Larval amphibian growth and development under varying density: are parasitized individuals poor competitors?

Population density and infection with parasites often are important factors affecting the growth and development of individuals. How these factors co-occur and interact in nature should have important consequences for individual fitness and higher-order phenomena, such as population dynamics of hosts and their interactions with other species. However, few studies have examined the joint effects of density and parasitism on host growth and development. We examined the co-influences of rearing density and parasitism, by the trematode Echinostoma trivolvis, on the growth and development of larval frogs, Rana (=Lithobates) pipiens. We also examined the potential role of parasite-mediated intraspecific competition by observing how unparasitized individuals performed when housed with other unparasitized tadpoles, versus housing with a combination of unparasitized and parasitized hosts. Mean mass and mean developmental stage were reduced under high rearing densities. The presence of parasitized conspecifics had no significant effect, but there was a significant interaction of density and parasitism presence on host mass, due to the fact that parasitized conspecifics grew poorly at high densities. Unparasitized individuals reared with parasitized and unparasitized conspecifics fared no better than unparasitized individuals reared only with one another. This result indicates that infected hosts compete as much as uninfected hosts for resources, even though infected individuals have reduced mass under high-density conditions. Resource acquisition and resource allocation are different processes, and parasitism, if it only affects the latter, might not have a discernible impact on competitive interactions.     

Are Sick Individuals Weak Competitors? Competitive Ability of Snails Parasitized by a Gigantism-Inducing Trematode

Parasitized individuals are often expected to be poor competitors because they are weakened by infections. Many trematode species, however, although extensively exploiting their mollusc hosts, also induce gigantism (increased host size) by diverting host resources towards growth instead of reproduction. In such systems, alternatively to reduced competitive ability due to negative effects of parasitism on host performance, larger size could allow more efficient resource acquisition and thus increase the relative competitive ability of host individuals. We addressed this hypothesis by testing the effect of a trematode parasite Diplostomum pseudospathaceum on the competitive ability of its snail host Lymnaea stagnalis. We experimentally examined the growth of snails kept in pairs in relation to their infection status and intensity of resource competition (i.e. food availability). We found that parasitized snails grew faster and their reproduction was reduced compared to unparasitized individuals indicating parasite-induced gigantism. However, growth of the snails was faster when competing with parasitized individuals compared to unparasitized snails indicating reduced competitive ability due to parasitism. The latter effect, however, was relatively weak suggesting that the effects of the parasite on snail physiology may partly override each other in determining competitive ability.     

中红侧沟茧蜂寄生对寄主粘虫血淋巴糖类、脂类和蛋白质含量的影响

为揭示寄生蜂寄生对其寄主的生理调控机制, 室内对中红侧沟茧蜂Microplitis mediator寄生与未被寄生寄主粘虫Mythimna separata幼虫血淋巴中糖类、脂类和蛋白含量变化进行了测定。结果显示: 在滞育与非滞育条件下, 被寄生的粘虫血淋巴中糖原浓度均比未被寄生的粘虫高。滞育条件下寄生后12 d差异显著（P<0.05）, 被寄生粘虫糖原含量为7.93 μg/mL, 未被寄生粘虫糖原含量为4.70 μg/mL; 非滞育条件下寄生后6 d差异显著（P<0.05）, 被寄生粘虫糖原含量为14.35 μg/mL, 未被寄生粘虫糖原含量为5.47 μg/mL。海藻糖含量测定结果显示, 在滞育条件下寄生蜂对被寄生粘虫无明显影响, 而非滞育条件下影响效果差异显著（P<0.05）, 寄生后4 d被寄生粘虫海藻糖含量为46.82 μg/mL, 未被寄生粘虫含量为26.72 μg/mL。在滞育与非滞育两种条件下, 寄生与未被寄生寄主脂类和蛋白含量没有显著性差异。结果说明: 寄生蜂的存在使寄主血淋巴中的糖原含量增高; 非滞育条件是影响被寄生粘虫海藻糖含量变化主要因素; 粘虫对中红侧沟茧蜂的寄生表现相当强的适应性和忍受力。     

