Essential versus alternative foods of insect predators: benefits of a mixed diet

Although many predatory insects appear to be opportunistic generalists in their selection of prey, only a subset of prey species may in fact serve as “essential foods” capable of supporting immature growth and adult reproduction. It has been suggested that other, “alternative foods” serve only to maintain the predator when essential foods are not available, but little research has evaluated the significance of a mixed diet of essential and alternative foods for predator growth or reproduction. Here we test the general hypothesis that although alternative prey may be inadequate to support reproduction when consumed alone by adult predators, consumption of such prey may enhance the predator's reproductive output when the predator also has access to essential prey. We compared egg production by two aphidophagous lady beetles, Coccinella septempunctata and C. transversoguttata, provided with diets of aphids (essential prey) and weevils (alternative prey). As predicted, female predators produced greater numbers of eggs when a diet of pea aphids in limited number was supplemented by alfalfa weevil larvae. The predators laid no eggs when provided only with weevils or only with sugar. But once aphids were added to the diet, females of C. transversoguttata (but not C. septempunctata) laid eggs in greater numbers when they had fed previously on weevils than on sugar. Females of both species also produced eggs in modest numbers when provided with both weevils in excess and sugar, but this diet supported a lower rate of egg production than did a diet of weevils in excess plus a limited number of aphids. Although C. septempunctata has a longer history of association with the alfalfa weevil than does C. transversoguttata, the former species was not more effective in exploiting this alternative prey in support of reproduction. The tendency of generalist predators such as adult lady beetles to consume alternative as well as essential prey probably enhances considerably their ability to capitalize on short-lived and scattered opportunities as they seek out suitable sites in which to reproduce. Received: 11 May 1998 / Accepted: 1 May 1999     

An artificial diet for continuous maintenance of Coccinella septempunctata adults (Coleoptera: Coccinellidae)

An artificial diet was developed for continuous maintenance of ladybird beetle (Coccinella septempunctata) adults. This beetle is an important biocontrol agent for greenhouse crops. Several biological parameters were measured and compared between colonies fed the artificial diet over eight successive generations, and colonies fed aphids. Weight, survival rates from 1st to 4th instar, pupation rate, and emergence rate were similar for both colonies, but the sex ratio (♀:♂) of the artificial diet-fed colony was higher than that of the aphid-fed colony. These results suggest that this artificial diet could meet the basic needs for the successive rearing of C. septempunctata adults. The prepared diet, which is semi-solid, provides suitable moisture for C. septempunctata, and the diet ingredients are inexpensive and relatively easy to prepare. This diet and rearing technique offers a very useful approach for mass production of C. septempunctata.     

Effect of diet and mating status on ovarian development and oviposition in the polyphagous predator Macrolophus caliginosus (Heteroptera: Miridae)

The mirid bug Macrolophus caliginosus is commercially reared on eggs of Ephestia kuehniella, constituting an effective but expensive factitious food. Artificial diets can decrease the rearing costs of this natural enemy, but developing and evaluating an artificial diet is a very time-consuming activity. In the current study, development and reproduction of M. caliginosus on two artificial diets based on egg yolk were investigated. The artificial diets resulted in longer development and lower adult weights, but survival was comparable with that of control insects fed E. kuehniella eggs. Reproductive potential of the predator reared on factitious and artificial foods was assessed using a dissection method. The influence of nymphal food on fecundity was less important than that of adult food. Adults fed E. kuehniella eggs had a preoviposition period of about 4 days, whereas adults offered only plant material started laying eggs about 7 days after emergence. Ovarian scores at day 7 were higher for females fed E. kuehniella eggs than for those given access only to a tobacco leaf. Ovarian scores were not significantly affected by mating status. In a final test, a parallel comparison of two methods for assessing reproductive response to diet was made. Here, adult couples were offered one of four diets: E. kuehniella eggs, one of two artificial diets or no food. Half of the females were dissected and the other half was held for determining lifetime oviposition. Females fed E. kuehniella eggs had superior ovarian scores and laid more eggs than those fed either artificial diet or those given no extra food. A good correlation (r = 0.97) was obtained between ovarian scores and oviposition data, indicating that dissecting females after 1 week provides a reliable estimate of fecundity as affected by diet quality. Rapid reproductive assessments as used in the current study will help to increase the rate of development of artificial diets and may contribute to more cost effective production methods for augmentative biological control agents.     

