Temperature-dependent development, diapause and cold tolerance of Gratiana graminea, a potential biological control agent of Solanum viarum in Florida, USA

The objectives of this study were to examine temperature-dependent development, diapause and cold tolerance of Gratiana graminea Klug (Chrysomelidae), a candidate biological control agent of tropical soda apple, Solanum viarum Dunal (Solanaceae). Immature development was examined at six constant temperatures ranging from 15°C to 30°C. Diapause induction was determined by exposing adults to either long or short photoperiods at 20°C and cold tolerance was assessed by exposing adults to 0°C. G. graminea completed development at temperatures ranging from 20°C to 30°C. Linear regression estimated a lower temperature threshold of 11.7°C and 312 degree-days were required to complete development. Diapause was induced when adults were exposed to short photoperiods (10:14 L:D h) at 20°C. The lethal times for diapausing adults of G. graminea at 0°C (LT₅₀?=?19?days, LT₉₀?=?41?days) were two times higher compared to Gratiana boliviana Spaeth, a biological control agent already established in south and central Florida, USA. The presence of diapause and the greater cold tolerance suggest that G. graminea may establish and perform better than G. boliviana in northern Florida.     

Effect of temperature on the life history and demographic parameters of Spalgis epius (Lepidoptera: Lycaenidae), a candidate biological control agent of mealybugs (Hemiptera: Pseudococcidae)

Many species of mealybugs (Hemiptera: Pseudococcidae) are serious pests of economically important crops worldwide. We evaluated the influence of constant temperatures: 14, 16, 18, 20, 22, 24, 26, 28, 30, 32 and 34°C on the life history and demographic parameters of Spalgis epius (Lepidoptera: Lycaenidae), a candidate biological control agent of various species of mealybugs. No eggs completed their development at 14 and 34°C. Egg-to-adult developmental time significantly decreased from 89.9 days at 16°C to 20.4 days at 32°C. The estimated lower temperature threshold of 10.2°C and 416.6 degree-days were required to complete egg-to-adult development. The mortality of immature stages was maximum at 16 and 32°C and minimum at 28°C. The highest lifetime fecundity was recorded at 28°C and it significantly decreased at 32°C. The longevity of adults was about three times more at 16°C than at 30 and 32°C. The net reproductive rate (R ₀) significantly increased with increased temperatures up to 28°C and significantly decreased at 32°C. The mean generation time (T) significantly decreased with increased temperature up to 30°C, but it significantly increased at 32°C. The intrinsic rate of population increase (r _m ) was highest at 30°C. The finite rate of increase (λ) was significantly greater at 30°C than at other temperatures. These data suggest that S. epius can develop, reproduce and survive in a wide range of temperatures and thus could be regarded a potential biological control agent of mealybugs.     

Biological parameters of Elasmus nudus (Hymenoptera,Eulophidae), a parasitoid of the pistachio fruit hull borer moth,Arimania komaroffi (Lepidoptera,Pyralidae)

Elasmus nudus (Nees) is known as a synovigenic, gregarious ectoparasitoid of the pistachio fruit hull borer moth, Arimania komaroffi Ragonot in pistachio plantations in southern Iran. The biological parameters of this parasitoid were studied using the Mediterranean flour moth, Ephestia kuehniella Zeller as a host and the resulting data were compared with that of its native host, A. komaroffi. The parasitoid developed successfully from egg to adult over a temperature range of 17–35°C. The developmental times of the parasitoid on A. komaroffi and E. kuehniella were not significantly different. The threshold value at which the development rate of E. nudus was zero was estimated to be 9.2°C and the estimated number of degree days (°D) for development from egg to adult was 238 above this threshold. The reproductive potential of E. nudus was similar on both hosts at 30°C. The intrinsic rate of natural increase (r _m ) for E. nudus did not vary much between 25°C and 32.5°C. In addition, the clutch size of the parasitoid on the two hosts was not statistically different. The present study provides some basic information on the biological characteristics of E. nudus.     