Brood‐parasite‐induced female‐biased mortality affects songbird demography: negative implications for conservation

Parasites, of all sorts, can profoundly affect host population dynamics. Parasites commonly cause sex‐biased mortality and this can add to their impact. Female‐biased mortality in particular can destabilize dynamics and promote population collapse. We previously reported in a correlative study that brown‐headed cowbird Molothrus ater brood parasitism of song sparrows Melospiza melodia appears to cause female‐biased host nestling mortality. Here, we report results from ‘infestation’ and ‘de‐infestation’ experiments designed to test whether brood parasitism causes female‐biased mortality, and we document the resulting demographic impact using a simulation model. Experimental cowbird infestation of song sparrow nests halved the proportion of female host nestlings (0.31±0.07 vs 0.59±0.06; infested vs unparasitized nests at day 6) replicating the halving reported in naturally cowbird‐parasitized nests (0.28±0.01 vs 0.57±0.05; parasitized vs unparasitized). De‐infestation of naturally cowbird‐parasitized nests in turn wholly eliminated any effect on the proportion of female host nestlings (0.53±0.13 vs 0.54±0.06; de‐infested vs unparasitized) confirming that brood parasitism is the cause. This halving of the proportion of females fledging is likely to be as significant as nest predation in affecting population dynamics, based on the elasticities derived from our demographic model (–0.50 vs –0.59). Experimental infestation reduced the testosterone levels, begging behaviour, and body mass of six day old female host nestlings, whereas males were largely unaffected, suggesting that it is the exacerbation of intra‐brood competition that may be primarily responsible for the resulting female‐biased mortality. The brown‐headed cowbird is invasive in most of North America and has been implicated in regional population declines of many native species. We suggest that female‐biased host offspring mortality is likely to be commonplace among the 144 host species the cowbird successfully parasitizes, and we discuss the negative implications for songbird conservation, given the projected demographic impact.     

Oxygen consumption in mosquito larvae parasitized by Romanomermis culicivorax (Nematoda)

The effect of parasitism by Romanomermis culicivorax on the oxygen consumption of three species of mosquito larvae was studied by manometric methods. The Q02 of Culex pipiens, both parasitized and unparasitized, decreased as the host larvae developed from the second to fourth instar. In the early stages there was no significant difference in Q02 in parasitized and unparasitized hosts but the Q02 increased significantly in parasitized fourth instar larvae. The Q02 of parasitized and unparasitized fourth instars of Aedes taeniorhynchus, Anopheles quadrimaculatus and C. pipiens were compared. The Q02 of all parasitized hosts was significantly increased and the Q02 of the two culicine species displayed a dramatic increase in oxygen consumption. The oxygen consumption of the parasitic mermithid larvae was estimated from that obtained earlier for the post-parasitic larvae but did not account for more than 9-36% of the increased Q02 of the parasitized hosts. On this basis it seems probable that the increased metabolic rate as reflected by oxygen consumption of the parasitized mosquito larve was principally a manifestation of the physiological and nutritional stress induced by the mermithid nematodes.     

Intraguild predation and interference by the mealybug predator Cryptolaemus montrouzieri on the parasitoid Leptomastix dactylopii

The compatibility of the encyrtid parasitoid Leptomastix dactylopii with the coccinellid beetle Cryptolaemus montrouzieri against the citrus mealybug, Planococcus citri, is determined by the extent of intraguild predation and interference by the predator. We tested the preference of the adults and fourth-instar larvae of C. montrouzieri for healthy mealybugs and parasitized mealybugs harboring 1-, 4-, 7- and 14-day-old parasitoid larvae. The experiments were conducted in no-choice (only unparasitized mealybugs or parasitized mealybugs of one parasitoid age were offered at one time) and choice (unparasitized and parasitized mealybugs of a specific age were offered simultaneously) tests. Both the adults and larvae of C. montrouzieri fed on unparasitized and parasitized mealybugs but strongly discriminated against the hardened mummies (14 days old). We also investigated the influence of the presence of C. montrouzieri to the foraging effectiveness of L. dactylopii. The level of parasitism by L. dactylopii was reduced from about 13 to 6% when the number of the C. montrouzieri was increased from zero to four. We recommended that the releases of C. montrouzieri should be conducted 14 days after the releases of L. dactylopii to reduce intraguild predation on the parasitized mealybugs and to avoid interference with the foraging parasitoids.     

