Sampling techniques for thrips (Thysanoptera: Thripidae) in preflowering tomato

Sampling techniques for thrips (Thysanoptera: Thripidae) were compared in preflowering tomato plants at the Coastal Plain Experiment Station in Tifton, GA, in 2000 and 2003, to determine the most effective method of determining abundance of thrips on tomato foliage early in the growing season. Three relative sampling techniques, including a standard insect aspirator, a 946-ml beat cup, and an insect vacuum device, were compared for accuracy to an absolute method and to themselves for precision and efficiency of sampling thrips. Thrips counts of all relative sampling methods were highly correlated (R > 0.92) to the absolute method. The aspirator method was the most accurate compared with the absolute sample according to regression analysis in 2000. In 2003, all sampling methods were considered accurate according to Dunnett's test, but thrips numbers were lower and sample variation was greater than in 2000. In 2000, the beat cup method had the lowest relative variation (RV) or best precision, at 1 and 8 d after transplant (DAT). Only the beat cup method had RV values <25 for all sampling dates. In 2003, the beat cup method had the lowest RV value at 15 and 21 DAT. The beat cup method also was the most efficient method for all sample dates in both years. Frankliniella fusca (Pergande) was the most abundant thrips species on the foliage of preflowering tomato in both years of study at this location. Overall, the best thrips sampling technique tested was the beat cup method in terms of precision and sampling efficiency.     

Spatial dependence, dispersion, and sequential sampling of Anaphothrips obscurus (Thysanoptera: Thripidae) in timothy

The spatial distribution and dispersion of Anaphothrips obscurus (Müller) (Thysanoptera: Thripidae) was examined with the goal of establishing a sequential sampling plan for this pest in timothy, Phleum pratense L. (Poaceae). Approximately 16 different California timothy fields were sampled twice yearly from 2006 to 2008 using direct observation and the beat cup method. For direct observation, the number of thrips on each leaf of the plant was counted. For the beat cup method, tillers were tapped into a cup and dislodged thrips were counted. Samples were separated by ≈3 m in 2006 and 2007 and exactly 3 m in 2008. Spatial autocorrelation of intrafield population distribution was tested for significance in 2008 using Moran's I, but autocorrelation was not detected. The population dispersion was assessed by Taylor's power law and was determined to be aggregated and density-dependent. Intraplant population dispersion and distribution for each year were also evaluated for adults, larvae, and total thrips. All lifestages were highly spatially dependent and more thrips were found near the top of the plant than the bottom. Direct observation proved to be a more accurate and precise method than the beat cup method, especially when thrips abundances were greater than one. However, the number of samples required to provide an accurate level of precision was unrealistic for both methods. A sequential sampling plan was evaluated, but was not practical for the beat cup method because few thrips were found using this method. Because there was no spatial autocorrelation at sampling distances of 3 m, samples can be taken at intervals at 3 m to obtain spatially independent population abundance estimates.     

Quantifying seasonal thrips population dynamics in relation to temperature and wheat senescence

The incidence of thrips in the High Plains of Texas (USA) was investigated using sticky traps during the 2021 and 2022 seasons. Yellow sticky traps were placed in wheat fields and collected and replaced weekly and thrips were counted under a dissecting scope. Weekly wheat reflectance measurements were taken using a hyperspectral radiometer from which normalized difference vegetation index (NDVI) was calculated for each measurement. Temperature (degree day) and NDVI values were then related to weekly thrips incidence using regression. Thrips incidence curvilinearly increased over time during each of the two seasons and reached a maximum in the middle of June, after which it declined sharply. There was a strong positive relationship between degree days and thrips incidence until the incidence reached a maximum, whereas the incidence was negatively related to NDVI values. Analysis of the thrips changes over time progress with the two variables together showed that degree day has greater impact on thrips incidence than NDVI. However, the steep decline in thrips abundance after its peak in mid-June suggests that senesced wheat fields with NDVI values near zero are not significant sources of thrips, signifying the importance of wheat growth stages in the seasonal population dynamics of thrips. Overall, the 2-year results were generally consistent in trends of thrips incidence during the season, which may need to be considered when choosing vegetable planting dates in the region.     

