Biological Control 58 (2011) 127–132 Contents lists available at ScienceDirect Biological Control journal homepage: www.elsevier .com/locate /ybcon Comparative biology and production costs of Podisus nigrispinus (Hemiptera: Pentatomidae) when fed different types of prey Sergio A. De Bortoli a, Alessandra K. Otuka a, Alessandra M. Vacari a,⇑, Maria I.E.G. Martins b, Haroldo X.L. Volpe a a Laboratório de Biologia e Criação de Insetos, Departamento de Fitossanidade, Universidade Estadual Paulista FCAV/Unesp, Via de Acesso Prof. Paulo Donato Castellane s/n, 14884-900 Jaboticabal SP, Brazil b Departamento de Economia Rural, Universidade Estadual Paulista FCAV/Unesp, Via de Acesso Prof. Paulo Donato Castellane s/n, 14884-900 Jaboticabal SP, Brazil a r t i c l e i n f o Article history: Received 9 February 2011 Accepted 18 April 2011 Available online 24 April 2011 Keywords: Heteropteran predator Mass rearing Biological control Asopinae 1049-9644/$ - see front matter Crown Copyright � 2 doi:10.1016/j.biocontrol.2011.04.011 ⇑ Corresponding author. E-mail address: amvacari@gmail.com (A.M. Vacari a b s t r a c t The objective of our work was to identify the prey that offers optimum production of the predator, Podisus nigrispinus (Dallas). The cost of predator production was determined for each prey. We test the hypothesis that prey types supporting optimal development do not facilitate cost-effective rearing of P. nigrispinus. Second-instar nymphs of P. nigrispinus were reared in 1000-ml transparent plastic containers and maintained under controlled conditions (temperature, 25 ± 1 �C; photophase duration, 12 h; relative humidity, 70 ± 10%), and the following biological aspects were evaluated: nymph viability, length of the nymphal period, and weight of fifth-instar nymphs. The adults were isolated in pairs (one female and one male) in Petri dishes, and the following biological aspects were evaluated: lifespan, adult weight after emergence, number of eggs per female, egg mass in females, number of eggs per egg mass, durations of the pre-oviposition, oviposition, and post-oviposition periods, viability, and length of the embryonic period. On the basis of our results, we conclude that the prey that leads to optimal development of P. nigrispinus is the larvae of Diatraea saccharalis (Fabr.), and the most cost effective prey for rearing P. nigrispinus is the larvae of Musca domestica L. Crown Copyright � 2011 Published by Elsevier Inc. All rights reserved. 1. Introduction factitious prey has been investigated. Feeding an artificial diet Since integrated pest management in agricultural crops was introduced to reduce the use of pesticides, entomologists have faced major challenges in marketing and increasing the use of nat- ural enemies as a method of biological control. Solutions to these challenges include the reduction in the production costs of these agents by improving rearing techniques and increasing their effec- tiveness in biological control in the field (Tauber et al., 2000). The effectiveness of using natural enemies as biological control agents depends on how efficiently they can be reared in the labo- ratory. Chambers (1977) stated that mass rearing is the production of insects with an acceptable cost/benefit ratio. The study of tech- nical and economic aspects of rearing a natural enemy is crucial to its effective use as a biological control agent in the field or green- houses. According to van Lenteren (2000), the development of techniques of mass production, quality control, storage, transport, and release of natural enemies can lead to a decrease in production costs and improvement in the quality of the product. The potential of using artificial diets to mass produce generalist predators, including predatory heteropterans, in lieu of natural or 011 Published by Elsevier Inc. All r ). rather than natural or factitious prey to the predator, Podisus nigri- spinus (Dallas) (Hemiptera: Pentatomidae), does not have any ben- efits (Lemos et al., 2003). For example, predator nymphs reared on an artificial diet weighed significantly less than cohorts fed their natural prey, Alabama argillacea Hübner (Lepidoptera: Noctuidae) larvae. Moreover, fresh weight of the ovaries of newly emerged P. nigrispinus females was the least for individuals