Parasitism may enhance rather than reduce the predatory impact of an invader

Invasive species can have profound impacts on communities and it is increasingly recognized that such effects may be mediated by parasitism. The ‘enemy release’ hypothesis posits that invaders may be successful and have high impacts owing to escape from parasitism. Alternatively, we hypothesize that parasites may increase host feeding rates and hence parasitized invaders may have increased community impacts. Here, we investigate the influence of parasitism on the predatory impact of the invasive freshwater amphipod Gammarus pulex. Up to 70 per cent of individuals are infected with the acanthocephalan parasite Echinorhynchus truttae, but parasitized individuals were no different in body condition to those unparasitized. Parasitized individuals consumed significantly more prey (Asellus aquaticus; Isopoda) than did unparasitized individuals. Both parasitized and unparasitized individuals displayed Type-II functional responses (FRs), with the FR for parasitized individuals rising more steeply, with a higher asymptote, compared with unparasitized individuals. While the parasite reduced the fitness of individual females, we predict a minor effect on population recruitment because of low parasite prevalence in the peak reproductive period. The parasite thus has a large per capita effect on predatory rate but a low population fitness effect, and thus may enhance rather than reduce the impact of this invader.     

Host selection and larval interactions among three primary parasitoids ofPlathypena scabra (Lep.: noctuidae)

Three hymenopteran parasitoids that attackPlathypena scabra (F.) larvae often oviposit into consecutive instars of the host. We investigated host discrimination by adults and competitive interactions among larvae of these three parasitoid species. Avoidance of superparasitism byCotesia marginiventris (Cresson) andDiolcogaster facetosa Ashmead was tested. Females of each species were presented either withP. scabra parasitized by a different female of the same species 6 h earlier or unparasitizedP. scabra. Under these conditions,C. marginiventris attacked similar numbers of parasitized and unparasitized hosts.D. facetosa attacked 31% fewer parasitized than unparasitizedP. scabra. The avoidance of multiple parasitism byD. facetosa was studied in a similar bioassay. AlthoughD. facetosa females parasitized fewerP. scabra that had been attacked byC. marginiventris 6 h previously, the reduction in parasitism was only about 23%. In competition studies, immatureD. facetosa were better competitors than immatureC. marginiventris. Aleiodes (=Rogas) nolophanae (Ashmead) was an inferior competitor against bothC. marginiventris andD. facetosa when the duration between parasitism events was 1 h, but their competitive ability increased when they multiply parasitized hosts at least 32 h after the initial oviposition.     

Acanthocephalan (Oligacanthorhynchidae) parasitism of the Caribbean termite <Emphasis Type="Italic">Nasutitermes acajutlae</Emphasis>: implications for reproductive success

Summary Few studies have examined the impact of parasitism on free-living social insects. We documented the association between an acanthocephalan parasite and survival, reproduction and growth of a Caribbean termite (Nasutitermes acajutlae). We measured these parameters yearly in 100 termite colonies for 4 years. In 2001 and 2002 we also compared the rate at which parasitized and unparasitized colonies rebuilt damaged foraging trails because trails protect foragers from predation as well as fluctuations in temperature and humidity. Although there were no significant differences in growth or survival between parasitized and unparasitized colonies, parasitized colonies reproduced significantly less often and rebuilt foraging trails significantly more slowly than unparasitized colonies. The average parasitized colony may sustain a loss of alate production =30% in the year in which it is parasitized, and a loss of up to 19% if it is parasitized once in a 5-year period. We conclude that the acanthocephalan parasite is associated with significant alteration of fitness parameters of individual N. acajutlae colonies. However, prevalence of parasitism is low (6.8%) and alate production by non-parasitized colonies is likely to compensate for losses due to parasitism. Therefore, it seems unlikely that this parasite affects N. acajutlae at the population level.Received 5 October 2003; revised 19 November 2003; accepted 3 December 2003.     