Spatiotemporal population dynamics of the banana rind thrips,Elixothrips brevisetis (Bagnall) (Thysanoptera: Thripidae)

The understanding of how environmental factors and agricultural practices affect population dynamics of insect pests is necessary for pest management. Here, we provide insight into the ecology of the banana rind thrips Elixothrips brevisetis (Bagnall) (Thysanoptera: Thripidae) by collecting and analysing a spatiotemporal database of population estimates in Martinique (West French Indies). We assessed the influence of climatic variables (which were rainfall and temperature) and biotic variables (which were banana and three weed species) on the adult thrips abundance for different components of the banana plant (sucker, mother plant and bunch) and evaluated the effect of thrips abundance and standard bunch covers on damages. The abundance of thrips on the sucker, the mother plant, and the bunch was significantly related to the abundance on neighbouring banana plants, and spatial autocorrelation indicated that E. brevisetis dispersed for only short distances. The number of thrips on the mother plant and on the bunch was positively related to the number of thrips on the sucker, suggesting that the thrips may disperse from the sucker to the mother plant and then to the bunch. The abundance of thrips on the sucker increased with sucker height and was positively correlated with the mean daily rainfall during the 17 days before sampling; the length of that period might correspond with the time required for an individual to complete its life cycle. Covered bunches had 98% fewer thrips than non‐covered bunches, and the damage caused by thrips was linearly related to the number of thrips present between the 2nd and 4th week after flowering. Finally, we found that the presence of Alocasia cucullata, Dieffenbachia seguine and Peperomia pellucida is significantly related with a decrease in thrips abundance on banana plants, suggesting the use of these weeds as potential trap plants.     

Spatial and temporal pattern of colonization of Nabidae (Heteroptera) in alfalfa (Medicago sativa)

Abstract: Six species of Nabidae (Heteroptera) were collected by standardized sweep net sampling in alfalfa fields in Thuringia, Germany, from 1993 to 1995: Nabis pseudoferus , N. ferus , N. brevis , N. major , Nabicula flavomarginata and Aptus mirmicoides . Colonization of a newly cultivated field was studied over a 3-year period. The density of all the studied nabid species was low (less than five individuals per 100 sweeps) and not related to time since colonization started, or to the distance from the margin of the field. Macropterous species were able to colonize the whole field within one season. The density of one macropterous species, N. pseudoferus , varied between the years of study and was mainly affected by the harvest regime. The brachypterous species reached the margin within one season but for density it took three seasons to reach satiated values also in the centre of the field. The abundance of the brachypterous N. brevis was significantly different both between years and sampling sites. This indicates the importance of the surroundings on the succession of this species. Nabis major , a fully winged species, showed a migration pattern intermediate to macropterous and brachypterous nabids. These results suggest that the total abundance of nabid predators cannot be predicted by time or distance from the expansion source (shelter belts). The abundance of brachypterous nabid individuals can be predicted from time since colonization but is best analysed at the species level.     

西花蓟马在黄瓜和架豆上的空间分布型及理论抽样数

西花蓟马Frankliniella occidentalis（Pergande）已在北京部分蔬菜园区成功定居,对蔬菜生产造成危害。为了解该虫在蔬菜田的种群空间分布型,指导田间取样,本文应用几种聚集度指标的计算公式以及Taylor、Iwao的回归方程式,分析和测定了西花蓟马的种群空间分布格局。结果表明：在黄瓜和架豆上西花蓟马的空间分布型一致,均为聚集分布型;该虫的空间分布型不受种群密度的影响,并且也不随取样时间的变化而变化。种群数量动态研究显示西花蓟马成虫和若虫在黄瓜植株的中部叶片分布较多（从顶部向下数第4叶至第17叶片上成、若虫的分布数量多于其他叶片上的数量）,未展开的叶片、嫩叶和下部老叶危害较轻。用Iwao回归法中α、β参数计算出在允许误差范围内的理论抽样数。     

Evaluation of sampling methodology for determining the population dynamics of onion thrips (Thysanoptera: Thripidae) in Ontario onion fields