reared on artificial diet than on the yellow mealworm, Tenebrio molitor L. (Coleoptera: Tenebrionidae), house fly, Musca domestica L. (Diptera: Muscidae), or cotton leafworm, Spodoptera littoralis (Boisduval) (Lepidoptera: Noctuidae) larvae. In a subsequent study, Lemos et al. (2005) dis- covered that food source had a profound impact on ovarian devel- opment. Newly emerged P. nigrispinus females had ovaries containing oocytes in an advanced stage of development when fed A. argillacea larvae, intermediate stage oocytes when fed T. mol- itor or M. domestica larvae, and early stage oocytes when fed an artificial diet. Moreover, P. nigrispinus showed good growth and high fecundity when fed a factitious prey, Diatraea saccharalis (Fabr.) (Lepidoptera: Crambidae) (Vacari et al., 2007). Research on the usefulness of other potential factitious prey that can be pro- duced at low cost is being conducted (Riddick, 2009). There are some companies that sell Podisus spp. in the United States and Europe to control caterpillars. In the United States, 250 ights reserved. http://dx.doi.org/10.1016/j.biocontrol.2011.04.011 mailto:amvacari@gmail.com http://dx.doi.org/10.1016/j.biocontrol.2011.04.011 http://www.sciencedirect.com/science/journal/10499644 http://www.elsevier.com/locate/ybcon 128 S.A. De Bortoli et al. / Biological Control 58 (2011) 127–132 eggs of the predator Podisus maculiventris (Say) (Hemiptera: Pent- atomidae) are sold for $133.50 (http://www.arbico-organics.com/). In Minas Gerais State, Brazil, timber companies such as ‘‘CAF Florestal Ltda’’ (Belo Horizonte), ‘‘V & M Florestal Ltda’’ (Paraopeba), and ‘‘Refloralje’’ (Montes Claros) have established laboratories to produce Asopinae predators. The mass production and release of P. nigrispinus are based on the rotation cycle of eucalyptus forests (ca. 6–7 years) and forecasts of outbreak periods, allowing the adoption of inoculative release of this predator in areas susceptible to outbreaks (Torres et al., 2006; Vacari and De Bortoli, 2010). P. nigrispinus is generally reared and maintained in the labora- tory; it is fed on the larvae and pupae of T. molitor (Zanuncio et al., 2001; Oliveira et al., 2002; Vivan et al., 2002) because this in- sect is easy to rear. However, there are no reports in the literature on the production cost of this natural enemy. Therefore, the objec- tive of our study was to determine the prey that offers optimum conditions for the establishment of P. nigrispinus and to calculate the production cost for each prey tested. In this study, we test the hypothesis that prey types supporting optimal development do not facilitate cost-effective rearing of the predator P. nigrispinus. 2. Materials and methods The study was conducted at the Laboratory of Biology and Rear- ing of Insect (LBRI), FCAV/Unesp, Jaboticabal, São Paulo. The prey used included larvae of Anticarsia gemmatalis Hübner and Spodop- tera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae) obtained by mass rearing at Dupont Agrosciences in Brazil located in Paulinia, São Paulo; larvae of D. saccharalis, obtained by mass rearing at Usi- na Santa Adelia located in Jaboticabal, São Paulo; larvae of T. mol- itor, obtained by mass rearing at LBRI, FCAV/Unesp; and larvae of M. domestica, obtained by mass rearing at the Laboratory of Frog Culture, CAUNESP, Unesp/Jaboticabal. The nymphs of P. nigrispinus were obtained by mass rearing at the LBRI, Department of Plant Protection, FCAV/Unesp, Campus de Jaboticabal. 2.1. Comparative biology For the experiment, 100 second-instar nymphs were used per treatment. They were placed in two transparent 1000-ml plastic containers (50 nymphs per container); third-instar larvae were added in the container ad libitum as prey for the predators. The in- sects were maintained at a room temperature of 25 ± 1 �C, with a relative humidity of 70 ± 10% and a photophase duration of 12 h. Evaluations were performed daily. Water was provided using a tube fixed on the lid of the container, with the tip turned inwards and a piece of cotton placed in the tube hole to allow supply of water by capillary action. We measured the length of the nymphal period, nymphal consumption, and the weight of fifth-instar nymphs. After emergence, the adults were isolated in pairs (one female and one