Susceptibility of Sitophilus zeamais (Motsch.) (Coleoptera: Curculionidae) and Prostephanus truncatus (Horn) (Coleoptera: Bostrichidae) to Entomopathogenic Fungi from Ethiopia

The efficacy of 13 isolates of entomopathogenic fungi belonging to Beauveria , Metarhizium or Paecilomyces spp. was assessed against Sitophilus zeamais (Coleoptera: Curculionidae) and Prostephanus truncatus (Coleoptera: Bostrichidae) using a total immersion bioassay technique in the laboratory. Fungi were applied at concentrations of 1 ×10 7 and 1 ×10 8 conidia mL -1 for P. truncatus and S. zeamais , respectively. All isolates tested were virulent to P. truncatus (98-100% mortality, and median survival time (MST) ranged from 2.85-4.05 days). Metarhizium anisopliae and B. bassiana were also virulent to S. zeamais (92-100% mortality, MST ranged from 3.58-6.28 days). The isolate of Paecilomyces sp. was found to be the least virulent against S. zeamais , causing only 26.32 ±4.29% mortality with MST of 10.38 ±0.29 days. P. truncatus proved more susceptible to the entomopathogenic fungi tested than S. zeamais . One M. anisopliae (PPRC-EE) and three B. bassiana isolates (PPRC-HH, PPRC-9609 and PPRC-9614) were selected for further study and dose-mortality relationships were assessed on S. zeamais . The tested concentrations ranged from 1 ×10 4 -1 ×10 7 conidia mL -1 . M. anisopliae (PPRC-EE) showed the lowest LC 50 (3.39 ×10 5 conidia mL -1 ) followed by B. bassiana PPRC-HH (2.04 ×10 6 conidia mL -1 ). PPRC-9609 and PPRC-9614 showed slight differences in LC 50 but not at LC 90 . The results revealed the higher potency of M. anisopliae as compared with the B. bassiana isolates tested. The study suggests that the use of entomopathogenic fungi may hold promise as an alternative method to control pests of stored-products in Ethiopia.     

Grain injury models for Prostephanus truncatus (Coleoptera: Bostrichidae) and Sitophilus zeamais (Coleoptera: Curculionidae) in rural maize stores in West Africa

Prostephanus truncatus (Horn) and Sitophilus zeamais Motschulsky have been reported as the two most serious pests of stored maize in sub-Saharan Africa and smallholder farmers are in urgent need of guidelines for their proper management. In this article we investigate the injury rates attributable to these two species in terms of percentage weight loss and percentage grain damage, and we derive functional response models for the two species on maize. The models successfully described the progression of grain injury in an extensive data set compiled from previously published studies, comprising 46 time series of data relating maize injury and insect pest density. The grain injury models can be used in conjunction with predictive models of pest population dynamics to guide the development of integrated management strategies for postharvest maize pests in West Africa and comparable regions elsewhere.     

A novel 8.7 kDa protease inhibitor from chan seeds (Hyptis suaveolens L.) inhibits proteases from the larger grain borer Prostephanus truncatus (Coleoptera: Bostrichidae)

A novel trypsin inhibitor purified from chan seeds (Hyptis suaveolens, Lamiaceae) was purified and characterized. Its apparent molecular mass was 8700 Da with an isoelectric point of 3.4. Its N-terminal sequence showed a high content of acidic amino acids (seven out of 18 residues). Its inhibitory activity was potent toward all trypsin-like proteases extracted from the gut of the insect Prostephanus truncatus (Coleoptera: Bostrichidae), a very important pest of maize. This activity was highly specific, because among proteases from seven different insects, only those from P. truncatus and Manduca sexta (Lepidoptera: Sphingidae) were inhibited. This inhibitor has potential to enhance the defense mechanism of maize against the attack of P. truncatus.     

Biological control of the larger grain borer Prostephanus truncatus (Coleoptera: Bostrichidae) in Kenya using a predatory beetle Teretrius nigrescens(Coleoptera: Histeridae)

The establishment of the predatory histerid beetle, Teretrius nigrescens Lewis, following its field release in south eastern Kenya in 1992, and its impact on populations of the larger grain borer, Prostephanus truncatus (Horn), in woodland habitats is evaluated. Pheromone trap catch data show a single strong peak of P. truncatus from November to January, coincident with the short rains. Comparisons of P. truncatus pheromone trap catches before and after predator establishment, and in areas with and without T. nigrescens, show a sustained decline of over 80% in P. truncatus abundance over a 5-year period, following the establishment T. nigrescens. Concurrent laboratory studies of the impact of T. nigrescens on populations of P. truncatus raised on a wood substrate showed a 77% reduction in the growth of the pest population. These results are compared with published findings of the impact of T. nigrescens as a biological control agent of P. truncatus in west Africa.     