Onion thrips, Thrips tabaci Lindeman, are an economic pest of alliums worldwide. In Ontario onion-growing regions, seasonal abundance and population trends of onion thrips are not well known. The objectives of this research were to investigate onion thrips population dynamics by using both white sticky traps and plant counts, to gain insight into flight height, and to determine the genus and sex of thrips fauna present in monitored fields. Adult thrips were captured on white sticky traps placed in two commercial onion fields in the Thedford-Grand Bend Marsh region as early as mid-May in 2001, 2002, and 2003. Thrips were not recorded on onion plants in these fields until late June and early July. A comparison of sticky trap captures to plant counts revealed a strong, positive correlation, indicating that sticky traps, which consistently detected thrips earlier than plant counts, could be used instead of plant counts early in the season to monitor onion thrips populations. Pole traps placed in onion and an adjacent soybean, Glycine max (L.) Merr., field revealed that regardless of crop type, most thrips were captured 0.7-0.95 m above the soil surface. During this study, 70% of 137,000 thrips captured on sticky traps and 89% of 1,482 thrips captured in pan traps were female onion thrips. No male onion thrips were identified in this study: most of the remaining thrips were Frankliniella spp.     

Weather-induced changes in moth activity bias measurement of long-term population dynamics from light trap samples

Interpretation of light trap catches of moths is complicated by daily variation in weather that alters flight activity and numbers caught. Light trap efficiency is also modified by wind and fog, and daily weather may effect absolute abundance (numbers actually present). However, actograph experiments and other sampling methods suggest that changes in daily activity are large by comparison to changes in absolute abundance. Daily variation in weather (other than wind and fog) is therefore a form of sampling error in absolute abundance estimates. We investigated the extent of this sampling bias in 26 years of population dynamics from 133 moth species. In a subset of 20 noctuid and geometrid species, daily numbers caught were positively correlated with temperature in 14 species, and negatively correlated with rainfall in 11 species. The strength of correlations varied between species, making it difficult to standardize catches to constant conditions. We overcame this by establishing how weather variation changed with time and duration of the flight period. Species flying later in the summer and for shorter periods experienced more variable temperatures, making sampling error greater for these species. Of the 133 moth species, those with shorter flight periods had greater population variability and more showed significant temporal density dependence. However, these effects were weak, which is encouraging because it suggests that population analyses of light trap data largely reflect factors other than sampling error.     

Assessing abundance and distribution of an invasive thrips Frankliniella schultzei (Thysanoptera: Thripidae) in south Florida

Within-plant and within-field distribution of larvae and adults of an invasive thrips species, Frankliniella schultzei (Trybom) on cucumber, Cucumis sativus L. was studied in 2008 and 2009 in Homestead, Florida. The majority of thrips were found inhabiting flowers of cucumber plants and little or none was found on the other parts of the plant. Thrips were aggregated in the field, as indicated by the two regression models, Taylor's power and Iwao's patchiness regression. Iwao's patchiness regression provided a better fit than Taylor's power law. The distribution was clumped during the initial stages of infestation at the edges of the field and became random thereafter. However, with increase in population density, thrips again formed aggregates in the field. Based on the average pest density per flower in a ～0.25-ha field, minimum sample size (number of flowers) required at the recommended precision level (0.25) was 51. The number of samples required at two levels of predetermined pest density was also calculated, which would help growers in collecting optimum number of samples required to determine the correct threshold level of pest in fields. Results from seasonal abundance indicated that density of thrips peaked during the fifth week of sampling with an average of 25 and 34 adults per ten flowers during autumn 2008 and 2009, respectively. Results from these studies will help growers and extension personnel in understanding the abundance and distribution of F. schultzei in the field, which are important components required in developing a sound management program.     

Temporal variation in arthropod sampling effectiveness: the case for using the beat sheet method in cotton

Predatory insects and spiders are key elements of integrated pest management (IPM) programmes in agricultural crops such as cotton. Management decisions in IPM programmes should to be based on a reliable and efficient method for counting both predators and pests. Knowledge of the temporal constraints that influence sampling is required because arthropod abundance estimates are likely to vary over a growing season and within a day. Few studies have adequately quantified this effect using the beat sheet, a potentially important sampling method. We compared the commonly used methods of suction and visual sampling to the beat sheet, with reference to an absolute cage clamp method for determining the abundance of various arthropod taxa over 5 weeks. There were significantly more entomophagous arthropods recorded using the beat sheet and cage clamp methods than by using suction or visual sampling, and these differences were more pronounced as the plants grew. In a second trial, relative estimates of entomophagous and phytophagous arthropod abundance were made using beat sheet samples collected over a day. Beat sheet estimates of the abundance of only eight of the 43 taxa examined were found to vary significantly over a day. Beat sheet sampling is recommended in further studies of arthropod abundance in cotton, but researchers and pest management advisors should bear in mind the time of season and time of day effects.     