male) and transferred to Petri dishes (9 cm � 1.5 cm). The prey species provided to the predators in the nymphal stage were also provided to the predators in the adult stage, and water was supplied via a cotton swab placed in the lid. All eggs were col- lected, counted, and placed in Petri dishes (6 cm � 2 cm) until hatching. For the adults, the following were determined: weights of recently emerged males and females; lifespan; adult consump- tion; number of eggs per female; number of eggs per clutch; and durations of the pre-oviposition, oviposition, and post-oviposition periods. For the eggs, length of incubation period and viability were evaluated. From the biological data of P. nigrispinus, the following param- eters were determined to construct a fertility life table by using the methods of Birch (1948), Silveira Neto et al. (1976), Southwood (1978), and Price (1984): x, the average age of parental females since emerging from egg stage; lx, life expectancy to age x, ex- pressed as a fraction of females; mx, specific fertility or number of descendents per female produced at age x and that result in fe- males; lx.mx, total number of females born at age x. The growth parameters obtained from the life table were calculated as de- scribed by the authors, with R0 as the net reproductive rate (i.e., the rate of population increase), considering the females from one generation to another or the number of females produced per parental female per generation; T, mean generation time or average lifespan of one generation; rm, intrinsic rate of increase in number; and k, finite rate of increase, defined as the number of times the population multiplies in unit time. In addition to these parameters, TD, the time required for the population to double in number, was also determined according to the method of Krebs (1994). The growth parameters (R0, T, rm, k, and Dt) were calculated using the following equations: R0 ¼ Rðmx:lxÞ T ¼ ðRmx:lx:xÞ=ðRmx:lxÞ rm ¼ log R0=Tð0:4343Þ k ¼ anti-logðrm ¼ 0:4343Þ Dt ¼ Lnð2Þ=rm The experimental design was completely randomized, with five treatments; each treatment included two rearing containers, each containing 50 nymphs of P. nigrispinus. The fertility life table analyses and mean comparisons were done using PROC GLM (SAS Institute, 2002), as described by Maia et al. (2000). The proportion of surviving adults was compared be- tween the treatments by the Kaplan–Meier method, using PROC LIFETEST (SAS Institute, 2002). The values of the life table parameters were subjected to multi- variate cluster analysis (Sneath and Sokal, 1973), and all biological characteristic data were used in the two-way analysis to determine the maximum similarity or the maximum dissimilarity between the groups by using the program Statistica version 7.0 (StatSoft Inc., 2004). 2.2. Production cost In the second step, we calculated the predator production cost. For this, we used information on the establishment of P. nigrispinus in 1000-ml plastic containers with 50 nymphs each. In Brazil, the production cost consists of the total production cost and the operational cost (Scorvo-Filho et al., 2004). The first includes all operating expenses plus depreciation and the inter- est on working capital and land remuneration of the investment and of the entrepreneur. The second involves only the operating expenses, including finance charges and depreciation of the physical structure and equipment specific to the activity studied. The total production cost can be divided into fixed and variable costs. Fixed costs are those that do not change with the quantity produced. Variable costs are those that depend on the produc- tion volume. In this study, we considered only the total operational cost (effective operational cost + depreciation cost of specific items for the prey) because we assumed that the producer has a bio-factory where other natural enemies are reared. The following definition of total operating cost was established by Matsunaga (1976) because of the difficulties in evaluating some fixed-cost items, such as land remuneration of fixed capital and of http://www.arbico-organics.com/ S.A. De Bortoli et al. / Biological Control 58 (2011) 127–132 129 the entrepreneur, given the subjectivity of their meaning: it consists of items considered