BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Effect of Diets on Biology of Abaris basistriata and Selenophorus seriatoporus (Coleoptera: Carabidae) Author(s): C. L. Barbosa, F. J. Cividanes, D. J. Andrade, and T. M. Dos Santos- Cividanes Source: Annals of the Entomological Society of America, 105(1):54-59. 2012. Published By: Entomological Society of America DOI: http://dx.doi.org/10.1603/AN11039 URL: http://www.bioone.org/doi/full/10.1603/AN11039 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. http://dx.doi.org/10.1603/AN11039 http://www.bioone.org/doi/full/10.1603/AN11039 http://www.bioone.org http://www.bioone.org/page/terms_of_use - ARTHROPOD BIOLOGY Effect of Diets on Biology of Abaris basistriata and Selenophorus seriatoporus (Coleoptera: Carabidae) C. L. BARBOSA,1,2 F. J. CIVIDANES,1 D. J. ANDRADE,1 AND T. M. DOS SANTOS-CIVIDANES3 Ann. Entomol. Soc. Am. 105(1): 54Ð59 (2012); DOI: http://dx.doi.org/10.1603/AN11039 ABSTRACT Ground beetles or carabids are collective terms for the beetle family Carabidae. This family contains many species considered important predators associated with agricultural crops. The current study aimed to evaluate the effect of different diet types on consumption, fecundity, and egg viability of Abaris basistriata Chaudoir and Selenophorus seriatoporus Putzeys (Coleoptera: Carabi- dae). The diets assessed were as follows: larvae of Tenebrio molitor L.; minced beef; dry cat food; the greenbug,Schizaphis graminum(Rondani); seeds of signal grass,BrachiariadecumbensStapf; and a diet mixture. Five males and Þve females of each species were kept isolated in a plastic container divided by a silicon barrier, one side being Þlled with sifted soil that was moistened for oviposition and the other lined with Þlter paper to receive the diet.A. basistriata did not consume theB. decumbens seeds. The most consumed diet by A. basistriata and S. seriatoporus adults was T.molitor larvae. S. graminum and T.molitor larvae and diet mixture were considered the diets most favorable for the reproductive capacity of A. basistriata and S. seriatoporus, respectively. However, T.molitor larvae and diet mixture were the most favorable diets for rearing both carabid species in the laboratory. KEY WORDS ground beetle, predator, consumption, reproduction Carabids are cited as being predators of aphids, lepi- dopteran larvae, slugs, and herbaceous plant seeds (Kromp 1999, Holland and Luff 2000, Tooley and Brust 2002). Many species of these beetles have a role in the natural biological pest control for several crops, among which are prominent Alabama argillacea (Hübner) (Lepidoptera: Noctuidae) on cotton, Gos- sypium hirsutum L. (Allen 1977, Chocorosqui and Pasini 2000);Anticarsia gemmatalisHübner (Lepidop- tera: Noctuidae) on soybean, Glycine max (L.) Merr. (Fuller 1988);Diatraea saccharalis (F.) (Lepidoptera: Crambidae) on sugarcane (Saccharum spp.) and sor- ghum (Sorghum spp.) (Fuller and Reagan 1988); Plu- tella xylostella (L.) (Lepidoptera: Plutellidae) on cab- bage, Brassica oleracea L. (Suenaga and Hamamura 2001); and Spodoptera frugiperda (J.E. Smith) (Lep- idoptera: Noctuidae) on corn,ZeamaysL. (Wyckhuys and OÕNeil 2006). Adults and larvae of carabids may have diverse feed- ing habits, varying from carnivorous to granivorous, with the granivore species considered to have evolved from carnivorous ancestors (Hurka and Jarosik 2003, Sasakawa 2010). However, little is known of the effect of diet on the fecundity of these beetles. In general, a mixed diet stimulates egg production (Wallin et al. 1992, Bilde and Toft 1994), but fecundity also is af- fected by the quality of monospeciÞc diets (Bilde and Toft 1994, Bilde et al. 2000). Wallin et al. (1992) ob- served insect diet adequate for the reproduction in some polyphagous predatory carabid beetles. How- ever, the females of granivorous species generally have a higher oviposition rate when they are fed on a diet of