Host plant age and population development of a cereal aphid, Metopolophium dirhodum (Hemiptera: Aphididae)

The sudden decline following the peak in population abundance of aphids on crops of small grain cereals is attributed to the joint effect of natural enemies and plant senescence. To distinguish between these causes, a four year experiment was established in which the numbers of Metopolophium dirhodum (Walker) infesting spring wheat plots sown from April to June at c. 14 day intervals were determined. Aphid abundance in replicates sown at successive dates peaked within a period of 5-9 days (106-171 day degrees above a base temperature of 0 degrees C) although their sowing dates varied by 62-97 days (727-1106 day degrees). At the time of the aphid population peaks, plants in the different sowings differed in age (11-99 days), developmental stage (stage 15-65 on the Zadoks scale), leaf nitrogen content and shoot mass. Maximum abundance of M. dirhodum decreased with sowing date because the time available for its population increase was shorter on late than early sowings. The abundance of M. dirhodum on spring wheat was similar to its abundance on winter wheat. After reaching peak abundance, aphids declined in numbers within 3-7 days. The effect of host plant ageing on the M. dirhodumdecline thus appeared small. Natural enemies (largely mycoses), and timing of alata production may have contributed to the aphid decline.     

Population dynamics and within-plant distribution of the mite Calacarus flagelliseta (Acari: Eriophyidae) on papaya in Hawaii

An important element in developing a management strategy for a new pest is the study of its seasonal dynamics and within-plant distribution. Here, we studied the mite Calacarus flagelliseta Fletchmann, De Moraes & Barbosa on papaya, Papaya carica L. (Caricaceae), in Hawaii to quantify 1) patterns of seasonal abundance, 2) its distribution across different vertical strata of the papaya canopy, and 3) shifts in its use of the upper versus the lower surfaces of papaya leaves. Nondestructive sampling conducted in two papaya plantings revealed that 1) populations of C. flagelliseta peak during the summer; 2) mites are most abundant in the middle and lower strata of the plant canopy, and least abundant on the youngest leaves found in the upper canopy; and 3) mites are found more predominantly on the upper leaf surfaces when overall population density peaks, suggesting that individuals move from the lower to the upper leaf surfaces when food resources on the lower leaf surface have been exploited by conspecifics. These results have significant implications for the development of sampling plans for C. flagelliseta in papaya.     

Sampling methods, dispersion patterns, and fixed precision sequential sampling plans for western flower thrips (Thysanoptera: Thripidae) and cotton fleahoppers (Hemiptera: Miridae) in cotton

A 2-yr field study was conducted to examine the effectiveness of two sampling methods (visual and plant washing techniques) for western flower thrips, Frankliniella occidentalis (Pergande), and five sampling methods (visual, beat bucket, drop cloth, sweep net, and vacuum) for cotton fleahopper, Pseudatomoscelis seriatus (Reuter), in Texas cotton, Gossypium hirsutum (L.), and to develop sequential sampling plans for each pest. The plant washing technique gave similar results to the visual method in detecting adult thrips, but the washing technique detected significantly higher number of thrips larvae compared with the visual sampling. Visual sampling detected the highest number of fleahoppers followed by beat bucket, drop cloth, vacuum, and sweep net sampling, with no significant difference in catch efficiency between vacuum and sweep net methods. However, based on fixed precision cost reliability, the sweep net sampling was the most cost-effective method followed by vacuum, beat bucket, drop cloth, and visual sampling. Taylor's Power Law analysis revealed that the field dispersion patterns of both thrips and fleahoppers were aggregated throughout the crop growing season. For thrips management decision based on visual sampling (0.25 precision), 15 plants were estimated to be the minimum sample size when the estimated population density was one thrips per plant, whereas the minimum sample size was nine plants when thrips density approached 10 thrips per plant. The minimum visual sample size for cotton fleahoppers was 16 plants when the density was one fleahopper per plant, but the sample size decreased rapidly with an increase in fleahopper density, requiring only four plants to be sampled when the density was 10 fleahoppers per plant. Sequential sampling plans were developed and validated with independent data for both thrips and cotton fleahoppers.     