Effect of temperature on the development and consumption of Phaenochilus kashaya (Coleoptera: Coccinellidae), a predator of the cycad aulacaspis scale,Aulacaspis yasumatsui

The cycad aulacaspis scale (CAS), Aulacaspis yasumatsui Takagi (Hemiptera: Diaspididae), is a serious pest of ornamental cycads in the southeastern United States. In Florida, CAS was first reported in Miami in 1996 and is now present in 43 Florida counties. Chemical control of CAS can be effective, but it is expensive, and insecticides must be frequently and regularly applied. Recent surveys of natural enemies in Thailand identified a new potential biological control agent of CAS. The objective of this study was to examine the temperature requirements and consumption rates of Phaenochilus kashaya Giorgi and Vandenberg (Coleoptera: Coccinellidae). Survival, developmental time and number of scales consumed by P. kashaya were measured at four constant temperatures (20, 25, 30 and 35°C). In addition, adult longevity and consumption were recorded at three constant temperatures (20, 25 and 30°C). Phaenochilus kashaya completed development to adulthood at temperatures ranging from 20 to 30°C. The highest survival rate (48%) was obtained at 25°C, and the accumulated degree-days for this species was 714. Phaenochilus kashaya is a voracious predator of CAS, consuming 380 scales during the larval stage and an estimated 4700 scales during the adult stage. Adult longevity varied from 59 days at 30°C to 220 days at 20°C for females, and 31 days at 30°C to 148 days at 20°C for males. The intrinsic rate of increase (r _m) was 0.34 at 20°C, which is high compared with other coccinellid species. According to this study, P. kashaya has the potential to be an effective biological control agent of the CAS in Florida.     

Threshold temperature and heat unit requirements for the development of the granary weevil,Sitophilus granarius (L.)

The objective of the present investigation was to determine the effects of temperature on the granary weevil, Sitophilus granarius L., in the laboratory so that forecasting models based on heat accumulation units could be developed for the pest. Development of S. granarius reared on wheat grains was studied at three constant temperatures (15, 20 and 25?°C) and relative humidity of 70?±?5%. The developmental time was significantly decreased with the increase in temperature. The developmental threshold temperatures estimated for egg, larva, pupa and from egg to adult were 7.79, 7.31, 14.08 and 9.97?°C, respectively. Based on these thresholds, the developmental stages, respectively, needed 92.75, 393.78, 45.04 and 562.36?day degrees to complete their development. Also, about nine generations were calculated for the pest per year under Assiut prevailing conditions. It seems that temperature of 25?°C is the most favourable developmental temperature.     

Assessing the effects of low temperature on the establishment potential in Britain of the non-native biological control agent Eretmocerus eremicus

Abstract. Eretmocerus eremicus is a parasitoid wasp that is not native to Britain. It is a biological control agent of glasshouse whitefly and has recently been released under licence in Britain for the first time. This study assessed the effect of low temperature on the outdoor establishment potential of E. eremicus in Britain. The developmental threshold calculated by three linear methods was between 6.1° and 11.6 °C, with a degree‐day requirement per generation between 256.3 and 366.8° day^?1. The supercooling points of non‐acclimated and acclimated larvae were similar (approximately ?25 °C). Non‐acclimated and acclimated larvae were subject to considerable pre‐freeze mortality, with lethal temperature (LTemp₅₀₎ values of ?16.3 and ?21.3 °C, respectively. Lethal time experiments indicated a similar lack of cold tolerance with 50% mortality of both non‐acclimated and acclimated larvae after 7 days at ?5 °C, 10 days at 0 °C and 13 days at 5 °C. Field trials showed that neither non‐acclimated nor acclimated larvae survived longer than 1 month when exposed to naturally fluctuating winter temperatures. These results suggest that releasing E. eremicus into British greenhouses would pose minimal risk because typical British winter temperatures would be an effective barrier against establishment in the wild.     