The glucogenic response of a parasitized insect Manduca sexta L. is partially mediated by differential nutrient intake

Induction of gluconeogenesis is accelerated in larvae of the insect Manduca sexta L. parasitized by Cotesia congregata (Say), maintaining the concentration of the blood sugar trehalose, an important nutrient for parasite development. Investigation has demonstrated that when host larvae are offered a choice of diets with varying levels of sucrose and casein, parasitized insects consume a different balance of these nutrients, principally due to a decrease in protein consumption. The result is metabolic homeostasis, with normal unparasitized and parasitized larvae exhibiting similar levels of gluconeogenesis and blood sugar level. In the present study, normal unparasitized and parasitized larvae were maintained on individual chemically defined diets having the balance of protein and carbohydrate consumed by each when offered a dietary choice. Total dietary nutrient, the sum of carbohydrate and protein, was provided at six levels, composed of three pairs of diets. Each diet pair consisting of diets having equivalent overall nutrient ratios of 2:1 and 1:1 casein/sucrose. Host growth and diet consumption were significantly affected by dietary nutrient level and the magnitude of these effects was influenced by parasitism. Due to the effects of dietary nutrient level on diet consumption, none of the unparasitized and parasitized larvae within any of the three diet pairs consumed protein and carbohydrate at the levels predicted by the earlier choice experiments. Among insects on all of the diets, however, two groups of unparasitized and parasitized larvae consumed the expected levels of protein and carbohydrate. In each case, gluconeogenesis, as measured by 13C nuclear magnetic resonance spectroscopy (NMR) analysis of pyruvate cycling and trehalose synthesis from [2-13C]pyruvate, was evident in unparasitized and parasitized insects, confirming the conclusions of the earlier experiments. Generally, all larvae that consumed less than approximately 250 mg of sucrose over the 3-day feeding period, were gluconeogenic, regardless of diet. Differential carbohydrate consumption, therefore, was an important factor in inducing gluconeogenesis in both unparasitized and parasitized insects. The selective 13C enrichment in trehalose displayed by non-gluconeogenic larvae on some diets demonstrated trehalose formation from [2]pyruvate. The absence of net carbohydrate synthesis in these insects was likely due to an elevation of glycolysis. There was no significant effect of diet consumption or parasitism on blood trehalose level. Parasitized larvae displayed higher levels of gluconeogenesis than did unparasitized insects, a finding consistent with the conclusion that blood sugar is rapidly sequestered by developing parasites. The parasite burden, the total number of parasites developing within host larvae, as well as the number of parasites emerging from host larvae to complete development, was significantly less at the lowest dietary nutrient level, but was otherwise similar at all dietary nutrient levels. Moreover, the number of parasites that emerged increased with increasing diet consumption as reflected by host final weight.     