七星瓢虫基础研究现状

七星瓢虫对蚜虫等农林业中的重要害虫有较强的控制作用,利用七星瓢虫进行控害的生物防治技术正在逐步推广.本文就七星瓢虫的生物学特性、捕食、对农药的反应以及人工饲养等方面进行综述:七星瓢虫幼虫随龄期增长逐渐出现黄斑,温湿度对生长发育影响较大,以成虫越冬,有滞育现象.七星瓢虫捕食性强,捕食对象较多,包括多种蚜虫、木虱、蚧虫、粉...     

Avoidance of intraguild predation leads to a long-term positive trait-mediated indirect effect in an insect community

Intraguild predation among natural enemies is common in food webs with insect herbivores at their base. Though intraguild predation may be reciprocal, typically one species suffers more than the other and frequently exhibits behavioural strategies to lessen these effects. How such short-term behaviours influence population dynamics over several generations has been little studied. We worked with a model insect community consisting of two species of aphid feeding on different host plants (Acyrthosiphon pisum on Vicia and Sitobion avenae on Triticum), a parasitoid (Aphidius ervi) that attacks both species, and a dominant intraguild predator (Coccinella septempunctata) that also feeds on both aphids (whether parasitized or not). As reported previously, we found A. ervi avoided chemical traces of C. septempunctata. In population cages in the laboratory, application of C. septempunctata extracts to Vicia plants reduced parasitism on A. pisum. This did not increase parasitism on the other aphid species, our predicted short-term trait-mediated effect. However, a longer term multigenerational consequence of intraguild predator avoidance was observed. In cages where extracts were applied in the first generation of the study, parasitoid recruitment was reduced leading to higher population densities of both aphid species. S. avenae thus benefits from the presence of a dominant intraguild predator foraging on another species of aphid (A. pisum) on a different food plant, a long-term, trait-mediated example of apparent mutualism. The mechanism underlying this effect is hypothesized to be the reduced searching efficiency of a shared parasitoid in the presence of cues associated with the dominant predator.     

Solid artificial diets for the phytoseiid predator Amblyseius swirskii

Amblyseius swirskii (Athias-Henriot) (Acari: Phytoseiidae) is a key predator of a wide range of pests including thrips, whitefly and several mite pests. A number of artificial diets have been developed for this predator, but all of these are liquid, complicating their use in mass production. In the present study, we investigated the survival, development and reproduction of A. swirskii fed on several dry artificial diets: the tested diets were freeze dried forms of previously developed liquid meridic artificial diets supplemented with extracts of decapsulated cysts of Artemia franciscana Kellogg (Anostraca: Artemiidae) or with pupal hemolymph of Chinese oak silkworm Antheraea pernyi (Guérin-Méneville) (Lepidoptera: Saturniidae), and newly composed powdered meridic artificial diets supplemented with ground dry A. franciscana cysts or lyophilized pupal hemolymph of A. pernyi. Performance of the mite on the artificial diets was compared with that on cattail pollen (Typha latifolia L.). Developmental time of A. swirskii females offered lyophilized diets was significantly shorter than on powdered diets. Total fecundity was significantly higher for females fed on the lyophilized diets than for those maintained on the powdered diet with A. franciscana. Daily oviposition rates were similar on T. latifolia pollen and both lyophilized diets but lower on both powdered diets. The highest intrinsic rate of increase was observed when A. swirskii was fed on T. latifolia pollen (0.210 females per female per day), followed by the freeze dried diets enriched with A. pernyi and A. franciscana (0.195 and 0.184 females per female per day, respectively), and the lowest growth rates were observed on the powdered diets supplemented with A. franciscana and A. pernyi (0.159 and 0.158 females per female per day, respectively). In conclusion, the phytoseiid was able to effectively feed on solid, powdered artificial diets. Freeze-drying of liquid diets did not influence their value to support the development and reproduction of A. swirskii. For mass rearing purposes, these dry diets have several advantages over liquid ones, including more convenient application and storage. Furthermore, they are believed to have better potential for use as supplemental foods to sustain predatory mite populations in the crop after release.     