The Effects of an entomopathogenic fungus, Beauveria bassiana (balsamo) vuillemin (hyphomycetes), on Prostephanus truncatus (horn) (Col.: Bostrichidae), Sitophilus zeamais motschulsky (Col.: Curculionidae), and grain losses in stored maize in the Benin Republic

A fungal entomopathogen, Beauveria bassiana, was used to treat maize ears placed in traditional grain stores against Prostephanus truncatus in a field experiment conducted from September 1997 to March 1998 in the Benin Republic, West Africa. Treatments included oil-based spray with and without conidia, maize stored with and without the husk, and stores with and without artificial infestation. Additional treated ears kept in insect-proof cages under field conditions were sampled weekly and exposed to insects to estimate the virulence and persistence of the pathogen during the storage season. P. truncatus densities were significantly lower in treatments that included conidia, although densities were high in all artificially infested treatments and grain losses were severe. The effect of the pathogen was modeled with an exponential decay function and incorporated in a published P. truncatus simulation model. The effects of hypothetical pathogens with different virulence and persistence characteristics were evaluated in terms of insect density and percentage grain loss.     

Studies on predation of Prostephanus truncatus (Horn) (Col., Bostrichidae) and Sitophilus zeamais Mots. (Col., Curculionidae) at different densities on maize by Teretriosoma nigrescens Lewis (Col., Histeridae)

Laboratory experiments using whole cobs were conducted to examine the effect of varying densities of the larger grain borer, Prostephanus truncatus and the maize weevil, Sitophilus zeamais on rate of population increase by the histerid beetle, Teretriosoma nigrescens , a predator primarily of P. truncatus . Densities of all species of insects were determined at the end of the experiment, and an electrophoretic analysis of gut content was conducted on larval and adult T. nigrescens sampled during the experiments. Results indicated that T. nigrescens has a strong preference for P. truncatus and densities of T. nigrescens were associated only with densities of P. truncatus . The maize weevil played little role as an alternative prey or in interfering with T. nigrescens reproduction.     

Enumerative and binomial sequential sampling plans for the multicolored Asian lady beetle (Coleoptera: Coccinellidae) in wine grapes

To develop a practical integrated pest management (IPM) system for the multicolored Asian lady beetle, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), in wine grapes, we assessed the spatial distribution of H. axyridis and developed eight sampling plans to estimate adult density or infestation level in grape clusters. We used 49 data sets collected from commercial vineyards in 2004 and 2005, in Minnesota and Wisconsin. Enumerative plans were developed using two precision levels (0.10 and 0.25); the six binomial plans reflected six unique action thresholds (3, 7, 12, 18, 22, and 31% of cluster samples infested with at least one H. axyridis). The spatial distribution of H. axyridis in wine grapes was aggregated, independent of cultivar and year, but it was more randomly distributed as mean density declined. The average sample number (ASN) for each sampling plan was determined using resampling software. For research purposes, an enumerative plan with a precision level of 0.10 (SE/X) resulted in a mean ASN of 546 clusters. For IPM applications, the enumerative plan with a precision level of 0.25 resulted in a mean ASN of 180 clusters. In contrast, the binomial plans resulted in much lower ASNs and provided high probabilities of arriving at correct "treat or no-treat" decisions, making these plans more efficient for IPM applications. For a tally threshold of one adult per cluster, the operating characteristic curves for the six action thresholds provided binomial sequential sampling plans with mean ASNs of only 19-26 clusters, and probabilities of making correct decisions between 83 and 96%. The benefits of the binomial sampling plans are discussed within the context of improving IPM programs for wine grapes.     

