Daniel T. Dalton

Temperature-Related Development and Population Parameters for Drosophila suzukii (Diptera: Drosophilidae) on Cherry and Blueberry

ABSTRACT Temperature-related studies were conducted on Drosophila suzukii Matsumura (Diptera: Drosophilidae: Drosophilini). From 10–28°C, temperature had a significant impact on blueberries, Vaccinium corymbosum L. (Ericales: Ericaceae), and cherries, Prunus avium (L.) L. 1755 (Rosales: Rosaceae), important commercial hosts of D. suzukii. Temperature had a significant influence on D. suzukii developmental period, survival, and fecundity, with decreasing developmental periods as temperatures increased to 28°C. At 30°C, the highest temperature tested, development periods increased, indicating that above this temperature the developmental extremes for the species were approached. D. suzukii reared on blueberries had lower fecundity than reared on cherries at all temperatures where reproduction occurred. The highest net reproductive rate (Ro) and intrinsic rate of population increase (r m) were recorded on cherries at 22°C and was 195.1 and 0.22, respectively. Estimations using linear and nonlinear fit for the minimum, optimal, and maximum temperatures where development can take place were respectively, 7.2, 28.1, and 42.1°C. The rm values were minimal, optimal, and maximal at 13.4, 21.0, and 29.3°C, respectively. Our laboratory cultures of D. suzukii displayed high rates of infection for Wolbachia spp. (Rickettsiales: Rickettsiaceae), and this infection may have impacted fecundity found in this study. A temperature-dependent matrix population estimation model using fecundity and survival data were run to determine whether these data could predict D. suzukii pressure based on environmental conditions. The model was applied to compare the 2011 and 2012 crop seasons in an important cherry production region. Population estimates using the model explained different risk levels during the key cherry harvest period between these seasons.

Laboratory survival of Drosophila suzukii under simulated winter conditions of the Pacific Northwest and seasonal field trapping in five primary regions of small and stone fruit production in the United States.

BACKGROUND Drosophila suzukii was first found in Oregon in August 2009. The threat of this pest to regional small and stone fruit production industries led to investigations on its overwintering capabilities in fruit-growing regions in the Pacific Northwest. Knowledge of its cold tolerance will help in the development of computer models to forecast seasonal population growth and decline. RESULTS Of 1500 adults or pupae, 22 (1.4%) individuals survived the 84 day experimental chilling period. Most (86%) of the survivors were subjected to 10 °C temperature treatments. Survival decreased significantly at lower temperature treatments. Freezing temporarily increased the mortality rate but did not significantly affect overall mortality over the trial period. Flies that emerged from pupae are estimated to survive for up to 103-105 days at 10 °C and for shorter periods at lower temperatures. Field trapping in five fruit production areas has demonstrated overwintering survival in California and Oregon, but lower survival is predicted in Eastern Washington and Michigan. CONCLUSION The experiments reported here indicate that long-term survival of D. suzukii is unlikely at temperatures below 10 °C. Field data from five climatic regions indicated extended low initial D. suzukii field presence in 2010 in all regions except California, where field presence was recorded earlier.

Integrating temperature-dependent life table data into a matrix projection model for Drosophila suzukii population estimation

Temperature-dependent fecundity and survival data was integrated into a matrix population model to describe relative Drosophila suzukii Matsumura (Diptera: Drosophilidae) population increase and age structure based on environmental conditions. This novel modification of the classic Leslie matrix population model is presented as a way to examine how insect populations interact with the environment, and has application as a predictor of population density. For D. suzukii, we examined model implications for pest pressure on crops. As case studies, we examined model predictions in three small fruit production regions in the United States (US) and one in Italy. These production regions have distinctly different climates. In general, patterns of adult D. suzukii trap activity broadly mimicked seasonal population levels predicted by the model using only temperature data. Age structure of estimated populations suggest that trap and fruit infestation data are of limited value and are insufficient for model validation. Thus, we suggest alternative experiments for validation. The model is advantageous in that it provides stage-specific population estimation, which can potentially guide management strategies and provide unique opportunities to simulate stage-specific management effects such as insecticide applications or the effect of biological control on a specific life-stage. The two factors that drive initiation of the model are suitable temperatures (biofix) and availability of a suitable host medium (fruit). Although there are many factors affecting population dynamics of D. suzukii in the field, temperature-dependent survival and reproduction are believed to be the main drivers for D. suzukii populations.