Performance of Amblyseius herbicolus on broad mites and on castor bean and sunnhemp pollen

Amblyseius herbicolus (Banks) is found associated with broad mites Polyphagotarsonemus latus in crops such as chili pepper in Brazil. The species has a potential for controlling P. latus, but little is known about its development and reproduction on this pest as well as on other food sources. We studied biological, reproductive and life table parameters of A. herbicolus on three different diets: broad mites, castor bean pollen (Ricinus communis) and sunnhemp pollen (Crotalaria juncea). The predator was able to develop and reproduce on all diets. However, its intrinsic growth rate was higher on the diet of broad mites or on castor bean pollen than on sunnhemp pollen. Differences among pollen species may be due to their nutritional content. Feeding on alternative food such as pollen can facilitate the predator’s mass rearing and maintain its population on crops when prey is absent or scarce. Other strategies of using pollen to sustain predator population and reduce pest damage are discussed.     

Regulation of vitellogenin synthesis by juvenile hormone analogue in Coccinella septempunctata

Vitellogenesis in the lady beetle, Coccinella septempunctata, is controlled by juvenile hormone (JH). When immature females were reared on an artificial diet, they were characterized by hypertrophy of the fat bodies and slow development of the ovaries. Vitellogenin (Vg) synthesis in the fat body remained at a very low level throughout adult development. RNA synthesis also stayed at a relatively low level. Treatment with hydroprene induced vitellogenesis in these non-fecund females. Stimulation in Vg synthesis in hormone-treated females was demonstrated both in vivo and in fat body culture. Synthesis of fat body RNA also increased after hormone treatment. Fat body RNA from hormone-treated females directed the synthesis of Vg polypeptides both in Xenopus oocytes and in a reticulocyte lysate. Thus the induction of Vg synthesis by JH involves an increase in the level of translatable Vg mRNA.     

Why eat extrafloral nectar? Understanding food selection by Coleomegilla maculata (Coleoptera: Coccinellidae)

Methods of increasing predator abundance within a habitat include the incorporation of non-prey food items, yet the influence of this on predation intensity toward herbivores remains unknown. In order to gain an understanding of nectar feeding in the predaceous beetle, Coleomegilla maculata (DeGeer), laboratory studies were conducted evaluating prey consumption in the presence of extrafloral nectaries. The physiology of beetles with access to prey only and a mixed diet were compared. To elucidate results of beetle physiology, Y-tube olfactometer studies were conducted and preferences between food types evaluated. Coleomegilla maculata females consumed 9 % fewer aphids when nectar was available. Lipid and glycogen content, as well as oocyte volume were not increased upon consumption of a mixed diet. Evaluation of predator behavior when offered both food resources together and separately demonstrated that extrafloral nectaries are attractive.     

Development and reproduction of the predatory mite Amblyseius swirskii on artificial diets

Development, survival and reproduction of the predatory mite Amblyseius swirskii (Athias-Henriot) (Acari: Phytoseiidae) were assessed when fed on cattail pollen (Typha latifolia L.), dried fruit mite (Carpoglyphus lactis L.), or on two artificial diets. The basic artificial diet (AD1) was composed of honey, sucrose, tryptone, yeast extract, and egg yolk. This diet was enriched (AD2) by adding hemolymph from oak silkworm pupae (Antheraea pernyi (Guérin-Méneville)). Mites fed on C. lactis and AD2 had shorter immature and preoviposition periods than those fed on the other diets. The total number of deposited eggs was significantly higher for females fed on AD2 than for those maintained on the other diets. The intrinsic rate of increase (r_m) of A. swirskii was highest on AD2 and C. lactis, followed by T. latifolia pollen, and AD1. In conclusion, the artificial diet AD2 supported development and reproduction of A. swirskii to the same extent as a factitious prey which is routinely used in the mass rearing of the phytoseiid. Our findings indicate the potential of artificial diets for the mass production of this economically important predatory mite.     

Optimization an Optimal Artificial Diet for the Predatory Bug Orius sauteri (Hemiptera: Anthocoridae)

Background

The flower bug Orius sauteri is an important polyphagous predator that is widely used for the biological control of mites and aphids. However, the optimal conditions for mass rearing of this insect are still unclear, thus limiting its application.

Methodology

In this study, we investigated the optimal ingredients of an artificial diet for raising O. sauteri using a microencapsulation technique. The ingredients included egg yolk (vitellus), whole-pupa homogenate of the Tussah silk moth (Antheraea paphia), honey, sucrose, rapeseed (Brassica napus) pollen and sinkaline. We tested 25 combinations of the above ingredients using an orthogonal experimental design. Using statistical analysis, we confirmed the main effect factors amongst the components, and selected five optimal combinations based on different biological and physiological characters.

