Evidence for multiple introductions of Centaurea stoebe micranthos (spotted knapweed,Asteraceae) to North America

Invasive species’ success may depend strongly on the genetic resources they maintain through the invasion process. We ask how many introductions have occurred in the North American weed Centaurea stoebe micranthos (Asteraceae), and explore whether genetic diversity and population structure have changed as a result of introduction. We surveyed individuals from 15 European native range sites and 11 North American introduced range sites at six polymorphic microsatellite loci. No significant difference existed in the total number of alleles or in the number of private alleles found in each range. Shannon–Weaver diversity of phenotype frequencies was also not significantly different between the ranges, while expected heterozygosity was significantly higher in the invasive range. Population structure was similar between the native range and the invasive range, and isolation by distance was not significant in either range. Traditional assignment methods did not allocate any North American individuals to the sampled European populations, while Bayesian assignment methods grouped individuals into nine genetic clusters, with three of them shared between North America and Europe. Invasive individuals tended to have genetically admixed profiles, while natives tended to assign more strongly to a single cluster. Many North American individuals share assignment with Romania and Bulgaria, suggesting two separate invasions that have undergone gene flow in North America. Samples from three other invasive range sites were genetically distinct, possibly representing three other unique introductions. Multiple introductions and the maintenance of high genetic diversity through the introduction process may be partially responsible for the invasive success of C. stoebe micranthos.     

Influence of male aggregation size on female visitation in Bactrocera tryoni (Froggatt) (Diptera: Tephritidae)

Abstract  The importance of male aggregation size for female visitation and initiation of male pheromone-calling was investigated in Bactrocera tryoni (Froggatt) (Diptera: Tephritidae) using artificial male aggregations in large laboratory cages. Female B. tryoni visited the largest aggregation more frequently than single males in association with a higher proportion of calling males, but there was no correlation between aggregation size and female visitation. Female B. tryoni had a limited capacity to perceive a difference between the number of calling males. Calling propensity of male B. tryoni was increased by the presence of conspecific males. Increased calling propensity in larger groups of male B. tryoni may be due to social facilitation of male calling behaviour. Female visitation at aggregations was only weakly associated with male calling, suggesting that aggregation size and the number of pheromone-calling males are not the only factors important in locating mates in B. tryoni , and it is possible that low-density populations could persist so long as females can encounter single males.     

Insecticide,sugar, and diet effects on feeding and mortality in Rhagoletis indifferens (Dipt., Tephritidae)

The effects of spinosad bait and various insecticides, the presence of sugar in insecticides, and diet on feeding responses and mortality in western cherry fruit fly, Rhagoletis indifferens Curran (Dipt., Tephritidae), were determined. Numbers of feeding events on insecticides with sugar were greater than on insecticides alone, but there was only a small effect of diet on feeding responses to insecticides with sugar. Feeding durations on imidacloprid, thiamethoxam and acetamiprid with sugar were shorter than on sugar water and spinosad bait, as the neonicotinoids paralysed flies quickly. Flies that fed on sugar only (nitrogen‐starved) suffered higher mortalities when exposed to spinosad, thiamethoxam and azinphos‐methyl than to imidacloprid, acetamiprid and indoxacarb, and mortality in between these two groups of treatments when exposed to spinosad bait. Mortalities were greater when sugar was added to insecticides, and were higher in nitrogen‐starved than fully‐fed (yeast extract + sugar fed) flies. Flies that fed once on thiamethoxam were killed more quickly than those that fed once on spinosad bait and spinosad. Results suggest that thiamethoxam is comparable to spinosad in its effects on mortality, and that using it with sugar in bait may also have similar results as using spinosad bait or spinosad. One benefit of using thiamethoxam with sugar may be that it kills flies more quickly, before they can oviposit, than spinosad bait, although whether a fly will feed on it may depend on how much sugar or nitrogenous food it has eaten.     

