Seasonal biology and abundance of Psyllaephagus pistaciae,a biocontrol agent of the common pistachio psylla Agonoscena pistaciae

The solitary endoparasitoid Psyllaephagus pistaciae Ferrière (Hymenoptera: Encyrtidae), is the most widely distributed biological control agent of the common pistachio psyllid, Agonoscena pistaciae Burckhardt and Lauterer (Hemiptera: Psylloidea), in Iran. The pupation and overwintering sites of diapausing parasitoids and the psyllid were studied for 2 years using emergence traps in pistachio orchards in Rafsanjan, Iran. The psyllid mummies containing the overwintering parasitoid adhered to pistachio leaves and were carried on these leaves away from the tree when they latter senesced. The present results verified that plant litter which included dried grasses and old pistachio leaves tended to support a greater population of adult winter-form psyllid and psyllid mummies during the winter through early spring than other options examined. Adult parasitoids appeared in the field in early April, about 30 days after the emergence of adult psyllids, but almost at the same time as the hatching of the first generation psyllid nymphs in early April. Rates of parasitism of CPP were generally low throughout most of the year, ranging from 1 to 5%, but rose in late autumn to about 11%. Results suggest that the density of P. pistaciae is not great enough to keep pace with the psyllid populations in these orchards. They explain why growers consider it necessary to apply pesticides for this pest. However, this parasitoid undoubtedly does play an important role in the natural control of A. pistaciae late in the growing season, particularly in non-sprayed orchards. Conservation of these natural enemies should be one of the objectives in the development of sustainable pest management programs.     

Life history parameters of the coccinellid beetle, Oenopia conglobata contaminata, an important predator of the common pistachio psylla, Agonoscena pistaciae (Hemiptera: Psylloidea)

The predatory beetle, Oenopia conglobata contaminata, is associated with the common pistachio psylla, Agonoscena pistaciae, the major pistachio pest in Iran. Successful development and reproduction on both A. pistaciae and Aphis gossypii (the major weed aphid in the pistachio orchards) indicated they were suitable prey for O. conglobata contaminata. Under ample prey supply, larval development on A. pistaciae was shorter and mortality was lower compared to those reared on A. gossypii. Furthermore, this ladybird attacks and destroys a large number of psyllid nymphs during the whole of its larval period (620±17 fourth stage nymphs) and also in its adult stage (191±7.5 4^th stage nymphs daily). The optimum temperature for development was 30°C, the theoretical threshold for development was 13°C and thermal requirements from egg to adult was estimated as 196 degree-days (°D). Fecundity for the first 21 days of adult life was 387 and 355 eggs when females fed on A. pistaciae and A. gossypii nymphs, respectively. The intrinsic rate of increase (r_m) at 27.5°C was 0.19 and 0.18 when ladybirds were fed on psyllid and aphid nymphs, respectively. In a laboratory choice experiment, the adult ladybirds showed a strong preference for A. pistaciae compared to A. gossypii. This was in agreement with our field observation that O. conglobata contaminata is almost always found on psyllid colonies, even when aphids were plentiful on weeds in the pistachio orchards. The influence of the predator on psyllid seasonal population under natural condition was not studied in this investigation and this subject must be measured in subsequent trials.     

Parasitoids associated with the common pistachio psylla, Agonoscena pistaciae, in Iran

The parasitoids associated with the common pistachio psylla, Agonoscena pistaciae Burckhardt and Lauterer, were investigated at three pistachio plantations in Rafsanjan, Iran. Of the 6504 wasps emerging from mummified psyllids, 46% were the primary parasitoid Psyllaephagus pistaciae Ferrière, and the remaining 54% represented six species of hymenopterous hyperparasitoids, including Chartocerus kurdjumovi (Nikol’skaja), Marietta picta (André), Pachyneuron aphidis (Bouché), Pachyneuron muscarum (Linnaeus), Psyllaphycus diaphorinae (Hayat), and Syrphophagus aphidivorus (Mayr). Lysiphlebus fabarum Marshall, the parasitoid of Aphis gossypii Glover and Aphis craccivora Koch present on weeds, was found to be an alternative host for three major hyperparasitoids of A. pistaciae. The most abundant hyperparasitoid was S. aphidivorus, appearing during the growing season in all trial locations on psyllids and aphids in pistachio orchards. The weed-infesting aphids, along with their primary parasitoid, can act as a reservoir of A. pistaciae secondary parasitoids. Therefore, parasitized aphids allow populations of secondary parasitoids to increase and consequently to apply higher pressure on P. pistaciae. We detected that two primary parasitoid species, including P. pistaciae and L. fabarum, attacking different species of hosts interact indirectly through shared secondary parasitism. It is suggested that the community structure of A. pistaciae may be influenced by apparent competition, although more work is needed to provide firm evidence.     

