Effect of Two Opiine Parasitoids (Hymenoptera: Braconidae) Introduced for Fruit Fly Control on a Native Hawaiian Tephritid,Trupanea dubautiae(Diptera: Tephritidae)

Diachasmimorpha longicaudata(Ashmead) andPsyttalia fletcheri(Silvestri) are opiine parasitoids introduced into Hawaii for control of the Oriental fruit fly,Bactrocera dorsalis(Hendel) and the melon fly,Bactrocera cucurbitae(Coquillett), respectively. Both species have recently been mass-reared and released for research in augmentative biocontrol programs. Laboratory and field sleeve cage experiments were conducted to investigate the potential impact of mass-producedD. longicaudataandP. fletcherion a native Hawaiian tephritid,Trupanea dubautiae(Bryan), infesting the flowerheads of the native composite shrubDubautia raillardioidesHillebrand. Gravid females ofD. longicaudataandP. fletcheriwere confined with bloomingD. raillardioidesflowerheads infested with late instarT. dubautiae.BothD. longicaudataandP. fletcherilacked positive oviposition responses toT. dubautiaelarvae in infested flowerheads and caused neither parasitism nor mortality to the flies. However, when larvae were removed from the flowerheads and presented in screened dishes containing artificial diet of the parasitoids' normal rearing hosts (B. dorsalisandB. cucurbitae), bothD. longicaudataandP. fletcherireadily oviposited in the test larvae. Oviposition byD. longicaudatadid not significantly affect the percentage pupation ofT. dubautiae,but did reduce the emergence of adult flies. Oviposition byP. fletcherisignificantly reduced both pupation and adult fly emergence. All progeny of both parasitoid species died as eggs or first-instar larvae. Results from our experiments demonstrate that biological control programs targeted against frugivorous tephritid pests byD. longicaudataandP. fletcherihave no harmful impact on flowerhead-infestingT. dubautiae.     

Evidence for an oviposition-deterring pheromone in Tephritis bardanae (Schrank) (Diptera: Tephritidae)

Summary Female Tephritis bardanae (Schrank) exhibit a characteristic ovipositor-dragging behaviour immediately after laying eggs into a flowerbud of Arctium minus (Hill) Bernh. To establish whether this behaviour is associated with an oviposition-deterring pheromone (ODP), female T. bardanae were presented with clean, unattacked flowerbuds and fly activity was monitored using video recording equipment. The distribution of oviposition and fly behaviour were analysed. Avoidance of attacked buds was revealed by the oviposition pattern becoming overdispersed. Spacing out of eggs continued until, on average, each bud contained one egg-batch, after which the distribution became more random or clumped. Analysis of fly behaviour showed that attacked and unattacked buds received similar numbers of visits, but fewer oviposition attempts were made on attacked buds. This behaviour was consistent with use of a contact pheromone deterring repeated oviposition and applied to the bud surface during ovipositor-dragging. Avoidance of attacked buds should increase the efficiency of resource use by T. bardanae populations in the field; however, distributions of egg-batches in flowerhead samples collected from Monks Wood NNR, Cambridgeshire, during 1983–1985, were highly clumped and did not provide supportive evidence. This suggests that the effectiveness of oviposition deterrence in the field is reduced, because the active life of ODP under natural conditions is short, and/or because wild females frequently experience high oviposition drive due to scarcity of suitable flowerbuds.     

Rearing ofChaetorellia australis (Dipt., Tephritidae) on an artificial diet

Larvae of the tephritid flyChaetorellia australis Hering, which infests flowerheads of the yellow starthistleCentaurea solstitialis L. (Asteraceae), and which is a candidate species for biological control, of that weed, were reared for the first time and for two consecutive generations on an artificial diet. At a density of 0.5 eggs per g diet a yield of 18.5% F₁ adults (over hatched eggs), was obtained. When the density of eggs per g diet increased, the yield decreased. At 25°C the duration of development from egg to adult was 20–25 days for both generations. F₁ adults reared on artificial diet had an average longevity of 32.0 days for males and 34.9 days for females. The fecundity was 53.3 eggs per female and the egg hatchability 88.8%. These and other biological parameters are similar to those of wild flies.      