Seasonality and host utilization of the invasive fruit fly, Bactrocera invadens (Dipt., Tephritidae) in central Tanzania

Abstract:  The temporal occurrence of the invasive and economically important pest fruit fly, Bactrocera invadens was studied in three agro-ecological areas of Morogoro Region, central Tanzania, during 2004–2005. Weekly and monthly trappings were carried out with methyl eugenol, protein bait and synthetic food attractant. Bactrocera invadens was permanently present at low and mid-altitudes (380–520 m a.s.l.) with peak periods coinciding with the fruiting season of mango ( Mangifera indica ) and guava ( Psidium guajava ). At high altitude (1650 m a.s.l.) its incidence was only temporal and apparently the result of dispersal from lower altitudes after the mango fruiting season. Rearing results showed mango, loquat ( Eriobotrya japonica ), guava and grapefruit ( Citrus  ×  paradisi ) to be the favoured commercial host fruits. Other Citrus species, cucurbits, papaya ( Carica papaya ) and avocado ( Persea americana ) were less favoured.     

Feeding history effects on feeding responses of Rhagoletis indifferens (Dipt., Tephritidae) to GF-120 and Nulure

Abstract:  Effects of feeding history on feeding responses of western cherry fruit fly, Rhagoletis indifferens Curran, to the commercial protein baits GF-120 and Nulure were determined in the laboratory. Flies were kept on 5% sucrose alone or yeast extract and sucrose (Y + S) for 3–7 or 14–16 days and exposed to 24-h-old GF-120 or Nulure drops on artificial leaves. Numbers and durations of feeding events on leaves and durations of non-feeding events were recorded over 1-h periods. Experiments were also conducted to determine effects of Y + S feeding sequences on responses to Nulure, of starvation after sucrose or Y + S feeding on responses to Nulure, and of feeding history on mortality after exposure to GF-120 and Nulure. Protein-deprived flies consistently fed more times on GF-120 and Nulure than protein-fed flies and fed longer. One day of exposure to Y + S or 16 h of starvation after exposure to sucrose caused greater feeding on Nulure than 7 days of exposure to Y + S or 16 h of starvation after exposure to Y + S. Durations of non-feeding events on leaves with sucrose or bait were similar in protein-deprived and -fed flies. Responses of 4- to 6-day-old flies kept on sucrose to 0- and 24-h-old GF-120 or Nulure were similar. More flies kept on sucrose were paralysed or dead at 6–32 h after exposure to GF-120 or Nulure with spinosad than flies kept on Y + S. Results show that complete or long periods of protein deprivation and starvation after sucrose feeding increased feeding responses to GF-120 and Nulure. The general lack of differences in durations of non-feeding events on leaves with sucrose or GF-120 or Nulure in protein-deprived and -fed flies suggests that most protein-deprived flies found baits through chance encounters following normal movement.     

Alcohol dehydrogenase allele frequencies in Bactrocera oleae (Dipt., Tephritidae): effect of ethanol addition in larval diet

Abstract: The effect of ethanol in larval medium on Bactrocera oleae larvae was examined at four concentrations. Ethanol exerted a differential effect on the three alcohol dehydrogenase allele frequencies. While originally being at equilibrium under laboratory conditions, after three generations of larval development in a diet containing ethanol at 1% concentration, Adh -F allele frequency increased, that of Adh -I dropped significantly and the frequency of Adh -S remained unaltered. Adh -S allele seems to be adapted in nature where only minor quantities of alcohol are present in the insects' natural host, while Adh -I is best adapted in the alcohol-free laboratory culture medium. The frequency of Adh -F allele remains unaltered when feral populations are introduced in the laboratory.     

Prospects for improving biological control of olive fruit fly,Bactrocera oleae (Diptera: Tephritidae), with introduced parasitoids (Hymenoptera)

Olive fruit fly is a key pest of olive and consequently a serious threat to olive fruit and oil production throughout the Mediterranean region. With the establishment of Bactrocera oleae in California a decade ago, interest was renewed in classical (introduction) biological control of the pest. Here we discuss the prospects of identifying natural enemies of B. oleae in Africa and Asia that may help reduce B. oleae populations in California and elsewhere. Based on the current understanding of Bactrocera phylogenetics, early opinions that B. oleae originated in Africa or western Asia rather than the Mediterranean region or the Near East are taxonomically and ecologically supportable. Closely related to cultivated olive, the wild olive Olea europaea cuspidata is widely distributed in southern and eastern Africa, the Arabian Peninsula, and eastwards into Asia as far as southwestern China. Little is known regarding the biology and ecology of B. oleae in Africa and eastern Asia, especially in wild olives. While the diversity of parasitoids of B. oleae in the Mediterranean region is low and unspecialized, a diverse assemblage of parasitoids is known from B. oleae in Africa. Conversely, regions in Asia have remained largely unexplored for B. oleae and its natural enemies.     