seeds in comparison with an insect diet (Jor- gensen and Toft 1997a,b; Saska 2008). Despite the importance of this group of predatory insects, most information on carabids in the Brazilian agroecosystems pertains to population ßuctuation and faunistic analysis (Pegoraro and Foerster 1988, Pinto et al. 2000, Freitas et al. 2002, Merlim et al. 2006). Abaris basistriata Chaudoir, Selenophorus seriatoporus Putzeys, Tetracha brasiliensis (Kirby), Odontochila nodicornis (Dejean), and Calosoma granulatum Perty are indicated as predominant species in the northeast region of the state of São Paulo (Cividanes et al. 2009). This study aimed to evaluate the effect of different diet types on the consumption, fecundity, and egg viability of adults of the carabids A. basistriata and S. seriatoporus. Materials and Methods The study was conducted from January to September 2010 in the Departamento de Fitossanidade and in an experimental area of the Fazenda de Ensino, Pesquisa e Produção at the Faculdade de Ciências Agrárias e 1 Universidade Estadual Paulista, Faculdade de Ciências Agrárias e Veterinárias, Departamento de Fitossanidade, Laboratório de Ecolo- gia de Insetos, Via de Acesso Prof. Paulo Donato Castellane s/no, CEP 14884-900, Jaboticabal, SP, Brazil. 2 Corresponding author, e-mail: crislanyunesp@hotmail.com. 3 Agência Paulista de Tecnologia dos Agronegócios, Avenida Ban- deirantes, no 2.419, CEP 14030-670 Ribeirão Preto, SP, Brazil. 0013-8746/12/0054Ð0059$04.00/0 � 2012 Entomological Society of America VeterináriasÐUniversidade Estadual Paulista (FCAV/ UNESP), Jaboticabal Campus, São Paulo state, Brazil. Live adults of the carabids A. basistriata and S. seriatoporus were captured using trench-type traps (Clark et al. 1994) placed in soybean and corn crops. Each trap was made of galvanized guttering sheet 90 cm in length and 10 cm in width, folded in half along its length in a V shape, which was installed with the edge kept at soil level. The extremities of the sheet were kept in contact with the edges of the empty plastic cups 8 cm in diameter and 14 cm in height. The traps were checked daily for the collection of adults. The diets assessed were as follows: 1)Tenebriomoli- tor L. larvae (Jorgensen and Toft 1997b); 2) minced beef (Sasakawa 2007); 3) Whiskas dry cat food (Wal- lin et al. 1992); 4) the greenbug, Schizaphis graminum (Rondani) (Bilde and Toft 1994, 1997); 5) seeds of signal grass, Brachiaria decumbens Stapf (Jorgensen and Toft 1997b); and 6) a mixture of diets 1Ð5 (Jor- gensen and Toft 1997a). T. molitor larvae and S. graminum were obtained in accordance with the method cited by Santos and Boiça (2002) and Santos et al. (2009), respectively. The seeds of B. decumbens were collected in an experi- mental area at FCAV/UNESP, according to the Jor- gensen and Toft (1997a,b) method. The assessment of the diets was carried out using gerbox-type plastic containers measuring 11 cm in length by 11 cm in width by 3.5 cm in depth. Each container was divided in two by a silicon barrier, with one part being Þlled with sterilized, sifted, and moist- ened soil and the other part lined with Þlter paper. The soil acted as shelter for carabid oviposition, and the Þlter paper enabled the food and water was provided by moistened cotton wool (Jorgensen and Toft 1997b). To maintain the humidity inside the container, the soil and cotton wool were moistened every 2 d. For the entirety of the experiments, the containers were kept in a room maintained at 26 � 2�C, 70 � 10% RH, and a photoperiod of 12:12 (L:D) h. To standardize the hunger level among the studied carabids, all food was withheld four days before the experiment, with only water being offered. Subse- quently, Þve male and Þve female carabids were sep- arated in each container, the sex of species was determined by examining the shape of the protarsi, with the aid of a magnifying lens. The protarsi of male are dilated and pronounced than in the