Thermal biology and establishment potential in temperate climates of the aphid parasitoid, <Emphasis Type="Italic">Lysiphlebus testaceipes</Emphasis>

Lysiphlebus testaceipes (Cresson) (Hymenoptera: Braconidae, Aphidiinae) is a parasitic wasp which plays an important role in the biological control of a number of aphid species. Through assessment of its thermal biology and low temperature tolerance, this study ascertains the establishment potential of L. testaceipes in cool temperate climates typical of northern Europe. The developmental threshold of L. testaceipes was 5.8°C. Rearing of parasitoids at shorter day lengths and lower temperatures indicated no ability to enter a diapause state. The supercooling points (SCP) of non-acclimated and acclimated parasitoid life stages were between −24.6°C and −17.7°C, with LTemp₅₀ temperatures approaching these values, indicating a high level of cold tolerance in short exposures. At 5°C the LTime₅₀ of acclimated larvae within parasitized aphids was 42.8 days. Acclimated pupae continued to develop with 54% adult emergence from mummies within 60 days. Acclimated parasitoid larvae and pupae, within living and mummified aphids, continued to develop during 70 days of winter field exposure and emerging adult parasitoids were reproductively viable under field conditions. These data indicate that where suitable host species are available throughout the year, L. testaceipes would be able to establish in northern Europe.     

Thermal biology of Typhlodromips montdorensis: implications for its introduction as a glasshouse biological control agent in the UK

The recent unexpected local establishment of a non‐native predatory mite, Neoseiulus californicus (McGregor) (Acari: Phytoseiidae), in the UK prompted us to undertake this study, which investigated the thermal biology of an alien species Typhlodromips montdorensis (Schicha) (Acari: Phytoseiidae). Laboratory and field experiments on its cold tolerance were used to assess its establishment potential outside of glasshouse environments in the UK. Currently, T. montdorensis is being tested as a glasshouse biological control agent against thrips and spider mites, but is not yet licensed for release in the UK. Typhlodromips montdorensis has a developmental threshold of between 10.3 and 10.7 °C, and a thermal budget of between 108.7 and 105.3 degree‐days when estimated by weighted and simple linear regression, respectively. Under outdoor conditions, T. montdorensis could theoretically complete up to six generations a year. The supercooling points of female and larval T. montdorensis were ?22 to ?24 °C with 100% pre‐freeze mortality apparent in both acute and chronic low temperature exposures. Typhlodromips montdorensis were unable to enter diapause under a selected laboratory regime. No reproduction occurred in the field from November to March, with 100% mortality within 7–14 days of release during this period. It is concluded that T. montdorensis would be a ‘safe candidate’ for introduction as a glasshouse biological control agent in the UK.     

Comparative assessment of the thermal tolerance of spotted stemborer,Chilo partellus (Lepidoptera: Crambidae) and its larval parasitoid,Cotesia sesamiae (Hymenoptera: Braconidae)

Under stressful thermal environments, insects adjust their behavior and physiology to maintain key life‐history activities and improve survival. For interacting species, mutual or antagonistic, thermal stress may affect the participants in differing ways, which may then affect the outcome of the ecological relationship. In agroecosystems, this may be the fate of relationships between insect pests and their antagonistic parasitoids under acute and chronic thermal variability. Against this background, we investigated the thermal tolerance of different developmental stages of Chilo partellus Swinhoe (Lepidoptera: Crambidae) and its larval parasitoid, Cotesia sesamiae Cameron (Hymenoptera: Braconidae) using both dynamic and static protocols. When exposed for 2 h to a static temperature, lower lethal temperatures ranged from ?9 to 6 °C, ?14 to ?2 °C, and ?1 to 4 °C while upper lethal temperatures ranged from 37 to 48 °C, 41 to 49 °C, and 36 to 39 °C for C. partellus eggs, larvae, and C. sesamiae adults, respectively. Faster heating rates improved critical thermal maxima (CT_max) in C. partellus larvae and adult C. partellus and C. sesamiae. Lower cooling rates improved critical thermal minima (CT_min) in C. partellus and C. sesamiae adults while compromising CT_min in C. partellus larvae. The mean supercooling points (SCPs) for C. partellus larvae, pupae, and adults were ?11.82 ± 1.78, ?10.43 ± 1.73 and ?15.75 ± 2.47, respectively. Heat knock‐down time (HKDT) and chill‐coma recovery time (CCRT) varied significantly between C. partellus larvae and adults. Larvae had higher HKDT than adults, while the latter recovered significantly faster following chill‐coma. Current results suggest developmental stage differences in C. partellus thermal tolerance (with respect to lethal temperatures and critical thermal limits) and a compromised temperature tolerance of parasitoid C. sesamiae relative to its host, suggesting potential asynchrony between host–parasitoid population phenology and consequently biocontrol efficacy under global change. These results have broad implications to biological pest management insect–natural enemy interactions under rapidly changing thermal environments.     