斑痣悬茧蜂寄生对甜菜夜蛾幼虫取食和食物利用的影响

为研究甜菜夜蛾Spodoptera exigua幼虫被斑痣悬茧蜂Meteorus pulchricornis寄生后的取食以及食物利用情况,在室内采用重量法测定了甜菜夜蛾4龄幼虫被寄生后取食量、体重增加量、营养指标的变化。结果表明：被寄生甜菜夜蛾幼虫的取食量、生长率和食物利用效率等明显受到抑制,幼虫被寄生后第3-6 d 取食量显著小于未被寄生幼虫,寄生后第4 d 的幼虫取食量只有正常幼虫的29.89%,第5 d只有48.69%。幼虫在寄生后的第3-5 d体重增加量显著小于未被寄生幼虫,分别为正常幼虫的21.51%,38.87%和14.42%,相对生长率则也显著低于后者。被寄生甜菜夜蛾幼虫的营养利用表现也明显不同于未被寄生幼虫,反映生长和代谢效率的食物利用率（ECI）和食物转化率（ECD）均显著降低,而反映吸收效率的近似消化率（AD）则提高。虽然在寄生后第4 d出现了相反的现象,其原因可能在于第4 d取食量明显减少,而体重仍在增加。本研究表明,斑痣悬茧蜂寄生明显抑制寄主甜菜夜蛾幼虫的取食、食物利用效率和生长。     

Consequences of Interspecific Competition on the Virulence and Genetic Composition of a Nucleopolyhedrovirus in Spodoptera frugiperda Larvae Parasitized by Chelonus insularis

Nucleopolyhedroviruses ( Baculoviridae ) are virulent insect pathogens that generally show a high degree of host specificity and have recognized potential as biological insecticides. Whenever viruses are applied for pest control, a proportion of the infected insects will also be parasitized by hymenopteran or dipteran parasitoids and interspecific competition for host resources will occur; the severity of such competition is likely to be modulated to a large degree by the virulence of each type of parasite. We examined the impact of parasitism by the solitary egg-larval endoparasitoid Chelonus insularis (Hymenoptera: Braconidae) on the speed of kill of nucleopolyhedrovirus-infected Spodoptera frugiperda (Lepidoptera: Noctuidae) larvae and the pattern of host growth and virus production in infected and/or parasitized hosts. We also examined the effect of parasitism on the virulence, infectivity and genetic composition of serially passaged virus. Both parasitism and viral infection resulted in a marked reduction in host growth. When third instar larvae were dually parasitized and virus-infected, the growth rate was even more severely affected compared to parasitized larvae. There was a significant increase in virus production in larvae infected at later instars. Interspecific competition resulted in a substantial decrease in pathogen production in parasitized larvae infected at the fourth instar, but not in parasitized larvae infected at earlier instars. The serial passage experiment resulted in the appearance of four distinct genetic isolates of the virus detected by restriction endonuclease analysis. Of the three isolates that appeared in nonparasitized larvae, two showed increased virulence, expressed by mean time to death, and for one of these the infectivity, expressed as LC 50 , was reduced. One isolate that appeared in parasitized larvae (isolate D) had increased virulence and infectivity. Southern blot analysis indicated that virus isolate D was most likely generated by point mutation of a restriction site or by alterations such as duplications, deletions or by recombination of two or more genotypic variants present in the wild-type nucleopolyhedrovirus isolate. Our study provides clear evidence of interspecific competition within the host, since, depending on the timing of inoculation, adverse effects were observed upon both the parasitoid and the virus.     

淡足侧沟茧蜂寄生对甜菜夜蛾幼虫血淋巴总糖、蛋白质及虫体脂质含量的影响

为探讨淡足侧沟茧蜂Microplitis pallidipes Szepligeti调控寄主的生理机制,测定了淡足侧沟茧蜂寄生后,甜菜夜蛾Spodoptera exigua(Hübner)幼虫血淋巴总糖、蛋白质及虫体脂类含量的变化。研究结果表明,寄生后连续5 d的观察时间内,甜菜夜蛾幼虫血淋巴总糖含量从寄生后第1天开始就高于未寄生寄主幼虫,且在寄生后第2至5天达到显著水平;除寄生后第3天外,被寄生寄主幼虫血淋巴总蛋白质含量始终低于未寄生幼虫,且在寄生后第1、4、5天达到显著水平;甜菜夜蛾被寄生后,虫体脂质含量始终高于未寄生幼虫,且从寄生后第2天开始达到显著水平。本研究揭示,淡足侧沟茧蜂寄生刺激了寄主甜菜夜蛾幼虫血淋巴总糖合成,但抑制了其蛋白质合成,同时也刺激了甜菜夜蛾虫体脂质合成。     