females, ac- cording to the method cited by Riddick and Mills (1994). After the start of the experiments, the diets offered were replaced with a new diet every 3 d. The diet not consumed by the insects was weighed using a Bel Engineering (0.001-g) precision scale. It should be noted that these carabids also were considered in the analysis of the effect of different diets on survival. The quantity used of T. molitor larvae, mincemeat and dry cat food was 0.2 g, whereas that of S. graminum was of 0.01 g and of B. decumbens seeds, 0.09 g. Before being given to the carabids, T. molitor larvae were cut into small pieces by using a scalpel, and S. graminum were died by freezing. It should be stressed that the quantity of the diets offered to the carabids was as- sessed in preliminary tests ad libitum. Consumption was assessed for each date of the diet replacement. Before being placed in the containers, the diets were weighed and the nonconsumed por- tions were dried in an oven at 75�C for 3 d and sub- sequently weighed with a precision scale. Means con- sumption was estimated using the following fresh (x), dry (y) weight regression model (y � a � bx), ob- tained considering the weight of 20 units of the diet without undergoing the drying process (x) and the corresponding dry weight (y), obtained from the aforementioned drying procedure (Riddick and Mills 1994). The linear regression equations obtained were T.molitor (y � Ð0.002 � 0.4786x; r2 � 0.99), meat (y � 0.0026 � 0.2898x; r2 � 0.97), dry cat food (y � 0.0002 � 0.9302x; r2 � 0.99), S. graminum (y � 0.0003 � 0.5203x; r2 � 0.97), and B. decumbens (y � Ð 0.0055 � 0.95x; r2 � 0.99). The consumption was evaluated during the entire time all the specimens were alive. Adults used in the evaluations of diets were the same individuals used in the formation of pairs for copulation and to obtain eggs, maintained on the cor- responding diets provided in the containers. These pairs were kept together for 12 h, three times a week. Subsequently, the containers with females were checked weekly, the soil was sifted to obtain eggs that were counted under a stereoscopic microscope trans- ferred to petri dishes containing moistened soil and kept in a climatized chamber at 26 � 2�C and a pho- toperiod of 12:12 (L:D) h (Jorgensen and Toft 1997b). The number of eggs observed in the containers was expressed as the mean egg production per female during a 12-wk oviposition period. The entirely randomized statistical design was used on a factorial scheme 5 � 2, with factor A (Þve diet types), factor B (male or female), and Þve repetitions per treatment. To analyze fecundity, the number of hatched larvae and egg viability was measured, with each species of carabid being assessed in six treat- ments (six diet types), with Þve repetitions. Before being analyzed statistically, the fecundity data and the number of larvae were transformed into ln(x � 5), and the egg fecundity data were trans- formed into arcsine √P/100. The data obtained were submitted to analysis of variance (ANOVA) using the F-test, and the means were compared with the Tukey test (P � 0. 05). Results and Discussion Considering separately the factors diet types and carabid sex, signiÞcant variation was found in the daily consumption of diets by A. basistriata and S. seriato- porus (Table 1). In the same way, the interaction between the factors diet type and sex of A. basistriata and S. seriatoporus was signiÞcant, indicating that the daily consumption of the diets varied as a function of the diet type and sex. In general, the diet consumed most on a daily basis by A. basistriata and S. seriatoporus adults was that consisting of T. molitor larvae, followed by beef, dry January 2012 BARBOSA ET AL.: EFFECT OF DIETS ON BIOLOGY OF CARABID BEETLES 55 cat food, and S. graminum (Table 1). The mean daily quantity of T. molitor larvae consumed by S. seriato- porus was higher than the quantity consumed by