Thermal biology and establishment potential in temperate climates of the predatory mirid <Emphasis Type="Italic">Nesidiocoris tenuis</Emphasis>

Nesidiocoris tenuis Reuter (Hemiptera: Miridae) is a polyphagous mirid currently used for the control of leafminers, thrips, whitefly and spider mites in Mediterranean regions to which it is indigenous. This study investigates the establishment potential of N. tenuis in cool temperate climates typical of northern Europe through assessment of its thermal biology and low temperature tolerance in laboratory and field experiments. The developmental threshold of N. tenuis was estimated to be 12.9°C with no indication of ability to diapause. Supercooling points of the acclimated and non-acclimated adults and nymphs of the mirid were between −17.6° and −21.5°C and the LTemp₅₀ was around −12°C, indicating a high level of pre-freeze mortality. The LTime₅₀ at 5°C was nine days and 100% mortality occurred after less than four weeks of winter field exposure. Collectively these data suggest that N. tenuis is unlikely to establish in northern Europe and would therefore have little or no non-target effects on native species in such regions.     

Effect of temperature on the reproductive success,developmental rate and brood characteristics of Ips sexdentatus (Boern.)

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Estimating temperature‐dependent developmental rates of potato tuberworm,Phthorimaea operculella (Lepidoptera: Gelechiidae)

Abstract The potato tuberworm, Phthorimaea operculella (Zeller) (Lepidoptera: Gelechiidae), is the most destructive pest of potato, Solanum tuberosum L. (Solanaceae), in tropical and subtropical regions in both field and storeroom situations. The modeling of temperature‐dependent development can be useful in forecasting occurrence and population dynamics of the pests. Published developmental parameters for this pest vary greatly for many reasons. We determined temperature‐dependent development of P. operculella at seven constant temperatures (16, 20, 24, 28, 32, 34 and 36 °C). Developmental period of whole immature stage (egg to the end of the pupal stage) varied from 75.5 days at 16 °C to 17 days at 32 °C. The population failed to survive at 36 °C. The observed data was modeled to determine mathematical functions for simulating P. operculella development in each stage of development and overall. Two linear models, ordinary linear regression and the Ikemoto linear model were used to describe the relationship between temperature and development rate of the different stages of P. operculella and estimating the thermal constant and lower temperature threshold. The lower temperature threshold (t) and thermal constant (k) of whole immature stage were estimated to be 11.6 °C and 338.5 DD by Ikemoto linear model, and the estimated parameters were not substantially different with those estimated by ordinary linear models. Different models provided a better fit to the various developmental stages. Of the eleven nonlinear models fitted, the Beriere‐1, Logan‐6 and Lactin‐1 model was found to be the best for modeling development rate of egg, larva and pupa of P. operculella, respectively. Phenological models based on these findings can be part of a decision‐support tool to improve the efficiency of pest management programs.     