Detection of the solanum fruit fly, Bactrocera latifrons (Hendel) in Tanzania (Dipt., Tephritidae)

Abstract:  The presence of the Solanum fruit fly, Bactrocera latifrons , in Africa is reported for the first time, based on trapped and reared specimens in Tanzania. Two new host records, Solanum aethiopicum and Solanum macrocarpon , are reported.     

Capture of mass‐reared vs. wild‐like males of Ceratitis capitata (Dipt., Tephritidae) in trimedlure‐baited traps

The sterile insect technique (SIT) is widely used to suppress or eradicate infestations of the Mediterranean fruit fly, Ceratitis capitata (Wied.), an insect whose broad polyphagy poses a serious threat to fruit and vegetable crops. The SIT involves the production, sterilization and release of sterile insects to obtain sterile male by wild female matings, thus yielding infertile eggs. Mass‐rearing over many generations is known to produce dramatic changes in the behaviour and life history of C. capitata. This study investigated the possibility that mass‐rearing also alters male response to trimedlure, a sex‐specific attractant widely used in detection and monitoring programmes. We compared captures of released males from a mass‐reared strain and a recently established colony of wild flies in trimedlure‐baited Jackson traps at three spatial scales – open field, large field enclosures (75 m²) and small field cages (7 m²) – in two separate years. In the first year, males were used independently of flight ability, while in the second year only males with demonstrated flight capability were used. Trap capture was scored 2 days after release for the open field and the large field enclosures but either 1 h or 1 day after release in the small field cages. The findings were consistent across these different experiments: wild‐like males were captured in significantly greater numbers than mass‐reared males in both years of study, except in the trials lasting 1 day in the small field cages where significantly more wild than mass‐reared males were captured in 1 year but not the other. These results are compared with other studies, and their implications for SIT are discussed.     

Revised methods for the mass-rearing of the spotted knapweed biological control agent,Cyphocleonus achates (Coleoptera: Curculionidae), in field corrals

Cyphocleonus achates (Fahraeus), a root-feeding weevil introduced from Eurasia, is an effective biological control agent against spotted knapweed, Centaurea stoebe L. ssp. micranthos. Because C. achates is univoltine and does not fly, distribution of the weevil has been slow. To hasten the weevil's distribution, a rearing effort using field corrals was initiated at a facility in Corvallis, Montana. Procedures for mass-rearing the weevil in field corrals are described, with an emphasis on improvements over earlier methods. The described field-corral approach is effective and appropriate for producing C. achates for distribution in the western United States.     

Identification of 21 polymorphic microsatellites in the African parasitoid wasp, Psyttalia lounsburyi (Silvestri) (Hymenoptera: Braconidae)

We have developed 21 dinucleotide repeat microsatellite loci from African populations of Psyttalia lounsburyi (Silvestri) (Hymenoptera: Braconidae), a parasitoid wasp of the olive fruit fly, as part of a study assessing the role of introgression/hybridization in the success of a biological control introduction. We proposed suitable conditions for polymerase chain reaction multiplexing. All 21 loci were polymorphic with two to 21 alleles per locus within the Kenyan and South African populations tested. Most of them were successfully amplified in two other Psyttalia species.     