as direct costs (labor, supplies, equip- ment maintenance, and other fees) and costs representing the depreciation of specific materials Matsunaga (1976). Thus, in this structure, the following costs are included: (1) effective opera- tional cost, which represents the annual expenditure incurred for the activity, and (2) total operating cost, which corresponds to effective operating costs, depreciation of the equipment, and spe- cific costs on furniture. The operating cost calculation allows a comparison of the product to that offered on the market. Such information is impor- tant to know whether it is feasible to produce the product and introduce it to the desired market segment (Reis, 2002). Our study considered effective operational costs, including expenses on diet, labor, energy consumed by the equipment, and other materials, as well as depreciation of the equipment and specific expenses on furniture. Effective operational costs on prey were estimated from the data obtained from the production unit of each insect. The depreciation cost of specific equipment and fur- niture was calculated as follows: D ¼ V VuHu � C where D , depreciation; V , value of the equipment; Vu, life of the equipment (in years); Hu, hours of use per year; and C, cycle length in hours. Adding the operational cost to the cost of depreciation gives the total operating cost of production per cycle (in dollars): TOC ¼ EOCþ depreciation cost Where TOC, the total operating cost and EOC, the cost-effective operational cost. The costs of each prey and each predator were also obtained by dividing the total operating cost by the number of individuals pro- duced per cycle: Insect Costi ¼ TOCi Ni where TOCi = total operating cost of the insect species i; Ni = num- ber of individuals of species i produced; and i = number of species (1–5). N ym ph al p er io d N ym ph al v ia bi lit y M al e w ei gh t O vi po si tio n P os t- ov ip os iti on E gg m as s/ fe m al e F em al e w ei gh t E gg s/ eg g m as s M. domestica T. molitor A. gemmatalis D. saccharalis S. frugiperda Fig. 1. Two-way analysis of the biological characteristics 3. Results 3.1. Comparative biology To identify which prey led to optimum development of the predator P. nigrispinus, we used multivariate statistical techniques that allow the simultaneous evaluation of different biological char- acteristics. Thus, applying a 2-way analysis allowed us to observe the influence of different types of prey on the biological character- istics of the predator. Feeding D. saccharalis larvae resulted in a higher number of nymphs (F4,15 = 5.17; P = 0.0081), and heavier adult females (F4,15 = 5.46; P = 0.0064) with greater fecundity (F4,15 = 10.84; P = 0.0002). In addition, adult females had a shorter pre-oviposition period (F4,15 = 7.52; P = 0.0016) and better repro- ductive characteristics in a shorter time (F4,15 = 4.16; P = 0.0183) (Fig. 1). Feeding on different types of prey showed differential effects on the parameters of the fertility life table of P. nigrispinus (Table 1). The net reproductive rate (R0) of the predator was higher when it was fed larvae of D. saccharalis (213.1), S. frugiperda (162.3), and M. domestica (165.3). With regard to the mean generation time (T), the highest values were found for M. domestica (41.6) and D. saccharalis (39.3), and the lowest value for S. frugiperda (36.3). The intrinsic rate of increase (rm) and the finite rate of increase (k) were significantly higher for S. frugiperda (0.140 and 1.150, respectively), but similar between A. gemmatalis, D. saccharalis, and T. molitor, with values of 0.126 and 1.134, respectively, for A. gemmatalis, 0.137 and 1.146, respectively, for D. saccharalis, and 0.131 and 1.140, respectively, for T. molitor. The population dou- bling time (Dt) was significantly shorter for S. frugiperda (5.0), although it was similar among A. gemmatalis (5.5), D. saccharalis (5.1), and T. molitor (5.3). The fertility life table of P. nigrispinus showed that D. saccharalis and S. frugiperda had higher net repro- ductive rates, indicating production of a greater number of off- spring per generation. Females that fed on M. domestica also showed higher mean generation time (T) and longer population doubling time (Dt) because of longer lifespan and lower offspring production