Humidity affects populations of Drosophila suzukii (Diptera: Drosophilidae) in blueberry

Temperature and humidity affect insect physiology, survival, fecundity, reproductive status and behaviour. Complementing previous work investigating the effects of temperature on adult survival and fecundity of the invasive frugivorous pest, Drosophila suzukii (Matsumura), this study was conducted to determine the effect of humidity on D. suzukii larval development, adult survival, fecundity and reproductive status using blueberry as a host substrate. The five constant humidity levels in laboratory bioassays were 20, 33, 71, 82 and 94% RH at 20.6 ± 0.2°C. As RH increased, fecundity and longevity increased. At the higher humidity levels, RH had limited impact on mean generation times (T), larval development and eclosion times. The highest net reproductive rate (Ro = 68) and highest intrinsic rate of population increase (rm = 0.17) were both recorded at 94% RH. The reproductive status of females, as indicated by the number of mature oocytes per female, was significantly greater at 82 and 94% RH, compared to 71% RH. In addition to the laboratory procedures, we correlated field trap captures over an 81‐day summer period to relative humidity (RH) levels in close proximity to those traps. In the field, low ambient humidity levels resulted in decreased trap captures. A humidity‐dependent population model predicted lower densities of D. suzukii relative to populations at higher humidity. This study supports the hypothesis that cultural practices that minimize lower humidity levels in crops can contribute to the management of D. suzukii. Such methods may include open pruning, drip irrigation and field floor management.

Distribution and activity of Drosophila suzukii in cultivated raspberry and surrounding vegetation

Spotted wing drosophila, Drosophila suzukii (Matsumura) (Diptera: Drosophilidae), may utilize wild ‘Himalaya’ blackberry (HB) Rubus armeniacus Focke or other non‐crop plants as refugia and possibly exploit adjacent field margins before colonizing cultivated fruiting crops. Studies were conducted to determine the role of field margins containing HB and their effect on D. suzukii activity, density and distribution in an adjacent commercial red raspberry crop. One‐ha plots adjacent to field margins containing HB or known non‐host (NH) grass crops were established in 2011 and 2012 and replicated three times. Each plot contained two transects with monitoring traps for D. suzukii in the field margin (0 m) and spaced approximately 10 (crop boundary), 40, 70 and 100 m into the adjacent crop (n = 10 traps/plot). Field margin vegetation was treated with a 10% chicken egg white mark solution weekly from pre‐harvest until the end of harvest using a cannon sprayer. Adult D. suzukii were collected from traps weekly and analysed for the presence of the egg white mark using an egg white‐specific enzyme‐linked immunosorbent assay (ELISA). During both years, marked flies and total flies were captured in higher numbers in HB field margins, whereas virtually no flies were captured in field margins containing no known alternative host. Similarly, more flies were captured in the crop near HB than near NH. Spatial Analysis by Distance IndicEs (SADIE) and mean D. suzukii trap captures additionally displayed significantly higher fly densities in the raspberry field near HB than near NH. These results suggest that HB may contribute to elevated D. suzukii populations and pest pressure in comparison with field margins containing no known alternate host vegetation for D. suzukii. Having closely adjacent non‐crop alternate host landscapes may result in increased D. suzukii pest pressure.

Drosophila suzukii (Diptera: Drosophilidae) and its potential impact to wine grapes during harvest in two cool climate wine grape production regions

ABSTRACT Drosophila suzukii (Matsumura) is a global pest attacking various berry crops. D. suzukii lays eggs in damaged and in intact wine grape berries of the most soft-skinned varieties. Here, we describe the relative host utilization of different wine grape cultivars grown in Northern Italy and Oregon. Assessments of host berry utilization were performed in both field and laboratory settings. Results were correlated to physiological changes occurring during grape berry development starting at véraison and concluding during harvest. We found that oviposition increased with an increase in sugar content and a decrease of acidity levels. Oviposition increased with a decrease of penetration force. Penetration force, as a measure of skin hardness, is a critical component of host selection among the D. suzukii-exposed cultivars. We demonstrated that incised berries are more favorable for D. suzukii oviposition and as a nutrient substrate. Increased presence on wine grapes, as indicated by egg laying and increased longevity, was observed for flies that were exposed to incised berries as opposed to fully intact berries. D. suzukii flies can be found feeding on damaged wine grapes during the harvest period, especially when the skins of berries are negatively impacted due to cracking, disease, hail injury, and bird damage. Such an increase of feeding and oviposition may increase the likelihood of spoilage bacteria vectoring due to D. suzukii.

Trap Designs for Monitoring Drosophila suzukii (Diptera: Drosophilidae)

ABSTRACT Drosophila suzukii (Matsumura), an invasive pest of small and stone fruits, has been recently detected in 39 states of the United States, Canada, Mexico, and Europe. This pest attacks ripening fruit, causing economic losses including increased management costs and crop rejection. Ongoing research aims to improve the efficacy of monitoring traps. Studies were conducted to evaluate howphysical trap features affect captures of D. suzukii.We evaluated five colors, two bait surface areas, and a top and side position for the fly entry point. Studies were conducted at 16 sites spanning seven states and provinces of North America and nine crop types. Apple cider vinegar was the standard bait in all trap types. In the overall analysis, yellow-colored traps caught significantly more flies than clear, white, and black traps; and red traps caught more than clear traps. Results by color may be influenced by crop type. Overall, the trap with a greater bait surface area caught slightly more D. suzukii than the trap with smaller area (90 vs. 40cm2). Overall, the two traps with a side-mesh entry, with or without a protective rain tent, caught more D. suzukii than the trap with a top-mesh entry and tent.