A. basistriata. However, despite the adults of S. seriato- porus being larger (10.4 mm) than those of A. basis- triata (6.2 mm), the consumption of dry cat food and of S. graminum by that species was less than the con- sumption observed for A. basistriata. It should be stressed that there was no consumption of B. decum- bens seeds byA.basistriata,whereas for S. seriatoporus, this was the least consumed diet. Some studies also assessed the feeding preferences of carabid species. Johnson and Cameron (1969) observed that Pterosti- chus melanarius Illiger and Amara apricaria Paykull preferred to feed on larvae and pupae of a curculionid (Hyperodes sp.) instead of seeds, whereas Jorgensen and Toft (1997a) observed that seeds of the weed Taraxacum sp. were the most preferred by adults of Harpalus rufipesDegeer, followed by diets aphidMeto- polophium dirhodumWalker and Drosophila melano- gasterMeigen. Fawki and Toft (2005) studied the food preference of Amara similataGyllenhal and observed higher consumption mixed seedÐinsect diet. There was a signiÞcant difference by sex in the consumption of diets by A. basistriata and S. seriato- porus (Table 1). Mean daily consumption by the fe- males was higher than that of males. Males and females of A. basistriata consumed similar daily quantities of beef, whereas for the diets of T. molitor larvae, dry cat food, and S. graminum, the females consumed signif- icantly more than the males (Table 1). For S. seriato- porus, there was no signiÞcant difference for the mean daily consumption of dry cat food by males and fe- males, but for the other diets, the females consumed signiÞcantly more than the males. The higher con- sumption by the females was probably due to the need to produce eggs, which uses considerable energy (OÕNeil and Wiedenmann 1990). The females of A. basistriata showed higher fecun- dity when were fed on T. molitor larvae and lower when fed on beef and dry cat food (Table 2). The fecundity of this carabid fed on S. graminum and diet mixtures did not differ signiÞcantly from the other diets. Females of this species did not feed, nor oviposit when kept on seeds of B. decumbens.The diets of beef and dry cat food enabled A. basistriata to produce an intermediate quantity of eggs in relation to the other diets; however, they did not differ signiÞcantly from the diets of S. graminum and diet mixtures. The fe- males of S. seriatoporus fed on beef, S. graminum and seeds ofB. decumbens did not produce eggs (Table 3), whereas the diet mixtures, dry cat food, and T. molitor larvae enabled S. seriatoporus to produce a higher quantity of eggs. Saska (2008) also found that a diet of Table 1. Consumption of different diets (milligrams per day, mean � SE) by males and females of A. basistriata and S. seriatoporus Diet Consumption (mg/d) A. basistriata S. seriatoporus � � � � T. molitor larvae (n � 5) 9.8 � 0.29aB 12.0 � 0.63aA 13.2 � 0.13aB 17.0 � 0.09aA Beef (n � 5) 7.3 � 0.07bA 7.7 � 0.11bA 7.0 � 0.19bB 10.6 � 0.13bA Dry cat food (n � 5) 4.5 � 0.11cB 5.6 � 0.08cA 3.7 � 0.25cA 3.6 � 0.14cA S. graminum (n � 5) 2.3 � 0.07dB 3.3 � 0.24dA 1.6 � 0.03dB 2.1 � 0.01dA B. decumbens seeds (n � 5) 0.0 � 0.00eA 0.0 � 0.00eA 0.2 � 0.12eB 2.0 � 0.15dA Diets (A) F-test 607.5** 3,440.0** Sex � and � (B) F-test 39.0** 466.8** (A � B) F-test 6.0** 75.6** CV % 10.3** 5.2** Values followed by same letter, lowercase in columns and uppercase in rows, are not different by Tukey test (P � 0.05). **, P � 0.01. Table 2. Mean fecundity, number of hatched larvae, and egg viability of A. basistriata females fed on different diets Diet Fecundity (mean � SE)a,b No. hatched larvae (mean � SE)a,c Egg viability (%, mean � SE)a,d T. molitor larvae (n � 5) 9.0 � 0.09a 5.0 � 0.07a 47.2 � 4.73a Beef (n � 5) 3.8 � 0.09b 1.2 � 0.07c 34.0 � 4.73a Dry cat food (n � 5) 3.9 � 0.09b 1.8 � 0.07bc 51.5 � 4.73a S. graminum (n � 5) 7.1 � 0.09ab 4.5 � 0.07a 51.8 � 4.73a B. decumbens seeds (n � 5) 0.0 � 