Enumerative and binomial sequential sampling plans for soybean aphid (Homoptera: Aphididae) in soybean

Since the discovery of the soybean aphid, Aphis glycines Matsumura, in midwestern U.S. soybean, Glycine max L., in 2000, the aphid has become a significant economic pest. Basic information about estimating population density within fields is unknown. Therefore, we developed two sampling plans to efficiently characterize A. glycines densities. Enumerative and binomial sequential plans were developed using 89 data sets collected from 10 commercial fields sampled during 2001-2003. Re-sampling software was used to validate the enumerative plan on whole plant counts, based on Taylor's power law parameters (a = 9.157 and b = 1.543). For research applications, the enumerative plan was modified to provide an actual precision level of 0.10 (SE/mean), which resulted in an average sample number of 310 individual plants. For integrated pest management (IPM) purposes, we developed an enumerative plan with an actual precision of 0.25, which resulted in an average sample number of 38 individual plants. For IPM applications, the binomial plan will likely be more practical. Binomial plans were developed using two tally thresholds at five action thresholds. Final analysis of the operating characteristic curve for each plan indicated that the tally threshold of > or = 40 aphids per plant, and an action threshold of 0.837 (84% of the plants infested) provided the most correct treat (4%) and no-treat (95%) decisions, with very low incorrect treat (0.5%) and no-treat (0.5%) decisions. A tally threshold of > or = 40 aphids per plant and action thresholds of 84% of plants infested is equivalent to a mean density of 250 aphids per plant, a recently recommended economic threshold. Using this threshold, the minimum required sample number for the binomial plan was 11 plants.     

Isolation and characterization of microsatellite markers from Teretrius nigrescens Lewis (Coleoptera: Histeridae), predator of the storage pest Prostephanus truncatus (Horn) (Coleoptera: Bostrichidae)

Teretrius nigrescens is a predator of the larger grain borer (LGB) Prostephanus truncatus, an invasive post-harvest pest in Africa. We describe the isolation and characterization of 24 novel polymorphic microsatellite markers and their testing on a population from Honduras. Alleles per locus ranged between 2 and 12, and observed heterozygosity between 0.037 and 0.646. Six loci deviated significantly from Hardy-Weinberg equilibrium and showed evidence of null alleles. These markers will be useful for studies of the predator's population structure and characterizing populations for control of LGB.     

Presence-absence sequential sampling plan for northern fowl mite, Ornithonyssus sylviarum (Acari: Macronyssidae), on caged-layer hens

Caged-layer hens were scored as infested or uninfested by visual examination of the vent region, and the number of northern fowl mite, Ornithonyssus sylviarum (Canestrini & Fanzago), per hen was estimated. The proportion infested and average number of mites per hen were shown to have a highly significant, positive relationship (r = 0.936). Sampling among houses within a flock, and rows and sections within houses were analyzed to determine the reliability of sampling a representative portion of a flock. Low- and moderate-tolerance treatment thresholds, based on percentage of hens infested with mites, were developed from sampling 1 wk before and 1 wk after acaricide treatments determined necessary by the producer. These thresholds were used to compare a fixed (single) sampling plan, a curtailed procedure of the fixed sampling plan, and a sequential sampling plan based on a sequential probability ratio test, by sampling 174 hens (the maximum number needed for the single sampling plan). The sequential sampling plan required fewer hen examinations on average to reach a treatment decision than did the other plans, depending on the infestation tolerance limits. Using a low tolerance approach in which infestations below 15% are considered noneconomic (safe threshold) and infestations above 25% are considered economically important (action threshold), as few as 5 hens required examination to reach a treatment decision. Sequential sampling plan graphs are presented for 2 tolerance threshold scenarios (a 15% safe-threshold paired with a 25% action threshold and a 35% safe-threshold paired with a 45% action threshold). These sequential sampling plans using presence absence assessments should greatly facilitate monitoring and treatment decisions for this important pest.     

Sequential sampling plans for the hairy chinch bug (Hemiptera: Lygaeidae)

Two sequential sampling plans were developed to produce tools to reduce the use of pesticides for the control of the hairy chinch bug, Blissus leucopterus hirtus Montandon, on turfgrass lawns in the cool-humid region of Quebec, Canada. A first plan based on Wald's method was shown to be too conservative in a validation conducted on infested lawns. The second plan developed using Iwao's method yielded good results. The difference between the two plans may have occurred because no common k could be found for the negative binomial functions describing chinch bug distribution on lawns, thereby violating an essential assumption of Wald's approach. Application of the Iwao plan, which is based on visual sampling of 0.1-m2 quadrats, requires approximately 20 min when lawn evaluation is conducted by a single person. Estimation of the occurrence of chinch bug infestations (28% infested lawns in the regions of Montreal and Quebec cities), and results from validation of the Iwao plan on infested lawns, indicate that the error rates alpha and beta of the plan are lower than an upper limit of 0.05 and 0.14, respectively. The adequate precision and practicality of this plan suggest that it could become an essential tool for management of turfgrass lawns in regions where the hairy chinch bug completes less than two generations per growing season.     