Carbon dioxide assimilation in the flowerhead of Arctium

Summary The gas exchange of flowerheads was determined in Arctium tomentosum and A. lappa during their development. The light, temperature and CO₂ responses were used to estimate flowerhead photosynthesis and the in situ contribution of carbon assimilation to the carbon requirement of the plant for supporting a flowerhead. Changes in vapour pressure deficit had no effect on flowerhead photosynthesis rates and were not included in the model.In both species assimilatory capacity correlated with total bract chlorophyll content. Light, temperature and CO₂ response curves were very similar in form between species, differing only in absolute rates. During all stages of development, flowerheads always exhibited a net carbon loss, which was mainly determined by temperature. The respiration rate decreased in the light, the difference of CO₂ exchange in the dark and in the light was interpreted as photosynthesis. This rate was larger in A. lappa than in A. tomentosum. 30% of the total C requirement of A. lappa flowerheads was photosynthesized by its bracts, the total contribution offlowerhead photosynthesis in A. tomentosum was only 15%. The potential competitive advantages of variation in flowerhead photosynthesis are discussed.     

Distribution of Insect Attacks in Biological Control of Weeds: Infestation of Centaurea virgata Flowerheads by a Gall Fly

The success of biological control efforts to reduce weed density through release of insects may depend as much on the distribution of insect attacks among individual plants or plant parts as on the mean level of infestation. We used an index of dispersion to describe the distribution of Urophora quadrifasciata (Diptera: Tephritidae) galls among squarrose knapweed (Centaurea virgata) flowerheads at 18 west central Utah sites in the first 5 years following introduction of the biological control agent. Two thirds of the samples showed a significantly aggregated distribution of galls among flowerheads. Statistical analysis showed that site and year accounted for relatively small proportions of the variance in the index of dispersion. The degree of gall aggregation among flowerheads was positively correlated with the mean flowerhead quality (mean number of seeds per flowerhead; P = 0.013) and tended to be negatively correlated with the mean fly density per flowerhead at a site in a given year (P = 0.097). Our data suggest that higher quality flowerheads, and possibly higher quality plants, are preferentially attacked by U. quadrifasciata and therefore are more heavily subject to reduced reproductive potential through biological control. However, an aggregated distribution of fly attacks may undercut the potential of the fly to reduce seed production by the weed population as a whole. Understanding both the distribution of insect attacks among individual plants and the behavioral mechanisms producing such distribution patterns is important to the biological control of weeds.     

Biocontrol of the shore fly Scatella tenuicosta with Hypoaspis miles and H. aculeifer in peat pots

Hypoaspis miles Berlese and H. aculeiferCanestrini (Acari: Laelapidae) were usedagainst Scatella tenuicosta Collin(Diptera: Ephydridae) at the rates of 7, 14and 36 mites per 0.3-l peat pot (1000, 2000 and5000 per net-m² of peat) with mint (Mentha piperita L.) as the crop. The predatorswere applied either one day (preventivecontrol) or nine days after (curative control)exposing the pots to the flies in thegreenhouse. Based on the number of flies thatemerged from the pots, the control efficacy of preventive H. aculeifer treatments at therate of 36 mites per pot was 84–100% and 92–97% (range over four replicate blocks)after two and three weeks, respectively, fromthe application. The control efficacy ofcurative treatments was 93–100% and 83–94% after two and three weeks, respectively,from the application. Preventive treatmentswith H. miles applied at the rate of 36per pot controlled the flies by 92–100% and 68–97% two and three weeks, respectively, afterthe application. Curative treatments with 36H. miles per pot were less effective (–53–68% and –16–65%, respectively). All treatments with lower mite rates with eitherspecies were ineffective. Due to moreconsistent reduction of fly numbers by Hypoaspis aculeifer, the species was concluded to be a better biocontrol agent ofshore flies in short-term greenhouse cropsgrown in peat pots, probably due to its higherpredation capacity in comparison to H. miles.     