Demographic study of the reproductive potential of pear psylla, Cacopsylla pyri

Longevity and fecundity of pear psylla adults, Cacopsylla pyri (L.) (Homoptera: Psyllidae), were studied in Greece under field conditions, during 1991/92 and 1992/93. Net fecundity (mean fecundity of cohort) of the overwintered females (winter form) was low (24.1 and 43.4 eggs per female for 1991/92 and 1992/93 respectively) because only a small fraction of the females survive reproductive diapause. Surviving females, however, are capable of producing a relatively high number of eggs (135.0 and 151 9 eggs per female for the two seasons respectively). Net fecundity of summer form females was very high in spring (343.2 and 406.8 eggs per female for 1992 and 1993 respectively), relatively high in early autumn (130.4 and 192.5 eggs per female) and very low in summer (21.9 and 46.5 eggs per female). Male longevity was in all cases shorter than that of female. Longevity of both sexes of the summer form was higher in spring and autumn (for females it ranged from 22.3 to 28.4 days) than it was in summer (for females it ranged from 14.3 to 17.4 days).     

The seasonal cycle of Thermocyclops crassus (Fischer, 1853) (Copepoda: Cyclopoida) in a shallow,eutrophic lake

The seasonal cycle of Thermocyclops crassus was studied from 1985 to 1987, in the Gronne, a shallow, productive lake. T. crassus was present from late April to early October, while water temperature was above 10 °C. It produced three generations per year. Population peak was usually reached by the second generation, in August. Abundance was positively correlated with water temperature. Females carried 18.3 to 32.3 eggs on average. Reproduction rates were highest in July and August. Sex ratio was low, as females generally outnumbered males. Between October and April copepodites 4 went in diapause, predominantly in the deepest part of the lake and 0–4 cm deep in the bottom mud. Variations in body size were low across the year.T. crassus coexisted with the cyclopids, Cyclops vicinus and Mesocyclops leuckarti. In 1985 and 1986, abundances of M. leuckarti were low, while those of T. crassus and C. vicinus were high. In 1987 lower water temperatures, caused by cold weather, resulted in a marked decrease of the population density of T. crassus, while the abundance of M. leuckarti increased, and density of C. vicinus remained high. Comparison of the egg duration times of T. crassus, C. vicinus and M. leuckarti showed that at 15 °C and below T. crassus may be outcompeted by its comparatively longer egg development times. However, experiments in limnocorrals showed that T. crassus has an advantage over C. vicinus when fish predation is high.Department of Biology III, University of Ulm     

Life history and demography of Psyllaephagus zdeneki (Hymenoptera: Encyrtidae), a potential candidate for biological control of olive psylla,Euphyllura pakistanica (Hemiptera: Psyllidae)