during their lifetime (R0, net reproductive rate). For the parameters of the fertility life table, the dendrogram shows the formation of 2 groups, A and B (Fig. 2). M. domestica shows the greatest contrast to the other types of prey and does not belong to any group. The group formed by D. saccharalis and S. frugiperda provide better fertility life table parameters for E gg s vi ab ili ty F em al e lo nv eg ity P re -o vi po si tio n S ex r at io E m br io na ry p er io d M al e lo ng ev ity N ym ph al w ei gh t E gg s/ fe m al e 1.5 1 0.5 0 -0.5 -1 -1.5 of Podisus nigrispinus fed on different types of prey. Table 1 Parameters of the fertility life table of Podisus nigrispinus fed on different types of prey. Preys R0 T rm k Dt Spodoptera frugiperda 162.3 ± 35.85ab1 36.3 ± 1.05c 0.140 ± 0.0037a 1.150 ± 0.0043a 5.0 ± 0.13b (126.41–198.12)2 (35.23–37.34) (0.1360–0.1434) (1.1456–1.1542) (4.83–5.10) Anticarsia gemmatalis 126.3 ± 13.63b 38.4 ± 1.25bc 0.126 ± 0.0061ab 1.134 ± 0.0070ab 5.5 ± 0.27ab (112.67–139.94) (37.17–39.67) (0.1198–0.1321) (1.1273–1.1412) (5.24–5.78) Diatraea saccharalis 213.1 ± 24.29a 39.3 ± 1.90ab 0.137 ± 0.0086ab 1.146 ± 0.0099ab 5.1 ± 0.32ab (188.84–237.41) (37.42–41.23) (0.1279–0.1452) (1.1364–1.1562) (4.78–5.41) Tenebrio molitor 133.4 ± 59.21b 36.7 ± 0.72bc 0.131 ± 0.0096ab 1.140 ± 0.0110ab 5.3 ± 0.39ab (74.18–192.60) (36.03–37.46) (0.1216–0.1408) (1.1293–1.1512) (4.91–5.70) Musca domestica 165.3 ± 14.32ab 41.6 ± 0.69a 0.123 ± 0.0032b 1.131 ± 0.0036b 5.6 ± 0.14a (150.99–179.63) (40.90–42.29) (0.1196–0.1260) (1.1270–1.1342) (5.51–5.79) F(4,15) 3.91 12.01 4.18 4.17 4.22 P 0.0228 0.0001 0.0181 0.0182 0.0173 R0, net reproductive rate (female/female); T, mean generation time (days); rm, intrinsic rate of increase; k, finite rate of increase (female/day); Dt, doubling time. 1 Means and confidence intervals followed by the same letter in the same column did not differ significantly (Tukey’s test, P > 0.05; F(DF, Error)); 2 Confidence limits. 130 S.A. De Bortoli et al. / Biological Control 58 (2011) 127–132 predators, and is more suitable for the establishment of P. nigrispi- nus, with the band showing the largest Euclidean distance in rela- tion to other prey types. T. molitor and A. gemmatalis are similar, forming another group (B) that is considered intermediate prey for rearing P. nigrispinus (Fig. 2). M. domestica T. molitor A. gemmatalis D. saccharalis S. frugiperda 1.0 1.5 2.0 2.5 3.0 3.5 4.0 4.5 5.0 5.5 Eu cl id ia n di st an ce AB Fig. 2. Dendrogram showing the clustering of prey based on the life table parameters of Podisus nigrispinus fertility. Table 2 Operating costs per production cycle and per predator for Diatraea saccharalis, Anticarsia g Prey Diatraea saccharalis US$c/ cycle Anticarsia gemmatalis US$ cycle Matrices Labor cost 44.17 141.35 Costs on equipment and energy 70.55 291.47 Diet 29.21 29.21 Materials 2.48 0.49 Caterpillars Labor cost 128.83 212.02 Costs on equipment and energy 18.38 7.15 Diet 51.89 45.22 Materials 768.03 52.07 Operational cost 1117.59 778.96 Depreciation of specific materials 46.77 36.54 TOCa 1164.36 815.50 Productionb (individuals/ cycle) 39,520 50,000 TOC/insect 0.03 0.018 a Total operating cost (TOC) considering only the depreciation of specific materials. b Number of individuals produced per cycle was obtained from each production unit. c Currency Exchange quote for Brazilian Real (R$) to US Dollar (US$) 1.0 Brazilian Rea 3.2. Cost of production The quantitative planning in the rearing of P. nigrispinus was done considering the following data: a female predator fed on D. saccharalis produces an average of 797.7 eggs per cycle, S. fru- giperda averages 442.0, A. gemmatalis averages 441.6 T. molitor averages 328.6 and M. domestica averages 407.8. The data were estimated considering an initial rearing population density of 1200 predators. These eggs were kept in glass Petri dishes (12 cm � 20 cm) and had average viabilities of 90%, 86%, 81%, 70%, and 73% and nymphal viabilities of 76%, 60%, 55%, 61%, and 80%, respectively, with average productions of 654,760 adults per cycle with D. saccharalis, 236,079 with A. gemmatalis, 547,410 with S. frugiperda, 168,359 with T. molitor, and 285,758 with M. domes- tica. Of the total number of individuals produced, 1200 pairs were returned