Insect feeding on spores of a bracket fungus, Elfvingia applanata (Pers.) Karst. (Ganodermataceae, Aphyllophorales)

Insects visiting sporocarps of Elfvingia applanata, a wood-rotting bracket fungus, were examined in Kyoto, central Japan. Mycodrosophila flies (Drosophilidae: Diptera) were predominant and visited the spore-producing sporocarps exclusively. They were observed feeding on the spores, and a number of spores seemed to be alive even after having passed through insects digestive tracts. In addition, the insects attached a number of spores on their body surfaces. In a rearing experiment with insects caught from E. applanata sporocarps, Mycodrosophila flies excreted 7700–469 000 and dropped 10–000–329 000 of viable spores during 48 h after collection. They were supposed to migrate among the sporocarps of other bracket fungi growing on different logs or stumps, suggesting that Mycodrosophila flies may act as spore-dispersal agents for E. applanata.     

High aquatic niche overlap in the newts Triturus cristatus and T. marmoratus (Amphibia,Urodela)

We studied spatial niche metrics of large-bodied newts (Triturus cristatus and T. marmoratus) in three breeding ponds in western France. Adults and larvae were sampled with underwater funnel traps. Larvae were identified to the species with diagnostic microsatellite DNA markers. The distribution of adult T. cristatus and T. marmoratus across pond regions differed in one out of six cases, no differences were observed between larvae (two ponds studied). Niche overlap and niche breadth indices across resource states defined as pond regions or individual traps were high (Schoener's C: pond regions 0.60–0.98, traps 0.35–0.71; Levins' B: pond regions 0.71–0.98, traps 0.35–0.76). Adults of large-bodied newts significantly differed in resource use from small-bodied newts (T. helveticus). The results are discussed in view of the occurrence of interspecific breeding attempts, and the unpredictable ecological characteristics of newt breeding ponds.     

Activity of pupal parasitoids of the stable fly Stomoxys calcitrans and prevalence of entomopathogenic fungi in the stable fly and the house fly Musca domestica in Denmark

A survey was conducted on confined dairy cattle farms and a pig farm from May–October in 1999 to determine the activity and relative abundance of pupal parasitoids and the prevalence of entomopathogenic fungi in populations of the haematophagous stable fly, Stomoxys calcitrans (Diptera: Muscidae), in Denmark. Four species of pteromalids were found with Spalangia cameroni as the predominant. The other parasitoids were S. nigripes, S. nigra and Phygadeuon fumator (Ichneumonidae). Peak activity of the parasitoids was observed to be late in the summer and the beginning of autumn (August–September) when approximately 10% of the collected stable fly pupae were parasitised. Adult stable flies were infected with four species of entomopathogenic fungi: Entomophthora muscae, E. schizophorae, Beauveria bassiana and Verticillium lecanii. All fungi occurred in low percentages (max. 4%) and remained at this level throughout the sampling period. Likewise, adult house flies were infected with B. bassiana and V. lecanii,but Metarhizium anisopliae, Paecilomyces fumosoroseus and V. fusisporum were also recorded. The overall hyphomycete prevalence in house flies was 0.3%, and single species rarely exceeded 0.1%. The prevalence remained low in spite of increasing house fly numbers in August–September.     