Establishment and Dispersal ofUrophora affinis(Diptera: Tephritidae) andMetzneria paucipunctella(Lepidoptera: Gelechiidae) in Southwestern Virginia

Two European gall-producing insects,Urophora affinisFrfld. (Diptera: Tephritidae) andMetzneria paucipunctella(Zeller) (Lepidoptera: Gelechiidae) were introduced into Virginia in 1986 for biological control of spotted knapweed (Centaurea maculosaLam.). Adults ofU. affinis(n = 2625) andM. paucipunctella(n = 450) were released at two sites in Montgomery County, Virginia, and their populations were monitored yearly by dissecting spotted knapweed flower heads. Beginning in 1992, knapweed samples collected at various distances from the release sites were checked for dispersal.U. affinisis well established and is spreading slowly. The number of larvae per flower head and the seed numbers are inversely related as plants with the greatest number of larvae per spotted knapweed head had the lowest number of seeds. Knapweed density has declined at one of the release sites which had the highest rate of infestation byU. affinis.Establishment of the moth,M. paucipunctella,is less certain as it has been recovered at a very low level from only one site.     

Pupal and larval-pupal parasitoids (Hymenoptera) obtained fromAnastrepha Spp. andCeratitis capitata (Dipt.: tephritidae) pupae collected in four localities of Tucuman province,Argentina

Pupal and larval-pupal parasitoids were obtained from 5 % of the 1,413 tephritid puparia collected in four localities of the Tucumán province, Argentina, from April, 1991 to April, 1993.Ceratitis capitata (Wiedemann) was attacked byPachycrepoideus vindemmiae (Rondani) (Pteromalidae), a pupal parasitoid, andAganaspis pelleranoi (Brèthes) (Eucoilidae), a larval parasitoid.Anastrepha spp. were attacked byDoryctobracon areolatus (Szépligeti) (Braconidae), a larval parasitoid, and also byA. pelleranoi. Information about parasitism, percentage of emergence of tephritid species and pupal viability in different localities is provided.     

Effects of the Invasive Forb <Emphasis Type="Italic">Centaurea maculosa</Emphasis> on grassland Carbon and Nitrogen Pools in Montana,USA

Invasions by exotic forbs are changing large areas of North American grasslands, but their biogeochemical impacts are not well characterized. Additionally, although many invasive plants may alter biogeochemistry, an invasive species effects have rarely been evaluated across physically diverse sites. We sampled nine sites containing the perennial Eurasian forb Centaurea maculosa to determine if this invasive species alters soil C and N pools in native grasslands in Montana, USA. We sampled surface soil in adjacent microsites with C. maculosa and native grasses and analyzed soil C and N pools with slow to rapid turnover. None of the pools evaluated in the laboratory showed significant differences between C. maculosa and grass microsites when analyzed across all sites. Some differences were found at individual sites, but they were infrequent and inconsistent: Four sites had no differences, four had differences in one or two pools with intermediate (particulate organic matter C or N) or rapid turnover rates (potentially mineralizable N), and just one site had differences encompassing pools with rapid, intermediate, and slow (total C and N, silt-and-clay-associated N) turnover rates. Where they differed, pools were usually smaller under C. maculosa plants than under native grasses, but the opposite was found at one site. In situ N availability, estimated using ion exchange resins, was significantly lower under C. maculosa than under grasses at one of three sites sampled. Results indicate that C. maculosa may sometimes reduce soil C and N pools, including those related to N availability, but they argue against generalizing about the impacts of C. maculosa in grasslands.     

Supercooling capacity of Urophora affinis and U. quadrifasciata (Diptera: Tephritidae) on spotted knapweed: comparisons among plants, sites, time of season, and gall densities

Larval supercooling points of Urophora affinis Frauenfeld and U. quadrifasciata (Meigen) were compared among plants, six research sites in western Montana, four fall/winter time periods, and among gall densities. These two tephritid fly species are introduced biological control agents of spotted knapweed, Centaurea maculosa Lamarck, and diffuse knapweed, Centaurea diffusa Lamarck. Few differences in larval supercooling points for U. affinis and U. quadrifasciata were found among plants, and where differences were found, they were not consistent across fall/winter time periods. Significant differences in larval supercooling points were found among sites and across fall/winter time periods. No relationship was found between larval supercooling points and site elevation. Larval supercooling points of both U. affinis and U. quadrifasciata showed no relationship with the density of Urophora galls within spotted knapweed capitula. Mean larval supercooling points of U. affinis were consistently lower than those of U. quadrifasciata across sites and fall/winter time periods. In conclusion, temporal differences in temperature over the fall/winter time periods and microclimatic differences among sites appear to be the most important abiotic factors influencing the supercooling points in U. affinis and U. quadrifasciata.     