IMPACT OF IMMUNE RESPONSE OF A PARASITIC BEETLE Dastarcus helophoroides ON ITS HOST BEETLE Monochamus alternatus

Dastarcus helophoroides is an ectoparasitoid beetle of Monochamus alternatus, and the parasitism by D. helophoroides larvae remarkably influenced on the immune responses of M. alternatus larvae in many aspects. The hemolymph melanization reactions in the hosts were inhibited 1 h and 24 h postparasitization. The phenoloxidase activities of hemolymph were significantly stimulated 4 h postparasitization and inhibited 12 h postparasitization, and back to control level. The antibacterial activities of hemolymph in the parasitized hosts were significantly lower than that in the unparasitized ones 1 h postparasitization. By 72 h postparasitism, the total hemocyte numbers of the parasitized larvae declined to not more than one‐seconds of the number collected from the unparasitized larvae. All sampled hemolymph held the capability of nodulation, and there were fluctuations in the number of nodules the hemocytes made. However, there were no significant differences between unparasitized and parasitized larvae at each time point in the hemagglutination activity and the ratios of spreading hemocytes. In conclusion, D. helophoroides larvae could regulate M. alternatus immune system and resulted in the changes in host immune responses.     

Effects of parasitization by Cotesia congregata on the brain-prothoracic gland axis of its host,Manduca sexta

The ability of prothoracic glands (PTGs) from parasitized and unparasitized Manduca sexta 5th-instars to respond to ecdysiotropic extracts prepared from day-5 5th instar brains was compared. An in vitro bioassay revealed that PTGs from parasitized animals were much less responsive to brain PTTH than glands from unparasitized larvae. However, when incubated in Grace's medium in the absence of brain extract, glands from day-3 and -4 hosts remained active for a much longer period of time than did those dissected from their unparasitized counterparts. Rather than exhibiting reduced (basal) levels of synthesis after the 3rd hour of incubation, glands from these parasitized larvae continued to synthesize/release ecdysteroid into the medium at relatively high rates. The timing of this enhanced secretory activity is coincident with the ecdysteroid peak that occurs just prior to and during wasp emergence. Following parasite emergence, gland activity decreased, and by the third day after emergence, was reduced to low levels. Results suggest that the requirement for PTTH to stimulate ecdysteroid production has been bypassed, i.e. that the parasite has uncoupled the normal mechanisms that permit brain regulation of PTG activity. The ability of brains from parasitized M. sexta to stimulate PTGs from unparasitized day-2 5th instars was also examined. Dose-response analyses performed for the first 7 days of the 5th instar showed that on a per brain basis ecdysiotropic activity in brains from parasitized and unparasitized animals was similar. However, when differences in brain size were considered, ecdysiotropic activity appeared to be more concentrated in brains from day-7 parasitized larvae than in brains from similarly aged unparasitized larvae. Analysis of the size distribution of the ecdysiotropic activity in brains from parasitized larvae revealed a unique form that was larger than the 29kDa standard. This suggests that parasitization may inhibit neuropeptide processing, particularly during the final stages preceding emergence of the wasps from the host. Thus, both an inhibition of prothoracicotropic hormone processing and the inability to respond to this neurohormone may contribute to the developmental arrest characteristic of parasitized 5th instars.     