Principal Findings

The results showed that, although different artificial diet formats significantly influenced the development and reproductive ability of O. sauteri, the complete development of O. sauteri to sexual maturity could only be achieved by optimizing the artificial diet according to specific biological characters. In general, pupae of A. paphia had more influence on O sauteri development than did artificial components. The results of a follow-up test of locomotory and respiratory capacity indicated that respiratory quotient, metabolic rate and average creeping speed were all influenced by different diets. Furthermore, the field evaluations of mating preference, predatory consumption and population dispersion also demonstrated the benefits that could be provided by optimal artificial diets.

Conclusions

A microencapsulated artificial diet overcame many of the difficulties highlighted by previous studies on the mass rearing of O. sauteri. Optimization of the microencapsulated artificial diet directly increased the biological and physiological characters investigated. Successive physiological tests and field investigations were used to evaluate the outcome of different artificial diet combinations on the quality of the reared O. sauteri.     

Performance of the Wheat Armyworm, Pseudaletia sequax Franclemont, on Natural and Artificial Diets

Armyworms in the genus Pseudaletia Franclemont are important pests of cereal crops worldwide. The development of pest management strategies is largely dependent on studies involving the relationship among the pest, its host plants and natural enemies, as well as its susceptibility to chemical and biological control strategies. Part of these studies requires rearing the species of interest under laboratory conditions, which can be greatly facilitated by the use of artificial diets. In the present study, the effects of three artificial diets on development, survival, reproduction and demographic parameters of P. sequax were compared to a natural host plant, Pennisetum clandestinum. A diet based on pinto beans, wheat germ and torula yeast provided the best results in comparison to those obtained with the host plant. The most important life history parameters used in the evaluation of the nutritional quality of the diets were fecundity, egg viability and pupae deformities. The intrinsic rate of increase and the net reproductive rate were higher when individuals were fed on the artificial diet if compared to the host plant, indicating the suitability of the tested diet for the rearing of P. sequax. This artificial diet has been used successfully for the rearing of P. sequax without any fitness effect for over 18 consecutive generations.     

Responses of midgut amylases of Helicoverpa armigera to feeding on various host plants

Midgut digestive amylases and proteinases of Helicoverpa armigera, a polyphagous and devastating insect pest of economic importance have been studied. We also identified the potential of a sorghum amylase inhibitor against H. armigera midgut amylase. Amylase activities were detected in all the larval instars, pupae, moths and eggs; early instars had lower amylase levels which steadily increased up to the sixth larval instar. Qualitative and quantitative differences in midgut amylases of H. armigera upon feeding on natural and artificial diets were evident. Natural diets were categorized as one or more members of legumes, vegetables, flowers and cereals belonging to different plant families. Amylase activity and isoform patterns varied depending on host plant and/or artificial diet. Artificial diet-fed H. armigera larvae had comparatively high amylase activity and several unique amylase isoforms. Correlation of amylase and proteinase activities of H. armigera with the protein and carbohydrate content of various diets suggested that H. armigera regulates the levels of these digestive enzymes in response to macromolecular composition of the diet. These adjustments in the digestive enzymes of H. armigera may be to obtain better nourishment from the diet and avoid toxicity due to nutritional imbalance. H. armigera, a generalist feeder experiences a great degree of nutritional heterogeneity in its diet. An investigation of the differences in enzyme levels in response to macronutrient balance and imbalance highlight their importance in insect nutrition.     

Egg Predation by the Introduced Lady Beetle,Coccinella septempunctata (Coleoptera: Coccinellidae), Lowers Mortality but Raises Relative Risk for the Native Lady Beetle,Coccinella novemnotata