0.00c 0.0 � 0.00c 0.0 � 0.00b Mixture with all Þve diets (n � 5) 7.9 � 0.09ab 4.1 � 0.07ab 46.0 � 4.73a F-test 18.0** 14.8** 17.7** CV (%) 8.8** 8.0** 27.6** Values followed by the same letter in the column do not differ signiÞcantly with the Tukey test (P � 0.05). **, P � 0.01. aOriginal fecundity data and number of hatched larvae were transformed into ln(x � 5). Egg viability data were transformed into arcsine P/100�. bMean number of eggs per female during a 12-wk oviposition period. cMean number of hatched larvae during a 12-wk period. dMean number of egg viability in percentage during a 12-wk period. 56 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 105, no. 1 T.molitorwas adequate for the reproduction ofAmara aenea Degeer. Similar results observed for Amara (Curtonotus) macronota (Solsky) and Anisodactylus punctatipennis Morawitz fed on T. molitor larvae fa- voring the production of eggs (Sasakawa 2009a,b). Wallin et al. (1992) also found that the diet with cat food was adequate for the reproduction of Bembidion lampros Herbst, Pterostichus cupreus Linnaeus and P. melanarius.There are little information is available on nonproduction of eggs by carabids fed on different diets. However, it should be noted that nonproduction of eggs by S. seriatoporus fed on seeds of B. decumbens is consistent with studies of Hurka and Jarosik (2003) and Saska (2008). Hurka and Jarosik (2003) observed that A. aenea not produce eggs when fed on seeds of Potentilla argentela,whereas Saska (2008) reported non- production of eggs by Amara familiaris (Duftschmid) and A. similata when fed on seeds of Capsella bursa- pastoris (L.) Medik. and Stellaria media (L.) Vill., re- spectively. In the current study, larval emergence rate hatched was greater when the species fed on diets that enabled higher egg production. Thus, T. molitor larvae and S. graminum allowed A. basistriata to produce more lar- vae, which occurred with S. seriatoporuswhen it fed of diet mixtures and T. molitor larvae (Tables 2 and 3). The egg viability of A. basistriata ranged from 34.0 to 51.8%, without a signiÞcant difference between the number of eggs and the diet types (Table 2). The greatest egg viability 52% was obtained for females fed S. graminum. With the exception of B. decumbens seeds, the test diets enabled A. basistriata to produce eggs in similar quantities. S. seriatoporus, the diets of dry cat food, T. molitor larvae and diet mixtures en- abled the carabid to produce eggs at viability of 32.3, 45.2, and 51.3%, respectively (Table 3). Mols (1988) reported that the quantity and quality of the diet directly inßuences carabid oviposition and egg viabil- ity, with reabsorption of formed eggs by the insect also being possible, should food be inadequate or scarce. When A. basistriata fed on diet mixtures, T. molitor larvae, beef and dry cat food �50% of the carabids survived up to 150, 135, 120, and 90 d, respectively (Fig. 1). The diet mixtures and T. molitor larvae also allowed individuals to survive until 225 d. Approxi- mately 50% of S. seriatoporus fed on T. molitor larvae and diet mixtures survive up to 240 and 210 d, respec- tively, whereas 50% of S. seriatoporus fed on beef and dry cat food only survive up to 90 and 45 d, respec- tively (Fig. 2). Therefore, the effects of T. molitor larvae and diet mixtures on survival are different from those of beef and dry cat food, although the last in- dividual of four groups all died at 255 d. Besides pro- viding the longest period of adult survival of A. basis- triata and S. seriatoporus (Figs. 1 and 2), the diet of T. molitor larvae and diet mixtures also allowed these carabid species reach the highest fecundity and egg viability (Tables 2 and 3). Therefore, T. molitor larvae and diet mixtures were the most favorable diets for rearing both carabid species in the laboratory. Acknowledgments WethankCoordenaçãodeAperfeiçoamentodePessoalde Nṍvel Superior (CAPES) and the Conselho