Influence of the braconid Glyptapanteles liparidis on the juvenile hormone titer of its larval host,the gypsy moth,Lymantria dispar

The increase in the juvenile hormone (JH) III titer in the hemolymph of Lymantria dispar larvae that were parasitized by the endoparasitoid braconid, Glyptapanteles liparidis, during the host's premolt to third instar, coincided with the molt of the parasitoid larvae to the second instar between day 5 and 7 of the fourth host instar. It reached a maximum mean value of 89 pmol/ml on day 7 of the fifth instar while it remained below 1 pmol/ml in unparasitized larvae. Only newly molted fifth instar hosts showed a low JH III titer similar to that of the unparasitized larvae. JH II, which is the predominant JH homologue in unparasitized gypsy moth larvae, also increased relative to controls in the last two samples (days 7 and 9) from parasitized fourth and fifth instars. Compared to unparasitized larvae, a generally reduced activity of JH esterase (JHE) was found in parasitized larvae throughout both larval stages. The reduction in enzyme activity at the beginning and at the end of each instar, when the JHE activity in unparasitized larvae was high, may be in part responsible for the increased JH II and JH III titers in parasitized larvae. Ester hydrolysis was the only pathway of JH metabolism in the hemolymph of unparasitized and parasitized gypsy moth larvae as detected by chromatographic assays. © 1996 Wiley-Liss, Inc.     

Sexually transmitted parasites and sexual selection in the milkweed leaf beetle, Labidomera clivicollis

Milkweed leaf beetles (MLBs; Labidomera clivicollis Chrysomelidae) are parasitized by a subelytral mite, Chrysomelobia labidomerae (Tarsonemina: Podapolipidae). We show that C. labidomerae is transmitted between MLBs when they copulate, and can reduce the survival of nutritionally stressed beetles. We investigated the effect of this sexually transmitted parasite on mate choice and male-male competition in MLBs, and the consequences of variation in these behaviours for mite transmission. We found no evidence of parasite avoidance by MLBs, and evidence for high rates of parasitism in the MLB populations we surveyed. In the absence of females, parasitized males contacted unparasitized males more often and for longer than controls, and they tended to displace rival males from females more often than did unparasitized males, a result consistent with the interpretation that parasitized beetles compensate for loss of fitness by increasing reproductive effort. These changes can also benefit mites, because longer and possibly more contacts between beetles provide more opportunities for transmission, but there is no evidence that these changes in male behaviour result from parasite manipulation.     

Foraging behavior of the coccinellid Nephus includens (Coleoptera: Coccinellidae) in response to Aphis gossypii (Hemiptera: Aphididae) with particular emphasis on larval parasitism

This study assessed the effect of parasitism of Nephus includens (Col.: Coccinellidae) larvae by Homalotylus flaminius (Hym.: Encyrtidae) on the predation rates of the predator on the cotton aphid, Aphis gossypii Glover (Hem.: Aphididae) by deriving functional responses for second- and fourth-instar predators at prey densities ranging from 10 to 80 aphids per arena. The relationship between the functional and numerical responses of adult females of N. includens also was determined for prey densities ranging from 10 to 140 aphids per arena. Predation rates of unparasitized and parasitized second-instar N. includens were both fit by a type II functional response model with parameters as follows: unparasitized (a = 0.0768 hours(-1) and T(h) = 0.975 h) and parasitized (a = 0.0787 hours(-1) and T(h) = 0.8823 hours). Predation rates of unparasitized and parasitized fourth-instar N. includens were fit by type III and II models, respectively, with the following parameters: unparasitized (b = 0.1702 hours(-1) and T(h) = 0.2369 hours) and parasitized (a = 0.038 h(-1) and T(h) = 0.539 h). The unparasitized fourth-instar was the most voracious stage, having the highest attack rate and lowest handling time. Considering these attributes, it would seem to be the most effective stage of this predator against A. gossypii. Adult female lady beetles (N. includens) showed a type III functional response and their numerical response increase to prey density was curvilinearly related to prey density, with the highest number of eggs being produced at highest prey densities. The maximum saturation level for both predation and egg production for adult females occurred at a prey density of 120 aphids. Thus, a ratio 1:120 (predator:prey) should be used when releasing this species for augmentative biological control. Release of either fourth-instar or adult stage N. includens should be minimized the potentially negative effect of parasitism by H. flaminius on early developmental stages, and hence increase its efficiency in biocontrol programs.