to rearing conditions; thus, this system produced around 652,360 adults per cycle with D. saccharalis, 233,679 with A. gemmatalis, 545,010 with S. frugiperda, 165,959 with T. molitor, and 283,358 with M. domestica, which can be commercially sold or released in the field. Since the insects were reared in an acclima- tized room, we obtained 7.5 cycles of 48–49 days per year with D. saccharalis and A. gemmatalis, 7.1 cycles of 51.2 days per year emmatalis, Spodoptera frugiperda, Tenebrio molitor, and Musca domestica (US$). / Spodoptera frugiperda US$/ cycle Tenebrio molitor US$/ cycle Musca domestica US$/ cycle 141.35 0.98 3.31 291.47 15.05 0.00 29.21 0.78 0.48 0.49 0.00 0.00 212.02 0.98 3.68 7.15 15.05 0.00 45.22 1.59 0.32 52.07 0.00 0.00 778.96 34.43 7.61 36.54 21.80 0.25 815.50 56.27 8.05 50,000 4000 8500 0.018 0.012 0.001 l = 0.601 US Dollar. Table 3 Operating costs (US$) (per production cycle and per predator) of Podisus nigrispinus fed on different types of prey. Treatments Podisus nigrispinus ?tul?> fed on Diatraea saccharalis US$b/cycle Podisus nigrispinus /ce:italic> fed on Anticarsia gemmatalis US$/cycle Podisus nigrispinus /ce:italic> fed on Spodoptera frugiperda US$/cycle Podisus nigrispinus /ce:italic> fed on Tenebrio molitor US$/cycle Podisus nigrispinus /ce:italic> fed on Musca domestica US$/cycle Matrices (1200 insects) Labor 39.78 39.78 39.78 39.78 39.78 Equipment and energy 49.52 49.52 49.52 49.52 49.52 Diet 45.01 26.84 27.06 17.32 2.39 Materials 1.47 1.47 1.47 1.47 1.47 Costs until adult emergence Labor 69.00 69.00 69.00 69.00 69.00 Equipment and energy 49.52 49.52 49.52 49.52 49.52 Diet 16,088.53 3314.66 8066.51 3789.42 499.64 Materials 2.10 2.10 2.10 2.10 2.10 Operational cost 16,345.03 3552.91 8304.97 4018.14 713.45 Depreciation of specific materials 14.74 14.74 14.74 14.74 14.74 TOCa 16,359.78 3567.65 8319.72 4032.89 728.19 Production (individual/cycle) 652,360 233,679 545,010 165,959 283,358 TOC/insect 0.024 0.018 0.018 0.024 0.002 a Total operating cost (TOC) considering only the depreciation of specific materials. b Currency Exchange quote for Brazilian Real (R$) to US Dollar (US$) 1.0 Brazilian Real = 0.601 US Dollar. S.A. De Bortoli et al. / Biological Control 58 (2011) 127–132 131 with S. frugiperda, 9.2 cycles of 39.7 days per year with T. molitor, and 7.7 cycles of 47 days per year with M. domestica. For D. saccharalis, the cost of production was higher because of the high cost of materials (cotton, string, petri dishes, etc.) used in the experiments (Table 2). The higher cost of producing M. domes- tica was because of the high labor costs. The estimated production cost of the predator P. nigrispinus in a laboratory rearing system was US$ 0.024 when fed with the prey D. saccharalis, US$ 0.018 with A. gemmatalis, US$ 0.018 with S. frugiperda, US$ 0.024 with T. molitor, and US$ 0.002 with M. domestica (Table 3). The cost of producing the predator was lower with M. domestica larvae, but its development was better with D. saccharalis larvae, although the predator’s nymphal period was shorter with both types of prey. The predator production was higher when fed on D. saccharalis, almost double that when fed on M. domestica. 4. Discussion The determination of population increase due to reproductive capacity is a crucial component of population studies. This increase can be analyzed with a fertility life table, which shows the repro- ductive potential of females at different times (Medeiros et al., 2000). The intrinsic rate of increase is represented by the combina- tion of R0 and T, and when the predator has lower offspring produc- tion in the same time span and/or has a longer lifespan, the rm value becomes smaller (Vacari, 2006). Vacari (2006) obtained the following values for the fertility life table parameters for P. nigrispinus that fed on D. saccharalis larvae: net reproductive rate (R0), 78.18; mean generation time (T), 41.38; intrinsic rate of increase (rm), 0.1053; finite rate of increase (k), 1.1111; and doubling time of the initial population (Dt), 6.58. These values show that this prey provides the best parameters in the life table. These values are different from the values obtained in our study. The laboratory conditions were similar, and the prey were from the same population. However, the