Life history and demographic parameters of Psyllaephagus zdeneki Noyes and Fallahzadeh (Hymenoptera: Encyrtidae) were studied on its host the olive psyllid, Euphyllura pakistanica Loginova (Hemiptera: Psyllidae). Experiments were conducted in a growth chamber at 20±1 °C, relative humidity of 60±5%, and a photoperiod of 16:8 (L: D) hours. Four different olive cultivars (Fishomi, Shenge, Oil and Yellow) were used to test possible host plant influence on parasitoid performance. The pre-imaginal developmental period of female P. zdeneki varied from 24.96 (on Fishomi) to 26.34 (on Shenge) days, and for males from 21.63 (on Fishomi) to 24.44 (on Yellow) days. Adult female longevity differed significantly among the four cultivars, ranging from 12.46 (on Fishomi) to 14.97 (on Shenge) days. For each cultivar, adult female longevity was significantly greater than male longevity. Life table parameters showed survival rates (l _x ) in newly emerged females were 84.61, 82.25, 85.71 and 78.12% on Fishomi, Yellow, Shenge and Oil, respectively. Female egg deposition was highest on Yellow (138.4 eggs per female) and lowest on Fishomi (116.3 eggs per female). The highest and lowest intrinsic rate of increase were 0.28 (on Shenge) and 0.24 (on Oil), respectively. The mean generation time ranged from 14.6 (on Shenge) to 16.7 (on Oil) days. These results are discussed with respect to the potential impact P. zdeneki as a natural enemy of E. pakistanica, the most important pest of olive in the Fars province of Iran, as well as the influence of olive cultivar on parasitoid life table parameters.     

Diapause and seasonal life cycle strategy in the house spider, Achaearanea tepidariorum (Araneae, Theridiidae)

Abstract In order to elucidate the mechanism regulating its seasonal life cycle, the photoperiodic response of Achaearanea tepidariorum has been analysed. Nymphal development was faster in long-day and slower in short-day photoperiods. The combined action of low temperature, poor food supply and short daylength induced diapause at an earlier developmental stage than short days alone. Thus, photoperiod is a primary factor inducing nymphal diapause, but the diapausing instar is influenced by both temperature and food supply. Hibernating nymphs became unresponsive to photoperiod in late December. After hibernation, however, sensitivity was restored and the nymphs remained sensitive to photoperiod throughout their life. This spider could also enter an imaginal or reproductive diapause. Photoperiod was again a primary inducing factor and temperature modified the photoperiodic response to some extent. The induction of the reproductive diapause was almost temperature-compensated whereas development was not. So the involvement of a photoperiodic counter system was suggested. Irrespective of whether the nymph had experienced diapause or not, the imaginal diapause was induced in response to a short-day photoperiod after adult moult. Based on these observations, the seasonal life cycle and the adaptive significance of nymphal and imaginal diapause are discussed.     

Interactions between citrus psylla,Trioza erytreae (Hem. Triozidae), and spiders in an unsprayed citrus orchard in the transvaal lowveld

During a two year survey, a total of 3,054 spiders represented by 21 families were sampled in an unsprayed citrus orchard in the Transvaal Lowveld. Numerically the Salticidae was the dominant family (34.4%) followed by the Theridiidae (21.9%), Thomisidae (11.9%), Araneidae (7.9%), Clubionidae (7.0%) and the Tetragnathidae (3.7%). Eighteen species of spiders were observed to prey on citrus psylla,Trioza erytreae (Del Guercio) (Hemiptera: Triozidae), while six species trap nymphs and adults under their retreats and webs. There were significant positive correlations between the weekly psylla populations and the weekly populations of web-building spiders and wandering spiders present one to four weeks later but no significant correlation between the weekly spider populations and the weekly psylla populations present one to five weeks later. This seems to indicate that while spiders are unable to keep citrus psylla populations at acceptable low levels, they may contribute in reducing their numbers.      

The problems and prospects of integrated control of Agonoscena pistaciae Burck. and Laut. (Hom., Sternorrhyncha) in Greece

Abstract: A study of the role of natural enemies of Agonoscena pistaciae Burck. and Laut. populations, in a pistachio orchard located in the Makrakomi area of Central Greece, was carried out during 1999–2000. Agonoscena pistaciae is economically the most important psyllid species of pistachio in Greece and causes considerable damage. Its adult population evaluation is clearly distinguishable in two phases. The first phase, from early April until the end of July, is characterized by slow population development. In the second phase, from the end of July until the fall of the leaves, the population density of this insect increases rapidly and reaches high levels. During this second phase, the presence of the predator insects Chrysoperla carnea Steph., Anthocoris nemoralis F and of the parasitoid Psyllaephagus pistaciae Ferriere is noteworthy. Among these, the most important is the parasitoid, which appears in satisfactory numbers between the end of July and the end of September/beginning of October and plays a significant role in reducing the populations of A. pistaciae that are on the point of hibernating.     