Spatial distribution and sampling plans for Thrips palmi (Thysanoptera: Thripidae) infesting fall potato in Korea

The spatial distribution of adult and immature Thrips palmi Karny on fall potato, Solanum tuberosum L., on Cheju Island, Korea, was studied over a 2-yr period by visually inspecting potato leaves. The majority of thrips collected from the leaves were observed in the top one-third of the plant. The within-field spatial patterns of adults and immature thrips were aggregated. The slopes and intercepts of Taylor's power law did not differ among adults and immature thrips. A fixed-precision-level sampling plan was developed using the parameters from Taylor's power law and was tested with resampling simulations using eight independent data sets. Over a wide range of densities, the simulation demonstrated that actual sampling precision (d = SEM/mean) values at d = 0.25 averaged < 0.24 in all cases. A binomial sampling plan for estimating mean density was developed using an empirical model evaluated at tally thresholds (the minimum number of insects present before a leaf is considered infested) of one, three, five, and eight thrips per leaf. Increasing sampling size had little effect on the precision of the estimated mean regardless of tally threshold (T). However, increasing T had a dramatic effect on precision. The best tally threshold for estimating thrips density based on the applicable density ranges and the precision of the model was T = 5. A binomial sampling plan with a tally threshold of five and a fixed sample size of 30 leaves should be an effective replacement for enumerative counts when thrips average < 10 per leaf.     

Binomial sequential sampling plan for hemlock woolly adelgid (Hemiptera: Adelgidae) sistens infesting individual eastern hemlock trees

The hemlock woolly adelgid, Adelges tsugae Annand (Hemiptera: Adelgidae), is an exotic insect pest that is killing eastern hemlock, Tsuga canadensis (L.) Carrière, and Carolina hemlock, Tsuga caroliniana Engelmann, in the eastern United States. We used the sequential interval procedure to develop a binomial sequential sampling plan for A. tsugae sistens on individual eastern hemlock trees that uses nondestructive sampling of new shoots. The actual a (type I) and beta (type II) error rates were essentially 5 and 10%, respectively. Tallies of new shoots infested by at least one A. tsugae sistens were compared with stop values for thresholds of 10 and 30% of new hemlock shoots infested. Twenty to 80 new shoots had to be examined per tree to render a low, high, or indeterminate classification, which took < 2 min per tree regardless of the threshold used. This plan should be an efficient and cost-effective tool in the management of A. tsugae infestations on individual, high-value eastern hemlock trees.     

Efficacy of Xylocoris flavipes (Reuter) (Het., Anthocoridae) to suppress Prostephanus truncatus (Horn) (Col., Bostrichidae) in traditional maize stores in southern Togo

In the course of three storage periods (1988–90) the predacious insects that occur in traditional maize stores in southern Togo were observed and their relationship to the introduced pest Prostephanus truncatus was investigated. The bug Xylocoris flavipes , a predator on a wide range of post-harvest pests, was the only one observed regularly and in higher numbers. Laboratory trials with P. truncatus showed that the larvae were preyed on, and no preference for any one of the three instars was detected. On average, 7.3 pest larvae were killed by larvae or adults of the predator. In glass jar trials on loose maize X. flavipes reduced the numbers of P. truncatus by 57.6% after 8 weeks. The losses and damage were reduced by 50 and 17.6%, respectively. Observations in traditional maize stores indicated a tendency to decreasing populations of X. flavipes when populations of P. truncatus increased, presumably due to the unfavourable environmental conditions created by the pest. Therefore, it is concluded that X. flavipes does not play an important role in the control of this pest.     