Populations of the native ninespotted lady beetle, Coccinella novemnotata Herbst, have undergone precipitous declines in North America following the establishment of the exotic sevenspotted lady beetle, Coccinella septempunctata L. Recent volunteer efforts have made it possible to establish colonies of the now-rare C. novemnotata and test mechanisms contributing to its decline. We evaluated the relative frequencies of intraguild predation and cannibalism of eggs between these two species. A single C. novemnotata or C. septempunctata adult was exposed to conspecific and heterospecific eggs in either the presence or absence of pea aphids. The study revealed two expected results: 1) eggs of C. novemnotata were consumed more frequently than eggs of C. septempunctata by both species, and 2) egg consumption was higher when aphids were absent, independent of predator and egg species. There were also two unexpected results from the study: 1) the asymmetry between egg predation rates was higher when aphids were present, and 2) higher predation rates on C. novemnotata eggs in the absence of alternate prey was due to a relatively higher rate of intraspecific cannibalism. This implies that C. novemnotata would have suffered higher egg mortality rates before the invasion of C. septempunctata, but even though the aggregate rate of egg predation on C. novemnotata eggs is lower post-invasion, it is still significantly higher than the aggregate rate of predation of C. septempunctata eggs. This differential pattern of asymmetric predation could contribute to habitat compression and the overall decline of C. novemnotata.     

Predaceous Behavior and Life History of Odontopharynx longicaudata (Diplogasterida)

The behavior of a California isolate of the predaceous nematode, Odontopharynx longicaudata de Man, was studied in water agar culture. When feeding on an Acrobeloides sp. the predator completed its life cycle in 13 to 14 days at 25 C. Optimum temperature for reproduction of the predator was 25 C, few individuals survived at 10 C, and 30 C was lethal. Males were necessary for reproduction, and at 25 C the sex ratio was about 1:1. All postembryonic stages were voracious feeders. A single female predator consumed 30 individuals of another Acrobeloides sp. in 1.5 days. Juveniles must feed in order to complete their development. Three modes of feeding were observed depending on the prey selected. A high degree of prey selectivity occurred; 6 of 17 nematode prey species were readily consumed by the predator, but there was little or no feeding on the remaining 11 species. Predation percentage varied with prey species. Consumption of Anguina pacificae and the two Acrobeloides spp. was almost 100%, consumption of A. amsinckiae, Pratylenchus vulnus, and Trichodorus sp. was 70-78%. Difference in final predator population densities was obtained after feeding on the two species of Acrobeloides. Final predator population densities increased linearly with increasing inoculum levels of the first Acrobeloides sp.     

Assessment of the functional response parameters of Coccinella septempunctata to varying densities of Acyrthosiphon pisum

The functional response of a predator describes the rate at which it kills its prey at different prey densities and is, therefore, an important tool to determine the efficiency of a predator in regulating the population dynamics of a prey. We investigated the functional response of Coccinella septempunctata L. (Coleoptera: Coccinellidae) to varying densities of Acyrthosiphon pisum (Harris) (Hemiptera: Aphididae). The results revealed that the linear coefficient, P₁ in the reduced logistic model was found significantly negative for the 4^th instar larvae and adult male and female of C. septempunctata (?0.002, ?0.004 and???0.002, respectively) indicating a type II functional response. The parameter estimates of adult male, female and 4^thinstar larvae of C. septempunctata calculated through Holling's disc model revealed that the highest attack rate (a) (1.047?±?0.014), shortest handling time (T_h) (0.0984?±?0.024?h) and largest maximum capture rate (T/T_h) (243.902) was exhibited by female. Parameter estimates calculated through Rogers’s equation also showed the same sequence where maximum attack rate (a) of 0.00256?±?0.0003 was exhibited by female followed by 4^th instar larvae (0.00251?±?0.0005). The shortest handling time (T_h) (0.210?±?0.003?h) and highest maximum capture rate (T/T_h) (114.17) was also exhibited by female. Comparison of functional response curves of adult male, female and 4^th instar larvae revealed that maximum consumption of prey at all the densities offered was shown by female followed by 4^thinstar larvae. The study manifested that C. septempunctata could be an efficient biocontrol agent of pea aphid, A. pisum.     

A new diet for reproduction of two predaceous mitesAmblyseius gossipi andAgistemus exsertus [Acari: Phytoseiidae,stigmaeidae]

Laboratory studies revealed that the acarid mite,Tyrophagus casei Oudemans is an adequate diet for development and reproduction of the stigmaeid mite,Agistemus exsertus Gonzalez. A combination of pollen grains of castor beanRicinus communis L. andT. casei is a sufficient plant-animal diet for the development and reproduction of the phytoseiid mite,Amblyseius gossipi Elbadry. When the later predator was restricted to such animal prey it developed to the adult stage, but females failed to lay eggs.      