Characterization and manipulation of fruit susceptibility to Drosophila suzukii

Drosophila suzukii (Matsumura) is an economic pest of small fruits and cherries that attacks intact ripening fruits. Host susceptibility may be influenced by characteristics such as flesh firmness, penetration force of the skin, total soluble solids (TSS, also known as °Brix), and pH. Improved knowledge of factors affecting fruit susceptibility is needed for developing thresholds and risk prediction models for IPM. A combination of laboratory and field studies was conducted to develop prediction and potential management tools. First, a direct bioassay was used to calculate the probability of oviposition in a given fruit based on various characteristics as determined across laboratory and field trials in Oregon and North Carolina, US. When multiple characteristics were evaluated simultaneously, oviposition probability consistently increased as penetration force decreased and pH increased. Oviposition probability sometimes increased as TSS increased. Second, raspberries and blueberries in unsprayed fields had substantially lower infestation in ripening fruit compared to ripe fruit. There was no or minimal infestation in green fruit. Third, given that skin penetration force influences oviposition, practices used to improve fruit quality were examined in laboratory no-choice cages for potential reduction of oviposition. Blueberry fruit sprayed with calcium silicate in the field had greater penetration force and firmness and reduced number of eggs laid by D. suzukii compared to untreated fruit. Other calcium-based treatments increased Ca content and firmness of fruit relative to untreated fruit. Timing of insecticide spray for D. suzukii might be delayed until fruit become susceptible.

Analysis of the invasiveness of spotted wing Drosophila (Drosophila suzukii) in North America, Europe, and the Mediterranean Basin

The polyphagous Asian vinegar fly Drosophila suzukii (spotted wing Drosophila) is a native of Eastern and Southeastern Asia. It emerged as an important invasive insect pest of berries and stone fruits in the Americas and Europe beginning in 2008. Species distribution models are commonly used for analyzing the extant and potential range expansion of invasive species. Previous modeling efforts for D. suzukii include a degree-day model, a MaxEnt ecological niche model, a demographic model incorporating the effects of temperature, and a preliminary mechanistic physiologically-based demographic model (PBDM). In the present analysis, we refine the PBDM for D. suzukii based on biological data reported in the literature. The PBDM is used to assess the effects of temperature and relative humidity from a recently published global climate dataset (AgMERRA) on the prospective geographic distribution and relative abundance of the pest in the USA and Mexico, and in Europe and the Mediterranean Basin. Our focus is on areas of recent invasion and of predicted higher invasiveness in these areas. Although the species is native to Asia and is of putative temperate origins, it has established in subtropical to north temperate zones worldwide where it infests a wide range of wild and domesticated berries and stone fruits. The model captures the observed phenology of D. suzukii at specific locations, as well as the potential geographic distribution and relative favorability across larger regions. The main limiting factor is cold winter temperature in northern areas, though high temperatures and low relative humidity may be limiting in arid areas. The effect of greater cold tolerance in winter morph adults is explored.

Seasonal Reproductive Biology of Drosophila suzukii (Diptera: Drosophilidae) in Temperate Climates

Abstract Drosophila suzukii (Matsumura; Diptera: Drosophilidae) is a key pest of sweet cherry and small fruits worldwide. The present studies were designed to describe the reproductive physiology in both sexes, through dissections of their reproductive organs. We extensively dissected female D. suzukii throughout the season from 2013 to 2016 and classified the reproductive status flies based on five recognizable ovarian maturation stages: 1) no ovaries; 2) unripe ovaries 3) ripening eggs in ovarioles; 4) mature eggs in ovarioles; and 5) mature eggs in the abdomen. Development was examined as a function of calendar days as well as degree-days (DD). Results obtained from winter collections revealed that females collected from November to March contained a lower percentage of mature eggs than females collected from April to September. These data suggest that environmental conditions during the dormant period induce reproductive diapause. Oogenesis likely increased with an increase in mean monthly temperatures and DD. The first overwintered females with mature eggs were dissected as early as 21 February 2014 in Trento (7 DD). Additionally, we found that a low proportion of males (less than 50%) had sperm in their testes between January and March, yet during the same period females already have sperm stored in their spermathecal. Ivy berries was an alternative but unfavorable non-crop host during the late dormant period, as evidenced by emergence of smaller adults when compared to individuals emerging from cherry fruits. This study showed that D. suzukii females have great potential for oviposition early in the season, posing a risk to early season maturing crop hosts.