Operating characteristics of full count and binomial sampling plans for green peach aphid (Hemiptera: Aphididae) in potato

Counts of green peach aphid, Myzus persicae (Sulzer) (Hemiptera: Aphididae), in potato, Solanum tuberosum L., fields were used to evaluate the performance of the sampling plan from a pest management company. The counts were further used to develop a binomial sampling method, and both full count and binomial plans were evaluated using operating characteristic curves. Taylor's power law provided a good fit of the data (r2 = 0.95), with the relationship between the variance (s2) and mean (m) as ln(s2) = 1.81(+/- 0.02) + 1.55(+/- 0.01) ln(m). A binomial sampling method was developed using the empirical model ln(m) = c + dln(-ln(1 - P(T))), to which the data fit well for tally numbers (T) of 0, 1, 3, 5, 7, and 10. Although T = 3 was considered the most reasonable given its operating characteristics and presumed ease of classification above or below critical densities (i.e., action thresholds) of one and 10 M. persicae per leaf, the full count method is shown to be superior. The mean number of sample sites per field visit by the pest management company was 42 +/- 19, with more than one-half (54%) of the field visits involving sampling 31-50 sample sites, which was acceptable in the context of operating characteristic curves for a critical density of 10 M. persicae per leaf. Based on operating characteristics, actual sample sizes used by the pest management company can be reduced by at least 50%, on average, for a critical density of 10 M. persicae per leaf. For a critical density of one M. persicae per leaf used to avert the spread of potato leaf roll virus, sample sizes from 50 to 100 were considered more suitable.     

Binomial sampling for pest management of stable flies (Diptera: Muscidae) that attack dairy cattle.

A binomial sampling plan for pest management of the stable fly, Stomoxys calcitrans (L.), was developed. Counts of stable flies on front legs of the same animal were independent and each leg from the same animal was considered a sample unit. The relationship between the mean number of flies per leg and the variance was determined and did not vary among farms. The relationship between the mean number of flies per leg and the proportion of legs with zero, one or less, and two or less flies (P0, P1, and P2) was determined and used as the basis of the binomial sampling plan. Predicted values of the mean number of flies per leg from P0, P1, and P2 were close to observed values of the mean number of flies per leg. Equations are presented for calculating the variance of a predicted value of the mean number of flies per leg using values of P0, P1 and P2 determined by sampling. Operating characteristic (OC) curves are also presented for determining the probability of making a treatment decision error at an economic threshold of one fly per leg (P0 = 0.47) using binomial sampling or direct counting. OC curves for binomial sampling with n = 50 legs were close to those for direct counting with n = 10 legs. Recommendations concerning the use of binomial sampling for stable flies are presented.     

Pest management in traditional maize stores in West Africa: a farmer's perspective

Farmers in the Republic of Benin have few resources to invest in protection of stored maize, and prophylactic pesticide application is often recommended by extension and development agencies. Neither the efficacy nor profitability of such an application in traditional maize storage facilities has been addressed quantitatively. In this study, existing management options for stored maize were evaluated monthly over 6 mo in central and southern Benin with respect to their effects on grain injury and on densities of Prostephanus truncatus (Horn) and Sitophilus zeamais Motschulsky. P. truncatus infested 54% of the experimental stores in the study even though Teretrius nigrescens (Lewis), a natural enemy introduced against P. truncatus, was well established in the region. S. zeamais was the most common pest, found in 85% of the experimental storage facilities. Prophylactically treated maize was, on average, worth more than untreated maize for month 1 through 5 in southern Benin, after taking into account market price, pesticide costs, percentage grain damage and weight loss, but maize storage was not profitable overall. No difference was observed between treatments in central Benin. After 6 mo treated storage facilities were not significantly different from untreated storage facilities in terms of either percentage damage or profit in either region. A rapid scouting plan intended to provide farmers with a means for identifying storage facilities at greatest risk of severe P. truncatus infestation was field validated. Given that unsafe pesticide use is prevalent in Benin, research and extension services should clearly state the limitations to prophylactic treatment and increase the effort to educate farmers on appropriate pesticide use, store monitoring and marketing.     