Nacional de Desenvolvimento CientṍÞco e Tecnológico (CNPq) for pro- viding a scholarships to C.L.B. and F.J.C, respectively. References Cited Allen,R.T. 1977. Calosoma(Castrida)alternans granulatum Perty: a predator of cotton leaf worms in Bolivia (Co- leoptera: Carabidae: Carabini). Coleopt. Bull. 31: 73Ð76. Bilde, T., J. A. Axelsen, and S. Toft. 2000. The value of Collembola from agricultural soils as food for a generalist predator. J. Appl. Ecol. 37: 672Ð683. Bilde, T., and S. Toft. 1997. Consumption by carabid beetles of three cereal aphid species relative to other prey types. Entomophaga 42: 21Ð32. Bilde, T., and S. Toft. 1994. Prey preference and egg pro- duction of the carabid beetle Agonum dorsale. Entomol. Exp. Appl. 73: 151Ð156. Chocorosqui, V. R., and A. Pasini. 2000. Predação de pupas de Alabama argillacea (Hübner) (Lepidoptera: Noctu- idae) por larvas e adultos de Calosoma granulatum Perty (Coleoptera: Carabidae) em Laboratório. An. Soc. En- tomol. Bras. 29: 65Ð70. Table 3. Mean fecundity, number of hatched larvae, and egg viability of S. seriatoporus females fed on the different diets Diet Fecundity (mean � SE)a,b No. hatched larvae (mean � SE)a,c Egg viability (%, mean � SE)a,d T. molitor larvae (n � 5) 7.6 � 0.07a 4.1 � 0.05b 45.2 � 1.61a Beef (n � 5) 0.0 � 0.00b 0.0 � 0.00c 0.0 � 0.00c Dry cat food (n � 5) 8.3 � 0.07a 2.5 � 0.05b 32.3 � 1.61b S. graminum (n � 5) 0.0 � 0.00b 0.0 � 0.00c 0.0 � 0.00c B. decumbens seeds (n � 5) 0.0 � 0.00b 0.0 � 0.00c 0.0 � 0.00c Mixture of all Þve diets (n � 5) 11.6 � 0.07a 7.0 � 0.05a 51.3 � 1.61a F-test 61.2** 57.0** 227.5** CV (%) 7.7** 5.8** 16.8** Values followed by the same letter in the column do not differ signiÞcantly with the Tukey test (P � 0.05). **, P � 0.01. aOriginal fecundity data and number of hatched larvae were transformed into ln(x � 5). Egg viability data were transformed into arcsine P/100�. bMean number of eggs per female during a 12-wk oviposition period. cMean number of hatched larvae during a 12-wk period. dMean number of egg viability in percentage during a 12-wk period. January 2012 BARBOSA ET AL.: EFFECT OF DIETS ON BIOLOGY OF CARABID BEETLES 57 Cividanes, F. J., J. C. Barbosa, S. Ide,N.W.Perioto, andR.I.R. Lara. 2009. Faunistic analysis of Carabidae and Staph- ylinidae (Coleoptera) in Þve agroecosystems in north- eastern São Paulo state, Brazil. Pesqu. Agropecu. Bras. 44: 954Ð958. Clark, M. S., J. M. Luna, N. D. Stone, and R. R. Youngman. 1994. Generalist predator consumption of armyworm (Lepidoptera: Noctuidae) and effect of predator removal on damage in no-till corn. Environ. Entomol. 23: 617Ð622. Fawki, S., and S. Toft. 2005. Food preferences and the value of animal food for the carabid beetle Amara similata (Gyll.) (Col., Carabidae). J. Appl. Entomol. 129: 551Ð556. Freitas, F. A., T. V. Zanuncio, M. C. Lacerda, and J. C. Zanuncio. 2002. Fauna de Coleoptera Coletada com Ar- madilhas Luminosas em Plantio deEucalyptus grandis em Santa Bárbara, Minas Gerais. Rev. Árvore. 26: 505Ð511. Fuller, B. W. 1988. Predation by Calleida decora (F.) (Co- leoptera: Carabidae) on velvetbean caterpillar (Lepidop- tera: Noctuidae) in soybean. J. Econ. Entomol. 81: 127Ð 129. Fuller, B. W., and T. E. Reagan. 1988. Comparative preda- tion of the sugarcane borer (Lepidoptera: Pyralidae) on sweet sorghum and sugarcane. J. Econ. Entomol. 81: 713Ð 717. Holland, J. M., and M. L. Luff. 2000. The effects of agricul- tural practices on Carabidae in temperate agroecosys- tems. Integr. Pest Manag. Rev. 5: 109Ð129. Fig. 1. Survival for adults of A. basistriata fed on different diets. Fig. 2. Survival for adults of S. seriatoporus fed on different diets. 58 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 105, no. 1 Hurka, K., and V. Jarosik. 2003. Larval omnivory in Amara aenea (Coleoptera: Carabidae). Eur. J. Entomol. 100: 329Ð335. Johnson, N. E., and R. S. Cameron. 