Dynamics ofSitotroga cerealella (olivier) (lepidoptera: gelechiidae) andSitophilus zeamais motschulsky (Coleoptera: Curculionidae) populations in a small bulk of stored corn

The dynamics of Angoumois grain moth, Sitotroga cerealella (Olivier ), and maize weevil, Sitophilus zeamaisMotschulsky , populations breeding in a small bulk (initially 5.36 t) of shelled corn were studied over an 8-year period by monthly sampling. The weevil population showed wide fluctuations in density superimposed on a general decline with time. The moth population showed no upward or downward trend for the first 60 months, although it fluctuated widely. Following a decline that occurred between 56 and 60 months, the moth population fluctuated within a much narrower range, and there was a general decrease in density with time. The decline of the weevil population paralleled deterioration of the corn as did that of the moth population after ca 60 months, and the decline of both species probably resulted from increasing scarcity of suitable breeding sites. Both populations exhibited seasonal variation in density with minima in late summer and early fall, following periods of adversely high temperatures in the storage shed. The populations increased during the fall, leveled off or declined slightly during the winter months, and then increased to maximum levels in late spring or early summer. It thus appears that high temperatures had a greater adverse effect on the populations than low temperatures. The grain moth and the maize weevil both tended to be randomly dispersed at low population levels and moderately aggregated at intermediate and high levels, although the degree of aggregation was not correlated with population density when low population levels were considered separately, and the maize weevil showed a greater tendency for aggregation than did the grain moth. Analysis of individual samples at fixed points in time showed a conspicuous bias for negative correlation between numbers of the two species within sampling quadrats, suggesting a tendency for the two species to segregate within the grain mass. This process could have resulted from behavioral differences or from the destruction of one species by the other. Competitive displacement of the grain moth by the maize weevil has been demonstrated in laboratory experiments but has rarely been observed under natural conditions, and in our study the two species coexisted for 8 years in a relatively small grain bulk.     

Assessment of methods for estimating wild rabbit population abundance in agricultural landscapes

Various methods have been used to estimate rabbit abundance, but comparisons of standard methods are still lacking, and thus, results remain roughly comparable across studies. Ideally, a method should be applicable over a wide range of situations, such as differing abundances or habitat types. Comparisons of methods are required to evaluate the benefits of each of them, and survey methods should be validated for the conditions in which they will be used. In this study, we compare the performance of direct methods (kilometric abundance index and distance sampling) in two seasons and at two times of day (dusk and night) for estimating wild rabbit abundances in agricultural landscapes. Estimates based on direct methods were highly correlated and detected similar seasonal population changes. Night counts provided better estimates than did dusk counts and exhibited more precision. Results are discussed within the context of rabbit behaviour and their implications for rabbit population surveys.     

Impact of neonicotinoid seed treatments on thrips (Thysanoptera: Thripidae) and soybean yield in Virginia and North Carolina

Currently there are several neonicotinoid insecticide seed treatments registered for use on soybean (Glycine max L.), with disparity in adoption rates in the eastern United States. A complex of seedling insect pests is found in mid-south soybean, but thrips are the primary early season pest of soybean in Virginia and North Carolina. Published knowledge regarding their impact on soybean yield is minimal, as is the impact of thrips on soybean yield; thrips species composition is also understudied. In 2008 through 2010, nine field experiments in Virginia and North Carolina were conducted to evaluate the impact on thrips population dynamics; the influence on yield of neonicotinoid seed treatments, imidacloprid and thiamethoxam, was reported from nine of these experiments. Moreover, thrips species abundance was recorded in three of these experiments. Both imidacloprid and thiamethoxam reduced thrips densities compared with untreated soybean. Thiamethoxam was more effective than imidacloprid in reducing adult thrips densities at 5 wk after planting. Adult densities peaked at 3 wk after planting, followed by larval densities, which peaked at 4 wk after planting. The most abundant thrips species was Frankliniella fusca (Hinds), followed by Neohydatothrips variabilis (Beach). Other common species included F. occidentalis (Pergande) and F. tritici (Fitch). In general, F. fusca was more common earlier in the season, while N. variabilis was more common later in the season. There were no significant differences in yield among any of the treatments or in the untreated controls. Although neonicotinoid insecticides reduced thrips abundance, data collected in these studies demonstrated that there was no positive yield response.     