Cost-effective binomial sequential sampling of western bean cutworm, Striacosta albicosta (Lepidoptera: Noctuidae), egg masses in corn

Striacosta albicosta (Smith) (Lepidoptera: Noctuidae), is a native pest of dry beans (Phaseolus vulgaris L.) and corn (Zea mays L.). As a result of larval feeding damage on corn ears, S. albicosta has a narrow treatment window; thus, early detection of the pest in the field is essential, and egg mass sampling has become a popular monitoring tool. Three action thresholds for field and sweet corn currently are used by crop consultants, including 4% of plants infested with egg masses on sweet corn in the silking-tasseling stage, 8% of plants infested with egg masses on field corn with approximately 95% tasseled, and 20% of plants infested with egg masses on field corn during mid-milk-stage corn. The current monitoring recommendation is to sample 20 plants at each of five locations per field (100 plants total). In an effort to develop a more cost-effective sampling plan for S. albicosta egg masses, several alternative binomial sampling plans were developed using Wald's sequential probability ratio test, and validated using Resampling for Validation of Sampling Plans (RVSP) software. The benefit-cost ratio also was calculated and used to determine the final selection of sampling plans. Based on final sampling plans selected for each action threshold, the average sample number required to reach a treat or no-treat decision ranged from 38 to 41 plants per field. This represents a significant savings in sampling cost over the current recommendation of 100 plants.     

Binomial sequential sampling plans for late instars of European corn borer (Lepidoptera: Crambidae), corn earworm (Lepidoptera: Noctuidae), and damaged kernels in sweet corn ears

Late-season infestations of European corn borer, Ostrinia nubilalis (Hübner), and corn earworm, Helicoverpa zea (Boddie), were sampled to develop binomial sequential sampling plans for larval infestations and damaged kernels in sweet corn, Zea mays L., ears, near harvest. Fields were sampled to obtain a range of larval densities likely to be encountered over a range of infestation levels and field conditions. Binomial sampling plans were developed for O. nubilalis larvae, H. zea larvae, O. nubilalis, and H. zea larvae combined, and for damaged sweet corn kernels. Observed densities ranged from 0.01 to 4.40 larvae per ear for O. nubilalis, 0.005-1.62 larvae per ear for H. zea, and 0.004-36.12 damaged kernels per ear. Results of resampling analyses, based on the proportion of ears infested with one or more larvae, or damaged kernels, indicated an average sample size of 34-37 ears was necessary to classify whether larval infestations, or the incidence of damaged kernels, exceeded 5%. Two operating characteristic curves are presented for each of the four sampling plans. Initial results, with upper bounds of 0.10, and alpha (type I) and beta (type II) error rates at 0.10 and 0.05, respectively, resulted in a 90% probability of making the correct management decision at infestation levels >10%. To improve performance of the sampling plans, we modified the binomial plans by reducing the upper bound to 0.075, while maintaining the same error rates. This plan resulted in a higher probability (>95%) of making the correct management decision to reject a sweet corn load when infestation levels are >10%.     

Transect sampling to enhance efficiency of corn rootworm (Coleoptera: Chrysomelidae) monitoring in corn

Crop monitoring for adult corn rootworms, Diabrotica virgifera virgifera LeConte and Diabrotica barberi Smith and Lawrence, remains the best means to assess fields at risk from this pest if replanted to corn, Zea mays (L.). Improvements in sampling methodology, including the development of a sequential sampling plan, have reduced the minimum sampling time required to make a management decision to 20 min or less per field per visit. However, many growers and crop consultants still find this time commitment a constraint to repeated scouting. A common currently used sampling method involves systematically covering most of the field following a "W" pattern. The feasibility of replacing the current sampling pattern with a simpler and less time-consuming transect (straight line) pattern was assessed. When sampling methods were compared, computer simulations demonstrated that treatment decisions based on transect sampling would have an acceptably low error rate averaging 10% over a range of realistic corn rootworm densities (0-2 adults per plant). This error rate represented a decrease in accuracy of <1% compared with systematic sampling. Field trials using transect, systematic, and random sampling in each field were used to compare the categorization of adult corn rootworm densities into "above" or "below" threshold with a sequential sampling plan. Efficiency measured in time to reach a decision, number of corn plants evaluated, and time divided by plants observed were compared between sampling methods. The three methods did not differ significantly in the number of plants evaluated or in the categorization of corn rootworm populations. Transect sampling resulted in a significantly shorter time divided by plants observed (38 s), than either systematic (78 s), or random sampling methods (166 s). Based on these results transect sampling reduces sampling time 51% compared with systematic sampling and thus could be used to reduce total sampling times substantially.