Intraguild interactions between the entomopathogenic fungus Pandora neoaphidis and an aphid predator and parasitoid at the population scale

The interactions that occur between the entomopathogenic fungus Pandora neoaphidis and a predator (Coccinella septempunctata) and a parasitoid (Aphidius ervi) were assessed in microcosm and polytunnel experiments. Transmission of P. neoaphidis to the pea aphid, Acyrthosiphon pisum, was enhanced in the presence of both C. septempunctata and A. ervi in microcosm experiments done under fixed abiotic conditions. In contrast, the reproductive success of A. ervi was reduced in the presence of P. neoaphidis. Despite the increased fungal transmission in the presence of C. septempunctata, there was no additional decrease in the aphid population indicating that P. neoaphidis is functionally redundant in the presence of the coccinellid. In polytunnel experiments the reproductive success of A. ervi was not affected by P. neoaphidis. These results do not support those of the microcosm and may be due to the more natural abiotic conditions in the polytunnel reducing the competitive advantage of the fungus. Microcosms therefore provide an arena in which the interactions between fungal pathogens and other aphid-natural enemies can be assessed however, further assessments at increased spatial scales under more natural abiotic conditions are also required to accurately determine the outcome of these interactions.     

Effect of diet and mating status on ovarian development and oviposition in the polyphagous predator Macrolophus caliginosus (Heteroptera: Miridae)

The mirid bug Macrolophus caliginosus is commercially reared on eggs of Ephestia kuehniella, constituting an effective but expensive factitious food. Artificial diets can decrease the rearing costs of this natural enemy, but developing and evaluating an artificial diet is a very time-consuming activity. In the current study, development and reproduction of M. caliginosus on two artificial diets based on egg yolk were investigated. The artificial diets resulted in longer development and lower adult weights, but survival was comparable with that of control insects fed E. kuehniella eggs. Reproductive potential of the predator reared on factitious and artificial foods was assessed using a dissection method. The influence of nymphal food on fecundity was less important than that of adult food. Adults fed E. kuehniella eggs had a preoviposition period of about 4 days, whereas adults offered only plant material started laying eggs about 7 days after emergence. Ovarian scores at day 7 were higher for females fed E. kuehniella eggs than for those given access only to a tobacco leaf. Ovarian scores were not significantly affected by mating status. In a final test, a parallel comparison of two methods for assessing reproductive response to diet was made. Here, adult couples were offered one of four diets: E. kuehniella eggs, one of two artificial diets or no food. Half of the females were dissected and the other half was held for determining lifetime oviposition. Females fed E. kuehniella eggs had superior ovarian scores and laid more eggs than those fed either artificial diet or those given no extra food. A good correlation (r = 0.97) was obtained between ovarian scores and oviposition data, indicating that dissecting females after 1 week provides a reliable estimate of fecundity as affected by diet quality. Rapid reproductive assessments as used in the current study will help to increase the rate of development of artificial diets and may contribute to more cost effective production methods for augmentative biological control agents.     

Two Meridic Diets for Perillus bioculatus (Heteroptera: Pentatomidae), a Predator of Leptinotarsa decemlineata (Coleoptera: Chrysomelidae)

Two artificial diets, devoid of insect components, were developed for the predator Perillus bioculatus (Fabricius). The diets based on chicken liver and tuna fish were designed to approach the nutritional characteristics of the primary insect prey, Leptinotarsa decemlineata (Say) eggs. P. bioculatus was reared on diet 2 for 11 consecutive generations without cumulative detrimental effects on its biology. Developmental time and preovipositional period were significantly longer, and egg viability, survival from egg to adult, and fecundity were significantly lower in P. bioculatus individuals reared on artificial diets than in those reared on L. decemlineata eggs. A significant reduction in developmental time, increase in survival from egg to adult, and reduction in length of the preovipositional period were observed after 11 generations of in vitro reared P. bioculatus on diet 2. These changes may indicate that the predators experience some degree of adaptation to the diet after several consecutive generations of in vitro rearing. No changes in egg viability were observed after 11 generations of in vitro rearing. Adult female weight was unaffected when reared on artificial diet, but a significant increase was observed after 11 generations of in vitro rearing. Females under all treatments weighed an average of 11.6 mg more than the males.