheromone ('ODP') or sucrose. In contrast, good recordings were obtained from almost all sensilla in freshly emerged, 1-day-old flies. Ageing, nutritional state and contamination were excluded as possible major factors influencing the responsiveness of the D-sensilla. The study of the variability in responsiveness of the sensilla of 1-day-old flies among different tarsomers, among tarsi of the same individual and among individuals, revealed that the major source of variability was among individuals.
Individual flies were also affected by the humidity in the holding cages: relative air humidity of less then 75% for 4–6 days was shown to reduce the number of D-sensilla giving recordings with a good signal-to-noise ratio. The D-sensilla on the most distal tarsomers were affected most by the reduced (30–55% r. h.) air humidity. Flies which had contact with plant leaves showed less reduction in responsiveness at low air humidity (40% r. h.) then flies without leaf contact.     

Population genetic structure of the western cherry fruit fly Rhagoletis indifferens (Diptera: Tephritidae) in British Columbia,Canada

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Invasion genetics of American cherry fruit fly in Europe and signals of hybridization with the European cherry fruit fly

The American cherry fruit fly is an invasive pest species in Europe, of serious concern in tart cherry production as well as for the potential to hybridize with the European cherry fruit fly, Rhagoletis cerasi L. (Diptera: Tephritidae), which might induce new pest dynamics. In the first European reports, the question arose whether only the eastern American cherry fruit fly, Rhagoletis cingulata (Loew) (Diptera: Tephritidae), is present, or also the closely related western American cherry fruit fly, Rhagoletis indifferens Curran. In this study, we investigate the species status of European populations by comparing these with populations of both American species from their native ranges, the invasion dynamics in German (first report in 1993) and Hungarian (first report in 2006) populations, and we test for signals of hybridization with the European cherry fruit fly. Although mtDNA sequence genealogy could not separate the two American species, cross‐species amplification of 14 microsatellite loci separated them with high probabilities (0.99–1.0) and provided evidence for R. cingulata in Europe. German and Hungarian R. cingulata populations differed significantly in microsatellite allele frequencies, mtDNA haplotype and wing pattern distributions, and both were genetically depauperate relative to North American populations. The diversity suggests independent founding events in Germany and Hungary. Within each country, R. cingulata displayed little or no structure in any trait, which agrees with rapid local range expansions. In cross‐species amplifications, signals of hybridization between R. cerasi and R. cingulata were found in 2% of R. cingulata individuals and in 3% of R. cerasi. All putative hybrids had R. cerasi mtDNA indicating that the original between‐species mating involved R. cerasi females and R. cingulata males.     

Genetic structure of cherry fruit fly (Rhagoletis cingulata) populations across managed,unmanaged, and natural habitats

The cherry fruit fly (CFF), Rhagoletis cingulata Loew (Diptera: Tephritidae: Trypetini), is endemic to eastern North America and Mexico, where its primary native host is black cherry [Prunus serotina Ehrh. (Rosaceae)]. Cherry fruit fly is also a major economic pest of the fruit of cultivated sweet (Prunus avium L.) and tart (Prunus cerasus L.) cherries. Adult CFF that attack wild black cherry and introduced, domesticated cherries in commercial and abandoned orchards are active at different times of the summer, potentially generating allochronic isolation that could genetically differentiate native from sweet and tart CFF populations. Here, we test for host‐related genetic differences among CFF populations in Michigan attacking cherries in managed, unmanaged, and native habitats by scoring flies for 10 microsatellite loci. Little evidence for genetic differentiation was found across the three habitats or between the northern and southern Michigan CFF populations surveyed in the study. Local gene flow between native black cherry, commercial, and abandoned orchards may therefore be sufficient to overcome seasonal differences in adult CFF activity and prevent differentiation for microsatellites not directly associated with (tightly linked to) genes affecting eclosion time. The results do not support the existence of host‐associated races in CFF and imply that flies attacking native, managed, and unmanaged cherries should be considered to represent a single population for pest management purposes.     

It''s about time: the evidence for host plant-mediated selection in the apple maggot fly, Rhagoletis pomonella, and its implications for fitness trade-offs in phytophagous insects

The apple maggot fly, Rhagoletis pomonella, Walsh (Diptera: Tephritidae), provides a unique opportunity to address the issue of host-related fitness trade-offs for phytophagous insects. Rhagoletis pomonella has been controversial since the 1860's when Benjamin Walsh cited the fly's shift from hawthorn (Crataegus spp.) to apple (Malus pumila) as an example of an incipient sympatric speciation event. Allozyme and mark-release-recapture studies have subsequently confirmed the status of apple and hawthorn flies as partially reproductively isolated and genetically differentiated host races, the hypothesized initial stage in sympatric divergence. Here, we review the ecological and genetic evidence for host-plant mediated selection in R. pomonella. We reach the following three major conclusions: First, although developmental timing is not everything, it is a good deal of the story. Differences in the fruiting phenologies of apple and hawthorn trees exert different selection pressures on the diapause and eclosion time characteristics of the host races. In particular, the 3-week earlier mean fruiting phenology of apples in eastern North America appears to select for a slower rate of metabolism or deeper pupal diapause in apple than hawthorn flies. Second, host-related fitness trade-offs for R. pomonella may not be due to disruptive selection affecting any one specific life-history stage. Rather, it is the sum total of directional selection pressures acting across different life-stages that generates divergent selection on apple and hawthorn flies. For example, selection favors the alleles Me 100, Acon-2 95 and Mpi 37 (or linked genes) in the larval stage in both host races. However, these same alleles are disfavored in the pupal stage to follow, where they correlate with early adult eclosion, and by inference premature diapause termination. Because apple trees fruit an average of 3 weeks earlier than hawthorn trees, this counter-balancing selection is stronger on apple-fly pupae. The net result is that the balance of selective forces is different between apple and hawthorn flies, helping to maintain the genetic integrity of the host races in sympatry in the face of gene flow. Finally, natural R. pomonella populations harbor a good deal of genetic variation for development-related traits. This variation allows fly populations to rapidly respond to temporal vagaries in local environmental conditions across years, as well as to broad-scale geographic differences that exist across the range of the species. Perhaps most importantly, this variation gives R. pomonella the flexibility to explore and adapt to novel plants. Taken together, our results underscore how difficult it can be to document host plant-related fitness trade-offs for phytophagous insects due to the need to consider details of the entire life-cycle of a phytophagous insect. Our findings also show how reproductive isolation can arise as a by-product of host-associated adaptation in insects, a central theme for models of sympatric speciation via host shifts.