The interactions of tropical soda apple mosaic tobamovirus and Gratiana boliviana (Coleoptera: Chrysomelidae), an introduced biological control agent of tropical soda apple (Solanum viarum)

Tropical soda apple (Solanum viarum Dunal (Solanaceae) is a South American invasive plant of rangelands, pastures and natural areas in Florida. A chrysomelid beetle from South America, Gratiana boliviana Spaeth, has been released at >300 locations in Florida for biological control of tropical soda apple since 2003. Tropical soda apple is a host of several plant viruses, including the newly described tropical soda apple mosaic virus (TSAMV). We investigated the influence of TSAMV infection of tropical soda apple plants on developmental time, leaf tissue consumption, longevity, fecundity, and feeding preference of G. boliviana, and also tested transmission of the virus by the beetle. Developmental time was approximately 10% slower, and adults consumed only about 50% as much leaf tissue, for beetles fed on infected plants compared to uninfected plants. Longevity did not differ between females reared on infected and uninfected plants, but females fed on uninfected plants produced 71% more eggs than those fed on infected plants. Adult G. boliviana preferentially fed on uninfected plants when given a choice. There was no evidence of TSAMV transmission by G. boliviana. The potential impacts of TSAMV infection on the effectiveness of G. boliviana as a biological control agent are discussed.     

Effect of temperature on development, overwintering and establishment potential of Franklinothrips vespiformis in the UK

This study investigated the effect of temperature on the development and overwintering potential of the predatory thrips Franklinothrips vespiformis (Crawford) (Thysanoptera: Aeolothripidae), a biological control agent used against glasshouse pests in continental Europe and Israel. Developmental rates increased linearly with rearing temperatures. It was estimated that 304.9 degree days, above a lower threshold temperature of 11.9 °C, were required for F. vespiformis to complete development from egg to adult eclosion. The effect of low temperatures (–5, 0, and 5 °C) was examined on adult female and larval survival. Subsequent reproductive and developmental attributes of survivors were also investigated. Lethal time experiments indicated that larval stages are more cold tolerant than adult F. vespiformis females. Surviving larvae increased their developmental times to adults with decreasing temperature and increasing exposure periods and second instars were significantly more successful than first instars in reaching adulthood. Surviving adult females decreased their oviposition rate with decreasing temperature and increasing exposure periods, and exposures to low temperatures affected the number of viable eggs produced. The results are discussed in the context of overwintering and establishment potential of F. vespiformis in the UK in the event of introducing the predatory thrips as a biological control agent against glasshouse pests.     

Development of the willow sawfly,Nematus oligospilus,at different temperatures,and an estimation of voltinism throughout New Zealand

Abstract

Cohorts of willow sawfly (Nematus oligospilus) were reared in the laboratory on crack willow (Salix fragilis) at seven constant temperatures between 11.3 and 28.8°C. Larvae developed through 5–7 instars. Body size (length) of adult females was influenced by temperature and was strongly correlated with fecundity (R² = 0.8973). The developmental threshold (t_o), determined through linear regression, was calculated to be 8.1°C, and the thermal accumulation above t_o required to complete one generation was 321 degree‐days. The number of degree‐days available for seasonal development of the sawfly at 10 sites throughout New Zealand was calculated. Allowing for the period of diapause, it was estimated that N. oligospilus might potentially develop through up to seven generations per year in the North Island and three generations per year in the south of the South Island.     

Morphological and biological notes on Encarsia meritoria Gahan (Hymenoptera,Aphelinidae), a parasitoid of Trialeurodes vaporariorum (Westwood) (Homoptera,Aleyrodidae) new in Europe

Encarsia meritoria Gahan, a Neartic species recorded only in the USA, was found naturally occurring in Catalonia (north‐east Spain) in 1987. The morphology of immature stages, the rate of development in the range 12°C‐34°C, the longevity and fecundity at 24°C and some observations on its searching and host feeding behaviour are presented in this paper. Mean development times from egg laying to adult emergence ranged from 75 days at 12° C to 11 days at 28°C. The Lower Developmental Threshold computed from linear regression equations was 9°C. Females laid an average of 198 eggs (range 89–330) in an average of 34 days (longevity range 19–54 days). The intrinsic rate of increase at 24° C was 0.1717, slightly greater than the r_m of T. vaporariorum and smaller than the values reported for E. tricolor and E. formosa, which is not very promising for biological control. However, this may not be a definitive conclusion, because factors other than those considered in our experiments may play an important role infield conditions.     