Effect of Diachasmimorpha tryoni on two non-target flowerhead-feedingtephritids

Laboratory tests were conducted to evaluate the possible effect of a deliberately introduced fruit fly parasitoid, Diachasmimorpha tryoni, on 2 non-target flowerhead-feeding tephritid flies,Trupanea dubautiae and Ensina sonchi. The former is an nativecomposite endemic Hawaiian tephritid which feeds on flowerheads of the native composite shrub, Dubautia raillar dioides; the latter is an inadvertently introduced tephritid infesting flowerheads of the exotic weed, Sonchus oleraceus. Gravid females of D. tryoniwere confined in test cages with field-collected D.raillardioides and S. oleraceus flowerheads infested with late instars of T. dubautiae and E. sonchi,respectively. D. tryoni showed low levels of visiting and probing responses to D. raillardioides flowerheads and relatively higher responses to S. oleraceus flower heads in both the presence and absence of the parasitoid's normal host, the Mediterranean fruit fly, Ceratitis capitata. With 72-h exposure to D. tryoni, 13.9 and 2.6% of T. dubautiaewere attacked by test parasitoids in the absence and presence of C.capitata in the test cage, respectively; while 56.8 and34.2% of E. sonchi were attacked. In contrast, 94.0 and84.0% of C. capitata larvae presented in screened disheswith diet in test cages were attacked by D. tryoni in testswith T. dubautiae and E. sonchi, respectively. Attackof D. tryoni on flowerhead-feeding T. dubautiae andE. sonchi resulted in significant reduction in the emergence ofadult flies, especially in the absence of the parasitoid's normal host.While 8.8–12.8 adult D. tryoni per test (both males andfemales) successfully emerged from medflies, no adult D. tryoniprogeny emerged from T. dubautiae, and only 3 deformed males ofD. tryoni (much smaller than the normal wasp) emerged fromE. sonchi. The relevance of these findings to the safety offuture biological control programs against tephritid pests is discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Pollination of sevenPlectranthus spp. (Lamiaceae) in southern Natal, South Africa

The genusPlectranthus (Lamiaceae) shows remarkable radiation on the sandstones of southern Natal and northern Transkei in South Africa, where six endemic species occur. Two of these endemic species,P. hilliardiae andP. oribiensis, are included in this study, as well asP. reflexus, for which only limited data are available. The other species that were studied areP. ambiguus, P. ciliatus, P. ecklonii, P. madagascariensis andP. zuluensis. Four of these taxa,P. ambiguus, P. hilliardiae, P. reflexus andP. saccatus var.longitubus, have uniquely long corolla-tubes (20–30mm) and this is related to pollination by nemestrinid flies of the genusStenobasipteron that have proboscides of similar length. Other nemestrinid species of the genusProsoeca have shorter proboscides and pollinate two species ofPlectranthus with shorter corolla-tube lengths (6–15mm). Acrocerid flies, tabanid flies and anthophorid bees are also important visitors to these species. This study on the pollination of seven species of varying corolla-tube lengths shows a correlation between floral tube length and proboscis length of insect visitors, many of which are recorded for the first time as pollinators ofPlectranthus.     

Taxonomy, attack strategies and host relations in flowerhead Tephritidae: a review

Abstract. 1. Host relations of Palaearctic flowerhead-inhabiting tephritids (Diptera: Tephritidae) are reviewed in terms of the interaction between type of attack strategy and flowerhead size.
2. Three types of attack strategy are recognized, each associated with a particular phase of Compositae flowerhead development. These strategies are characteristic of a tribe or subfamily, viz the Tephritini (Tephritinae), Terelliini (Tephritinae) and Myopitinae.
3. Tephritini species show the most generalized feeding method, and this enables them to attack both large and very small flowerheads. In contrast, the more specialized requirements of members of the Terelliini and Myopitinae apparently restricts these groups to plant taxa with large flowerheads. As a result, over the whole range of Compositae flowerhead sizes, there exists a positive relationship between flowerhead size and the diversity of associated tephritids.
4. The occurrence of stem-feeding tephritids, and differences between a coevolutionary and functional (colonization) explanation for tephritid host-relations are discussed.     

Localization of mating behavior of releasedBactrocera dorsalis flies on host fruit in an orchard

Oriental fruit flies,Bactrocera dorsalis, cultured in the laboratory for six generations, were released when 12–14 days old in an orchard of nonfruiting host trees that were furnished with either food and water, nonpunctured host fruit, punctured host fruit, or no resources. Nearly all flies of both sexes, all sexual behavior of males, and all mating pairs were observed on trees with fruit. Moreover, on trees with fruit, nearly all flies of both sexes, nearly all sexual behavior of males, and nearly all mating pairs occurred on the fruit itself rather than on the foliage or branches. In a subsequent test, both sexes were found to be strongly attracted to the odor of host fruit. For a polyphagous species of a tropical tephritid, these findings are the first to show a high level of male aggregation and a high amount of male sexual behavior on the fruit of host trees. Findings are discussed in relation to current knowledge of tephritid mating behavior. An additional quality-control test for laboratory cultured males used in the sterile insect technique of fly management is recommended.     