Biology of Bactrocera carambolae (Diptera: Tephritidae) on four hosts

Bactrocera carambolae is a quarantine pest found in Brazil, restricted to the states of Amapá, Pará and Roraima. This fruit fly can potentially cause extensive socioeconomic and environmental damage in the country, if it disperse into areas where fruit is grown for exporting. The objective of this work was to study the biology of B. carambolae on fruits of Averrhoa carambola L. (Oxalidaceae), Psidium guajava L. (Myrtaceae), Spondias mombin L. (Anacardiaceae) and Eugenia stipitata McVaugh (Myrtaceae). The following parameters were investigated: duration of egg-larva, pupal, egg-adult, pre-oviposition, oviposition and post-oviposition periods, pupal weight and viability, sex ratio, fecundity, fertility and longevity. All parameters except pupal weight, oviposition and post-oviposition period, egg fertility and sex ratio were influenced by the host plant on which the larvae were reared. The carambola fruit fly completes its development on all those hosts studied here, with the highest fecundities on A. carambola and P. guajava.     

Morphometric divergence in populations of Anastrepha obliqua (Diptera,Tephritidae) from Colombia and some Neotropical locations

The West Indian fruit fly, Anastrepha obliqua, is one of seven species of quarantine importance of its genus and is one of the most economically important fruit fly pests in Colombia. The taxonomic status of this species is a key issue for further implementation of any pest management program. Several molecular studies have shown enough variability within Anastrepha obliqua to suggest its taxonomic status could be revised; however, there are no morphological studies supporting this hypothesis. The aim of this work was to describe the morphological variability of Colombian populations of Anastrepha obliqua, comparing this variability with that of other samples from the Neotropics. Measurements were performed on individuals from 11 populations collected from different geographic Colombian localities and were compared with populations from Mexico (2), Dominica Island (1), Peru (1) and Brazil (2). Linear morphometric analyses were performed using 23 female morphological traits, including seven variables of the aculeus, three of the thorax, and six of the wing; seven ratios among them were also considered. Discriminant function analyses showed significant morphological differentiation among the Colombian populations, separating them into two groups. Furthermore, in the comparisons between Colombian samples with those from other countries, three clusters were observed. The possibility of finding more than one species within the nominal Anastrepha obliqua population is discussed.     

Biological attributes of three introduced parasitoids as natural enemies of fruit flies,genus Anastrepha (Diptera: Tephritidae)

The biological attributes of three introduced species of parasitoids which attack the fruit fly Anastrepha ludens were evaluated. Larvae and eggs of A. ludens were exposed to larval parasitoids Diachasmimorpha longicaudata and D. tryoni and the egg parasitoid Fopius arisanus. Parasitoid longevity and fecundity were determined using larvae and eggs of A. ludens. Likewise, the parasitism rates of these parasitoid species in infested host fruits were recorded. The intrinsic rate of increase for F. arisanus was 0.1019 followed by D. tryoni with a rate of 0.1641 and D. longicaudata with the highest rate of 0.2233. Although F. arisanus females had the highest levels of fecundity, only 50% of them remained alive until reproductive age. These results in combination with the longer generation time (in comparison with D. longicaudata and D. tryoni), can be considered as the most important factors explaining F. arisanus reduced rate of increase. However, we note that oviposition activity caused egg mortality which reduced Anastrepha egg hatch by ca. 20%. This result suggests that F. arisanus has a high potential as a natural enemy of A. ludens, in accordance with our research efforts to develop a new F. arisanus strain specialized for development in Anastrepha eggs. The results show that D. tryoni is not a good candidate for biological control of Anastrepha. Meanwhile, D. longicaudata continues to be the most important exotic parasitoid for suppression of Anastrepha fruit fly populations.