Seasonal and habitat-related distribution pattern of Synechococcus genotypes in Lake Constance

The abundance and distribution of Synechococcus spp. in the autotrophic picoplankton of Lake Constance, were followed in the pelagic and littoral habitat by qPCR over 2 years. One genotype, represented by isolated phycoerythrin-rich strain BO 8807, showed a seasonal distribution pattern in both habitats. Before a stable thermal stratification, the maximum of both the Synechococcus population and genotype BO 8807 occurred at 15 or 20 m water depth in the pelagic habitat. During the summer stratification, when the absolute abundance of all Synechococcus spp. was highest above 15 m, the absolute and relative abundance of genotype BO 8807 was maximal at 20 m. These results indicate that Synechococcus spp. or single genotypes are present in deep maxima in Lake Constance. The in situ dynamics of genotype BO 8807 is consistent with the observation that isolated strain BO 8807 requires higher phosphate concentrations for maximum growth rates than a strain from the same phylogenetic cluster that dominates the pelagic summer population. In contrast to these findings, low genome numbers of phycocyanin-rich genotype BO 8805 were found temporarily only in both the littoral and pelagic plankton. Microscopy revealed that PC-rich cells in general occurred preferentially in the littoral habitat. We discuss our results with respect to the versatility of picocyanobacteria of the evolutionary lineage VI of cyanobacteria, and a habitat-related distribution pattern of Synechococcus genotypes.     

Six years of wild bee monitoring shows changes in biodiversity within and across years and declines in abundance

Wild bees form diverse communities that pollinate plants in both native and agricultural ecosystems making them both ecologically and economically important. The growing evidence of bee declines has sparked increased interest in monitoring bee community and population dynamics using standardized methods. Here, we studied the dynamics of bee biodiversity within and across years by monitoring wild bees adjacent to four apple orchard locations in Southern Pennsylvania, USA. We collected bees using passive Blue Vane traps continuously from April to October for 6 years (2014–2019) amassing over 26,000 bees representing 144 species. We quantified total abundance, richness, diversity, composition, and phylogenetic structure. There were large seasonal changes in all measures of biodiversity with month explaining an average of 72% of the variation in our models. Changes over time were less dramatic with years explaining an average of 44% of the variation in biodiversity metrics. We found declines in all measures of biodiversity especially in the last 3 years, though additional years of sampling are needed to say if changes over time are part of a larger trend. Analyses of population dynamics over time for the 40 most abundant species indicate that about one third of species showed at least some evidence for declines in abundance. Bee family explained variation in species‐level seasonal patterns but we found no consistent family‐level patterns in declines, though bumble bees and sweat bees were groups that declined the most. Overall, our results show that season‐wide standardized sampling across multiple years can reveal nuanced patterns in bee biodiversity, phenological patterns of bees, and population trends over time of many co‐occurring species. These datasets could be used to quantify the relative effects that different aspects of environmental change have on bee communities and to help identify species of conservation concern.     

Relationship between Euschistus conspersus (Hem., Pentatomidae) pheromone trap catch and canopy samples in processing tomatoes

Abstract:  This study identified statistically significant, and practically important, seasonal pheromone trap response dynamics of Euschistus conspersus (Uhler) in processing tomatoes in 12 fields over 5 years in California, USA. Commercial fields were monitored weekly using traps baited with Euschistus spp. pheromone and direct canopy shake samples. Regression analysis determined no significant relationship between the two sampling methods at any given time during the growing season. This result indicates that the number of E. conspersus recovered in pheromone traps cannot be expected to serve as a direct estimate of the surrounding field population density from week to week during the processing tomato growing season. However, E. conspersus response to pheromone traps was characterized by differential attraction across the sample dates and elevated response early in the growing season relative to the associated field population. Among the most important findings of this work are differential responses of female and male E. conspersus to pheromone traps by crop phenology stage, and the fact that female-biased trap catch coincided with elevated trap response relative to the surrounding field population early in the growing season during tomato vegetative growth through flowering phenology stages. These results, when coupled with further E. conspersus reproductive biology studies, suggest that growers and crop consultants will be able to rely on E. conspersus pheromone trap catch patterns identified in this study to implement pheromone trap monitoring and phenology-based timing of insecticide treatment targeting E. conspersus nymphal stages in processing tomatoes.