Interplant movement by pear psylla (Homoptera: Psyllidae): Effects of sex ratio and reproductive status

Sticky trap catch of pear psylla,Cacopsylla pyricola Foerster, is male biased during the reproductive generations, but not the diapausing generation. In cage studies, we monitored movement by male and female pear psylla between host plants, and tested whether reproductive and diapausing psylla exhibit similar rates of movement. We also experimentally varied sex ratio to determine whether sex ratio affected movement. Male-biased sex ratios prompted increased movement off of the original host by male psylla of the reproductive generations; no such effect was noted for diapausing insects. We interpret these results to indicate that male movements increased under male-biased conditions due to mate-searching activities. There was also evidence in two experiments that severely male-biased sex ratios prompted movement off of the original host plant by reproductive females; this effect may have been due to harassment of ovipositing females by males.     

Nettle extract effects on the aphid Myzus persicae and its natural enemy, the predator Macrolophus pygmaeus (Hem., Miridae)

Abstract:  The effect of water extract of Urtica urens L. on the biological characteristics and population parameters of Myzus persicae (Sulzer) and its natural enemy, the polyphagous predator Macrolophus pygmaeus (Rambur) was investigated on eggplants. Plant material was extracted in deionized water (1 : 5 w/w) for 24 h. Three applications were made at 5-day intervals using the nettle extract, with deionized water as the control. The effect of the extract on survival, duration of nymphal development, pre-reproductive period, fecundity and longevity of both insects was studied at conditions of 25 ± 1°C, 65 ± 5% relative humidity and photoperiod of 16 : 8 h (light : dark). The results showed that the application of nettle extract significantly reduced the fecundity of M. persicae (20% on average) but did not result in a substantial lowering in its intrinsic rate of population increase. However, the predator's biological characteristics and population parameters were not affected by the application of this plant extract. The importance of these results for the use of nettle extract in the management of M. persicae along with its compatibility with the use or conservation of M. pygmaeus is discussed.     

Modelling age‐ and density‐structured reproductive biology and seasonal survival of Apanteles galleriae Wilkinson (Hym., Braconidae)

Abstract: The biology of Apanteles galleriae Wilkinson, an important biological control agent of wax moths, is well described in the literature. We developed models simulating the functional response of fecundity and female progeny proportion of adult females as a function of age, host and/or parasitoid density to integrate current knowledge. Daily pattern of age‐related fecundity and female progeny proportion of different parasitoids was also examined. We investigated the effect of sex, mating status and seasonal time on adult longevity. We derived survival data of females in two different seasonal periods. We found that 50% of adult life span is important for an efficient fecundity. The patterns of host and/or parasitoid density‐dependent fecundity and sex ratio varied considerably. The highest fecundity and female progeny proportion occurred with one parasitoid and one female equivalent host. Longevity of adults was affected by sex and month periods, but mating status did not affect their longevity. Female survival was greater between December and May relative to June and November. Our results indicated that age, host and parasitoid density, and the timing of rearing influenced the life processes of parasitoids.     

Diapause induction in Trichogramma embryophagum Htg. (Hym., Trichogrammatidae): the dynamics of thermosensitivity

It is known that the low temperature is the most important factor inducing the pre-pupal diapause in Trichogramma species. The position of the thermosensitive period over the life cycle and temporal variation of the degree of responsiveness were investigated in T. embryophagum Htg. by transferring pre-imaginal stages between 'neutral' temperature of 15°C and 'diapause-inducing' temperature of 10°C. Our experiments showed that 6 days long exposure at 10°C significantly increased the percentage of diapausing pre-pupae when started during rather large part of development: from embryo up to early pre-pupa. The highest thermosensitivity was recorded during the embryo and the larval stages, with some decrease during the hatching period. Treatments with shorter cold exposures (2–3 days) gave similar results. Even 24 h long exposure at 10°C increased the percentage of diapausing pre-pupae when applied during egg or early larval stage. Being started at the same stage of development, longer cold exposures caused stronger increase in the percentage of diapausing individuals. The experiments did not reveal any significant daily changes in thermosensitivity: at 12 : 12 h light : dark, larvae subjected to the low temperature during six photophases showed practically the same percentage of diapausing individuals as those subjected to the low temperature during six scotophases, and as those subjected to the 3 days long uninterrupted cold exposure. Hence, in natural conditions even occasional short-term cold periods could be accumulated.     