predators were reared in different numbers and in different containers. Watanabe et al. (1997) also estimated the parameters for P. nig- rispinus that fed on A. gemmatalis larvae and found lower values, with the number of female offspring per female (R0) being only 13.02, mean generation time (T) being 13.02, the intrinsic rate of increase (rm) being 0.0809, and the doubling time of the initial population (Dt) being 8.85. The fertility life table showed that feed- ing on D. saccharalis larvae provided greater production of offspring per generation than feeding on A. gemmatalis larvae. The highest values of rm (combination of R0 and T) for the predators that fed on S. frugiperda and D. saccharalis can be explained by the greater production of offspring within the indicated time span and/or shorter lifespan (Evangalista Júnior et al., 2003). Evangalista Júnior et al. (2004) fed P. nigrispinus under condi- tions of partial prey scarcity (3 days without food) and provided T. molitor pupae and plants (Amaranthus hybridus L. [Amarantha- ceae], Desmodium tortuosum Sw. [Leguminoseae], Euphorbia hetero- phylla L. [Euphorbiaceae], Ageratum conyzoides L. [Compositae], Bidens pilosa L. [Asteraceae], Ricinus communis L. [Euphorbiaceae], and Gossypium hirsutum L. r. latifolium Hutch. [Malvaceae] cv. CNPA Early 1) as food supplements. They found that the life table param- eters were better with some plants (D. tortuosum and Ageratum conyzoides) than they were with A. argillacea larvae. The values of these parameters were higher for all the plants studied, even when there was no food scarcity and no plant supplements (the same as indicated above). The highest values for R0, T, rm, and k were 185.5 (A. conyzoides), 36.3 (A. conyzoides), 0.1525 (B. pilosa), and 1.1647 (B. pilosa), respectively (Evangalista Júnior et al., 2003). The values obtained by Peluzio (2008) for the parameters of the fertility life table showed that for this predator, T. molitor was more suitable than A. gemmatalis. This result was not consistent with those obtained by us. This difference was probably due to the use of T. molitor pupae instead of larvae, indicating that T. molitor pupae can be more nutritious than the larvae. According to the author, P. nigrispinus can be reared with T. molitor with the aim of increasing the populations of this predator in the field for biolog- ical control of A. gemmatalis. In terms of cost of production, one way to reduce material costs is to switch production to a larger scale, since rearing insects in large quantities can optimize the system, leading to economies of scale (i.e., reducing the cost per unit produced). M. domestica had the lowest cost (Table 2) because this insect is less demanding in terms of the production process; fewer devices are required to maintain the climatic conditions of the laboratory since this prey can be reared at ambient temperature and produces a large number of individuals per cycle. Mendes et al. (2005) calculated the cost of producing Orius insidiosus (Say) (Hemiptera: Anthocoridae) and observed that labor 132 S.A. De Bortoli et al. / Biological Control 58 (2011) 127–132 costs accounted for 62% of the total production cost. According to Parra (2002), in Brazil, labor costs represent 60–80% of the total cost of producing this predator in the laboratory. Therefore, rearing predators on D. saccharalis can lead to greater production at the same initial population and time interval. How- ever, economically, larvae of M. domestica are more viable because the final cost of rearing this prey is 10 times lower than that for D. saccharalis. The estimated cost of producing P. nigrispinus was low- er than that for the pirate bug O. insidiosus, as reported by Mendes et al. (2005). However, one should bear in mind that we did not compute the depreciation of buildings and equipment, which would increase the cost of production. In summary, producing the predator P. nigrispinus was almost twelve times as costly when they were fed D. saccharalis rather than M. domestica. Although D. saccharalis larvae are the best prey for the optimal development of P. nigrispinus. M. domestica larvae are the prey that gives the lowest cost of production of the preda- tor P. nigrispinus. 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