Field Studies of Entomophthora (Zygomycetes: Entomophthorales)—Induced Behavioral Fever in Musca domestica (Diptera: Muscidae) in Denmark

House flies were collected over 3 days (three to five times per day) from specific sites on a dairy farm with a range of high to low temperatures. Flies were held individually to determine whether the distribution of fungus-infected (Entomophthora muscae and E. schizophorae) house flies differed according to the stage of infection and temperature. All but 2 of 396 infected flies (99.5%) had E. muscae. More E. muscae-infected flies collected from cool areas were in later stages of infection (i.e., dying 0–2 days after capture), whereas flies collected on sun-exposed surfaces tended to be in earlier stages of infection (i.e., dying 6–8 days after capture). Most flies died 3–5 days after capture and were consequently in the middle stages of infection. A mark and release experiment was conducted to determine whether E. schizophorae-inoculated flies frequented surfaces with higher temperatures than did uninfected control flies. About 3000 yellow-marked house flies inoculated with E. schizophorae and 3000 blue-marked control flies were released in an enclosed swine farrowing barn. Significantly more inoculated flies were recorded on the heat lamps than flies in the control group. The results suggest that behavioral fever occurs in the field for flies infected with both E. muscae and E. schizophorae and that flies can cure themselves of infection through the use of artificial heat sources.     

Successful oviposition and reproductive biology of Aprostocetus vaquitarum (Hymenoptera: Eulophidae): A predator of Diaprepes abbreviatus (Coleoptera: Curculionidae)

Aprostocetus vaquitarum (Wolcott) causes 78–91 percent mortality to eggs of Diaprepes abbreviatus (L.), under field conditions in southern Florida. In the laboratory, A. vaquitarum was reared on D. abbreviatus eggs at 25 °C, a photoperiod of 12:12 (L:D) and with abundant hosts, A. vaquitarum adult females lived around 15 days. Oviposition was significantly affected by the age of the host egg mass. Egg masses aged 0- to 3-day-old were accepted significantly better than those aged 4–6 days. The mean number of eggs deposited per female was around 53, with extreme values of 124 and 19 eggs per female. Using these data in combination with the sex ratio observed in the field (0.16) and the duration of the preimaginal stages, r_m (0.168–0142 day⁻¹), T (22.39–22.89 days), and R₀ (43.03–25.81 females per female) were calculated.     

Changes in abundance of water birds species in southern Bohemia (Czech Republic) in the last 10 years

In 1981–1992, the breeding fauna of 153 fishponds was studies in three fishpond regions in south Bohemia (Czechoslovakia). On each pond, all water and wetland birds were censused by the Two-check method, which consists of adult birds counts during each breeding season (the first in the second part of May, and the second in the second part of June). We assessed the numbers of all species living in water and wetland habitats in the breeding season.The decline in abundances (expressed as number of individuals) of ducks (Anatinae), grebes (Podicipediformes), Coot (Fulica atra), Moorhen (Galinulla chloropus), Black-headed Gull (Larus ridibundus) were the most marked changes recorded. Marked increase in abundances was found in Cormorant (Phalacrocorax carbo) and Mute Swan (Cygnus olor) during the whole decade, whereas in Graylag Goose (Anser anser) the increase was restricted to the first half of the period under study. Numbers of charadriform and passeriform birds fluctuated in the dependence of water level variation.     

Commensals of Mytilus galloprovincialis in the Black Sea: Urastoma cyprinae (Turbellaria) and Polydora ciliata (Polychaeta)

The intensity of infection of the mussel Mytilus galloprovincialis Lamarck in the Black Sea by the turbellarian Urastoma cyprinae (Graff), which lives on its gills, was found to be higher in larger hosts, reaching a maximum in mussels of 50–70 mm length. Greater numbers occured in mussels inhabiting a silty bottom than in cultivated mussels suspended above the bottom. Over the period 1982–1987, U. cyprinae was most numerous in winter and especially so in years that were colder. The spionid polychaete Polydora ciliata (Johnston) also infects M. galloprovincialis, burrowing into the shell. Young spionids of up to 1 mm length occured in mussels with a shell length of 35 mm. Numbers of this commensal reached a maximum in mussels of intermediate size.     