Antioxidant activity and phenolic profile of pistachio (Pistacia vera L., variety Bronte) seeds and skins

Pistachio (Pistacia vera L.; Anacardiaceae) is native of aride zones of Central and West Asia and distributed throughout the Mediterranean basin. In Italy, a pistachio cultivar of high quality is typical of Bronte (Sicily), an area around the Etna volcano, where the lava land and climate allow the production of a nut with intense green colour and aromatic taste, very appreciated in international markets. Pistachio nuts are a rich source of phenolic compounds, and have recently been ranked among the first 50 food products highest in antioxidant potential. Pistachio nuts are often used after removing the skin, which thus represents a significant by-product of pistachio industrial processing. The present study was carried out to better characterize the phenolic composition and the antioxidant activity of Bronte pistachios, with the particular aim to evaluate the differences between pistachio seeds and skins. The total content of phenolic compounds in pistachios was shown to be significantly higher in skins than in seeds. By HPLC analysis, gallic acid, catechin, eriodictyol-7-O-glucoside, naringenin-7-O-neohesperidoside, quercetin-3-O-rutinoside and eriodictyol were found both in pistachio seeds than in skins; furthermore, genistein-7-O-glucoside, genistein, daidzein and apigenin appeared to be present only in pistachio seeds, while epicatechin, quercetin, naringenin, luteolin, kaempferol, cyanidin-3-O-galactoside and cyanidin-3-O-glucoside are contained only in pistachio skins. The antioxidant activity of pistachio seeds and skins were determined by means of four different assays (DPPH assay, Folin-Ciocalteau colorimetric method and TEAC assay, SOD-mimetic assay). As expected on the basis of the chemical analyses, pistachio skins have shown to possess a better activity with respect to seeds in all tests. The excellent antioxidant activity of pistachio skins can be explained by its higher content of antioxidant phenolic compounds. By HPLC-TLC analysis, gallic acid, catechin, cyanidin-3-O-galactoside, eriodictyol-7-O-glucoside and epicatechin appeared to be responsible for the antioxidant activity of pistachio skin, together with other unidentified compounds. In conclusion, our work has contributed to clarify some particular characteristics of Bronte pistachios and the specific antioxidant power of pistachio skins. Introduction of pistachios in daily diet may be of undoubted utility to protect human health and well-being against cancer, inflammatory diseases, cardiovascular pathologies and, more generally, pathological conditions related to free radical overproduction. On the other hand, pistachio skins could be successfully employed in food, cosmetic and pharmaceutical industry.     

Human‐induced changes in the reproductive traits of Lake Constance common whitefish (Coregonus lavaretus)

Size‐selective fishery harvest leads to phenotypic changes in fish reproductive traits. When these changes represent an evolutionary response of a stock, they may have severe consequences for future stock dynamics and yields. In freshwater ecosystems, reproductive traits may also be affected by other human impacts such as changes in system productivity. The present study uses regression analysis to evaluate the impacts of changes in lake trophy and of an intensive size‐selective harvest over several decades on the reproductive traits of common whitefish in Lake Constance between 1963 and 1999. Fecundity was strongly linked to lake trophy but was also related to the calendar year, suggesting an evolutionary response to size‐selective harvest and to massive stocking of the lake with hatchery‐reared larvae. The present study is an example of how fish reproductive traits are influenced by the combined action of various human impacts: changes in system productivity, size‐selective harvest and massive stocking.     