1969. Phytophagous ground beetles. Ann. Entomol. Soc. Am. 62: 909Ð914. Jorgensen, H. B., and S. Toft. 1997a. Food preference, diet dependent fecundity and larval development inHarpalus rufines (Coleoptera: Carabidae). Pedobiologia 41: 301Ð 315. Jorgensen, H. B., and S. Toft. 1997b. Role of granivory and insectivory in the life cycle of the carabid beetle Amara similata. Ecol. Entomol. 22: 7Ð15. Kromp, B. 1999. Carabid beetles in sustainable agriculture: a review on pest control efÞcacy, cultivation impacts and enhancement. Agric. Ecosyst. Environ. 74: 187Ð228. Merlim, A. O., A. M. Aquino, and E.J.B.N. Cardoso. 2006. Larvas de Coleoptera em ecossistemas de araucária no Parque Estadual de Campos do Jordão, SP. Cienc. Rural 36: 1303Ð1306. Mols, P.J.M. 1988. Simulation of hunger, feeding and egg production in the carabid beetle Pterostichus coerulescens L. (Poecilus versicolor Sturm). Agric. Univ. Wag. Pap. 88: 1Ð99. O’Neil, R. J., and R. N. Wiedenmann. 1990. Body weight of Podisus maculiventris (Say) under various feeding re- gimes. Can. Entomol. 120: 285Ð294. Pegoraro, R. A., and L. A. Foerster. 1988. Abundância e distribuição de larvas e adultos de Calosoma granulatum Perty, 1830 (Coleoptera: Carabidae) dentre cultivares de soja em diferentes épocas de semeadura. Ann. Soc. En- tomol. Bras. 17: 237Ð248. Pinto, R., J. S. Zanuncio Junior, J.A.M. Ferreira, and J. C. Zanuncio. 2000. Flutuação populacional de Coleóptera em plantio de Eucalyptus urophylla no municṍpio de Três Marias, Minas Gerais. Floresta Ambiente 7: 143Ð151. Riddick, E.W., and N. J. Mills. 1994. Potential of adult cara- bids (Coleoptera: Carabidae) as predators of Þfth-instar codling moth (Lepidoptera: Tortricidae) in apple or- chards in California. Environ. Entomol. 23: 1338Ð1345. Santos, N.R.P., T. M. Santos-Cividanes., F. J. Cividanes., A.C.R. Anjos., and L.V.L. Oliveira. 2009. Aspectos bi- ológicos de Harmonia axyridis alimentada com duas es- pécies de presas e predação intraguilda com Eriopis con- nexa. Pesqu. Agropecu. Bras. 44: 554Ð560. Santos, T. M., and A. L. Boiça, Jr. 2002. Biological aspects and predatory capacity of Podisus nigrispinus (Dallas) (Hemiptera: Pentatomidae) fed on Alabama argillacea (Hübner) (Lepidoptera: Noctuidae) reared on cotton genotypes. Sci. Agric. 59: 671Ð675. Saska, P. 2008. Effect of diet on the fecundity of three cara- bid beetles. Psychol. Entomol. 33: 188Ð192. Sasakawa,K. 2007. Laboratory studies on larval feeding hab- its ofAmaramacronota (Coleoptera: Carabidae: Zabrini). Appl. Entomol. Zool. 42: 669Ð674. Sasakawa, K. 2009a. Diet affects male gonad maturation, fe- male fecundity, and larval development in the granivo- rous ground beetle Anisodactylus punctatipennis. Ecol. Entomol. 34: 406Ð411. Sasakawa, K. 2009b. Effects of maternal diet on fecundity and larval development in the ÔprimitiveÕ granivorous carabid Amara (Curtonotus) macronota. Entomol. Exp. Appl. 130: 106Ð112. Sasakawa, K. 2010. Field observations of climbing behavior and seed predation by adult ground beetles (Coleoptera: Carabidae) in a lowland area of the temperate zone. Environ. Entomol. 39: 1554Ð1560. Suenaga, H., and T. Hamamura. 2001. Occurrence of cara- bid beetles (Coleoptera: Carabidae) in cabbage Þelds and their possible impact on lepidopteran pests. Appl. Ento- mol. Zool. 36: 151Ð160. Tooley, J., andG.E.Brust. 2002. Weed predation by carabid beetles, pp. 215Ð229. In J. M. Holland (ed.), The agro- ecology of carabid beetles. Intercept, Andover, United Kingdom. Wallin, H., P. A. Chiverton, B. S. Ekbom, and A. Borg. 1992. The diet, fecundity and egg size in some polyphagous predatory carabid beetles. Entomol. Exp. Appl. 65: 129Ð 140. Wyckhuys, K.A.G., and R. J. O’Neil. 2006. Population dy- namics of Spodoptera frugiperda Smith (Lepidoptera: Noctuidae) and associated arthropod natural enemies in Honduran subsistence maize. Crop Prot. 25: 1180Ð1190. Received 21 February 2011; accepted 31 May 2011. January 2012 BARBOSA ET AL.: EFFECT OF DIETS ON BIOLOGY OF CARABID BEETLES 59