Effect of temperature on reproduction,development, and phenotypic plasticity of Drosophila suzukii in Brazil

Temperature is a determining factor for the development and establishment potential of insect pests. The present study describes the impact of temperature (13, 18, 23, 25, 28, 30, and 33 °C) on the life cycle parameters and phenotypic plasticity of South American populations of Drosophila suzukii (Matsumura) (Diptera: Drosophilidae) in the laboratory. Secondary objectives were to determine the lower thermal threshold and thermal constant to estimate the number of annual generations of the insect in small-fruit-producing regions in Brazil. The highest egg-to-adult survival was recorded at 23 and 25 °C. At 30 and 33 °C, no emergence of D. suzukii was observed. The egg-to-adult development time was shortest at 25 and 28 °C (ca. 10 days). The net reproductive rate (R₀), and the intrinsic rate of population increase (r_m) were highest at 23 and 25 °C. In contrast, temperatures of 13 and 28 °C generated largest and smallest body sizes, respectively, and caused reductions of 99 and 93% in R₀. The estimated lower thermal threshold was 7.8 °C for egg-to-adult survival. The estimated thermal constant was 185.8 degree days, and the estimated annual number of generations of D. suzukii ranged from 17.1 in cold regions to 27.2 in warm regions. The results of the present study are important for understanding D. suzukii occurrence in the field, contributing to more informed and precise pest management.     

Multiple generation effects of high temperature on the development and fecundity of Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae) biotype B

Insects are ectotherms and their ability to resist temperature stress is limited. The immediate effects of sub‐lethal heat stress on insects are well documented, but longer‐term effects of such stresses are rarely reported. In this study, survival, development and reproduction of the whitefly, Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae) biotype B, were compared over five consecutive generations at 27, 31 and 35 °C and for one generation at 37 °C. Both temperature and generation significantly affected the fitness of the whitefly. These impacts were more dramatic with increasing generations and temperatures. Among the experimental temperatures, the most favorable for development and reproduction were 27 °C and 31 °C. At 27 °C, survival, development and fecundity were all stable over these five generations. At 31 °C, immature survival rate was the highest in the fifth generation, but female fecundities decreased in the fourth and fifth generations. At 35 °C, egg hatching rate, immature survival rate and female fecundity decreased significantly in the fourth and fifth generations. At 37 °C, survival of B. tabaci was not adversely affected, but female fecundity at 37 °C was less than 10% of that at 27 °C or 31 °C. These results demonstrate that the lethal high temperature for B. tabaci is over 37 °C, and the whitefly population continued expanding in the five generations at 35 °C. The ability of B. tabaci biotype B to survive high temperature stress will play an important role in its population extension under global warming.     

Thermal biology of the spider mite predator Phytoseiulus macropilis

Phytoseiulus macropilis Banks (Acari: Phytoseiidae) is a specialist predator of the two-spotted spider mite Tetranychus urticae Koch (Acari: Tetranychidae), and has been used to control the pest in its native range: Mediterranean regions, the tropics and Florida, USA. This study investigates the thermal biology as a proxy for establishment potential of glasshouse escapees in cooler northern European climates, using a combination of laboratory and field trials. High mortality rates at 10 °C indicated limited acclimation ability. Mites displayed continuous oviposition in conditions that have previously been shown to induce a hiatus in other phytoseiid species, supporting previous findings that Phytoseiulus is a genus with no diapause state. Adult P. macropilis supercooled to between ?17.2 and ?24.0 °C, but the lower lethal temperature₅₀ was ?5.7 °C, resulting in a high level of pre-freeze mortality. The lethal time₅₀ at 5 °C was 2.6 days, and maximum survival in winter field trials across 2010 and 2011 was 21 days. The thermal biology data indicate that P. macropilis is unlikely to establish in northern Europe, and will therefore make a suitable glasshouse biological control agent in temperate climates.