Morphometry of the sexual forms of Acromyrmex subterraneus brunneus Forel, 1911 (Hym., Formicidae) in queenright and queenless laboratory colonies

Abstract:  A morphometric study was performed on the sexual forms of Acromyrmex subterraneus brunneus in colonies kept under laboratory conditions. The males studied were obtained from queenright and queenless colonies, i.e. they were produced by either the queen or the workers. Cluster analysis revealed a wide distribution in the size of the sexual forms produced in queenless colonies compared with queenright colonies, and that some of the worker-produced males were significantly smaller than those produced in queenright colonies. However, we found no indication that the males produced in queenless colonies are unable to mate, as smaller and larger males had identical genitalia that varied only in size, and therefore probably represent an alternative route of reproduction in this species.     

The postmarsupial development of Porcellio siculoccidentalis, with some data on reproductive biology (Crustacea, Isopoda, Oniscidea)

In the broader context of research on the Sicilian Porcellio imbutus-complex, the postmarsupial development of Porcellio siculoccidentalis Viglianisi, Lombardo & Caruso, 1992 was studied in detail. This research was conducted in the laboratory under controlled conditions, allowing us to follow the stages of development, from the formation of the marsupium in ovigerous females until the larval stages and development of the seventh pair of legs. The timing of developmental stages and the morphological modifications of appendages in the postmarsupial manca stages (M I-M III) are described. The manca stage M I had a duration of about one hour. Ovigerous females were collected and reared separately, and the number of parturial molts in the absence of males was counted. The results showed a maximum of four successive parturial molts. Fecundity and fertility were evaluated as the number of eggs and embryos, respectively, inside the marsupium of the ovigerous females. Both parameters were positively correlated with the size of the females. The maximum numbers of eggs and embryos in the marsupium were 113 and 141, respectively. Data describing the total number of postmarsupial mancas released per month indicated that the highest release occurred in April.     

Variation in diapause potential and strength in eggs of the Australian plague locust, Chortoicetes terminifera (Walker) (Orthoptera: Acrididae)

Samples of eggs of Chortoicetes terminifera were incubated under 3 temperature regimes which would allow non-diapause eggs to develop about 50% and so take them beyond the diapause stage. Even so, many more eggs entered diapause when reared at 20°C for 3 weeks than at 32°C for 1 week. By collecting and incubating eggs at intervals after laying in autumn, diapause potential or strength of eggs at different stages of development was estimated. For eggs in pre-diapause, the proportion in diapause after rearing was used to estimate diapause “potential”; for those in diapause, the proportion was used to estimate diapause “strength”. At laying, eggs varied greatly in their potential to enter diapause. However, those with lower potential at laying often increased their potential during the pre-diapause stage so that by the time diapause was reached diapause strength varied but over a lesser range. In all eggbeds, diapause strength decreased by 7–9 weeks after laying; little or none remained by mid-winter.These variations in diapause potential and diapause strength seem to reflect how much the temperature threshold for development during diapause is increased above that for non-diapause. Both diapause potential and strength may reflect the value of some factor whose level at laying is determined by the environment experienced by the parents but which changes subsequently.     

Influence of thermoperiod and photoperiod on reproductive diapause in Bruchidius atrolineatus (Pic) (Coleoptera, Bruchidae)

Abstract Bruchidius atrolineatus (Pic) is a common pest infesting cowpea seeds ( Vigna unguiculata Walp) in Africa. In the Sahelian zone, the adults are in reproductive diapause during the dry season. Diapause induction depends on the climatic conditions during embryonic and post-embryonic development. A strain of B. atrolineatus originating from the Niamey (13^oN) was reared in different thermoperiodic and photoperiodic conditions. In the thermoperiodic regime 40:25^oC, in continuous darkness, induction of diapause was dependent on the duration of the thermophase. Photoperiod could also influence diapause induction but the response depended on the temperature. When the beetles were reared in conditions of LD 12:12 h, 40:25^oC, for seven generations, the proportion of diapausing adults decreased and their sensitivity to photoperiod increased. In LD 14:10 h, 40:25^CC, the proportion of diapausing adults remained high regardless of the photoperiod. In these conditions, the first emerging adults were sexually active and were used for selection of a strain with a low incidence of diapause. After twelve generations, the proportion of diapausing beetles was lower than 5%. The responses of this selected strain to photo- and thermoperiods were reduced. The sensitivity of B. atrolineatus to photoperiod and thermoperiod could be controlled by genetic systems as observed in other insect species.