Development and reproduction of Stratiolaelaps scimitus (Acari: Laelapidae) with fungus gnat larvae (Diptera: Sciaridae), potworms (Oligochaeta: Enchytraeidae) or Sancassania aff. sphaerogaster (Acari: Acaridae) as the sole food source

Stratiolaelaps ( = Hypoaspis) miles (Berlese) (Acari: Mesostigmata: Laelapidae) is a polyphagous soil-dwelling predatory mite that is widely marketed for use in greenhouse production systems to manage populations of dark-winged fungus gnats, Bradysia spp. (Diptera: Sciaridae) and for supplemental control of thrips. The suggestion by Walter and Campbell (2003, Biol. Control 26: 253–269) that North American commercial cultures of S. miles may actually be S. scimitus was confirmed. The development and reproduction at 21–23 °C of S. scimitus provided ad libidum with one of three different prey – the fungus gnat Bradysia aff. coprophila (Lintner), potworms (Enchytraeidae), or Sancassania aff. sphaerogaster (Zachvatkin, 1937) (Acari: Astigmata: Acaridae) – were compared. Developmental duration of the egg and non-feeding larval stages were 2.47 and 1.11 days, respectively; mortalities were 8.3 and 5.5%. Stratiolaelaps scimitus failed to develop beyond the protonymphal stage when provided with S. aff. sphaerogaster alone, although some feeding was observed. Development and reproduction of S. scimitus was successful on both fungus gnat larvae and enchytraeids, with no influence of prey on protonymphal duration (4.70 days) and mortality (8.3%), or on deutonymphal duration (4.61 days) and mortality (6.1%). Adult female S. scimitus feeding on potworms, compared to those feeding on fungus gnat larvae, had a significantly shorter pre-oviposition period (2.69 vs. 4.59 days). However, diet did not influence other adult female developmental or reproductive characteristics (oviposition period, 18.6 days; post-oviposition period, 6.2 days; total adult longevity, 27.3 days; total number of eggs, 26.5). S. scimitus reared on potworms tended (p = 0.06) to have a higher intrinsic rate of increase, a higher finite rate of increase and a shorter doubling time (r _m = 0.142 day⁻¹, λ = 1.153, Dt = 4.85 days) than those reared on fungus gnat larvae (r _m = 0.105 day⁻¹, λ = 1.110, Dt = 6.58 days), but differences in net reproductive rate (R ₀) and generation time (G) were not significant.     

Flowering biology ofGeonoma irena andG. cuneata var.sodiroi (Arecaceae)

The phenology and flowering of two sympatric understory palms from western Ecuador, until recently regarded as conspecific, is described.Geonoma irena Borchs. flowers throughout the year. Anthesis lasts 11–14 weeks per inflorescence, with overlapping male and female phases. Flowers open in the midmorning, and are visited mainly by meliponid and halictid bees.Geonoma cuneata var.sodiroi (Burret)Skov flowers from December to April, with a distinct peak in Feburary. Anthesis lasts 6–8 days per inflorescence, with non-overlapping male and female phases. Flowers open at dawn, and are visited mainly by drosophilid and sphaerocerid flies. The flowering pattern of the two species match different specific behavioural features of their insect visitors. The study provides an example of how differences in reproductive biology may act as an important barrier to gene flow between related, co-occurring taxa.     

Suboptimal oviposition of tephritid flies supports parasitoid wasps

1. Female insects lay eggs on low‐quality plants, resulting in progeny with no or low reproductive potential; this is referred to as ‘suboptimal oviposition’ in this study. 2. Here, it is reported that suboptimal oviposition of a tephritid fly species (Tephritis femoralis) supports the populations of parasitoid wasp species in a Tibetan alpine meadow. 3. Suboptimal oviposition was confirmed because the adult flies emerging from Asteraceae species Anaphalis flavescens flowerheads did not survive cold winters. 4. DNA barcoding of puparia indicated that tephritid larvae developing from suboptimal oviposition hosted populations of at least five parasitoid wasp species. Consequently, suboptimal oviposition has significant effects on parasitoid population dynamics and species diversity.