Evidence for a marking pheromone in host discrimination byCephalonomia Stephanoderis (Hym.: Bethylidae)

The bethylidCephalonomia stephanoderis Betrem is an ectoparasitoid that prefers to oviposit on the prepupae and pupae of the coffe berry borerHypothenemus hampei (Ferrari) (Coleoptera: Scolytidae). It has the ability to distinguish unparasitized from parasitized hosts and rarely lays more than one egg per host. The mechanism of this host discrimination byC. stephanoderis was investigated under laboratory conditions. For this, parasitoid eggs that had been deposited on host pupae were removed and pupae were then offered (individually and collectively) to individual female wasps. A total of 92% of individually offered hosts and 93% of collectively offered hosts were not parasitized. It is concluded thatC. stephanoderis recognizes a marking pheromone deposited into or onto the host, preceding, during, or after oviposition which enables female parasitoids to avoid self and conspecific superparasitism.     

Alternative Hosts for Bethylid Parasitoids of the Coffee Berry Borer, Hypothenemus hampei (Coleoptera: Scolytidae)

Three species of bethylid wasps, Prorops nasuta Waterston, Cephalonomia stephanoderis Betrem, and C. hyalinipennis Ashmead, attack the coffee berry borer, Hypothenemus hampei (Ferrari) (Coleoptera: Scolytidae), by both predation and parasitism. C. hyalinipennis has only recently been reported to attack H. hampei. Its previously recorded hosts belong to the coleopteran families Curculionidae, Anobiidae, and Scolytidae. We evaluated five further coleopteran species, in the families Curculionidae, Bostrichidae, and Bruchidae, as alternative hosts. High proportions of all five species were both fed upon and oviposited on (normally by concurrent host feeding). Offspring developmental mortality varied greatly (12–95%) on different host species. Offspring production per host was about three times greater than on H. hampei when females were presented with the curculionids Caulophilus oryzae (Gyllenhal) and two species of Sitophilus. C. stephanoderis and P. nasuta are currently the main biological control agents of H. hampei: they have long been thought to be monophagous but C. stephanoderis has recently been reported to attack H. obscurus. We evaluated C. oryzae, as a potential host or prey for P. nasuta, and both C. oryzae and Sitophilus sp. for C. stephanoderis. Host feeding was very common in both parasitoid species. The majority of females also oviposited on C. oryzae (71% P. nasuta, 60% C. stephanoderis) and 20% of C. stephanoderis parasitized Sitophilus sp. Fifty-two percent of C. stephanoderis and 7% of P. nasuta offspring survived to adulthood when developing on C. oryzae. The lifetime fecundities of these parasitoids presented with these host species were, however, low (maximum 17 eggs, mean ≤ 8.25) and preoviposition periods were generally long (ca. 19 days). Adult female C. stephanoderis lived longer when presented C. oryzae rather than Sitophilus sp. Host feeding also increased longevity. We discuss these results in the context of mass rearing these bethylids for release as biological control agents.     

Influence of host density on the reproductive strategy of Cephalonomia stephanoderis, a parasitoid of the coffee berry borer

The effect of the density of the coffee berry borer, Hypothenemus hampei (Coleoptera: Scolytidae) on reproductive and host-feeding behaviours of the parasitoid Cephalonomia stephanoderis (Hymenoptera: Bethylidae) was evaluated under laboratory conditions. The number of hosts used for oviposition was density-dependent at low host density. Beyond a density of six hosts/day, the oviposition rate reached a maximum of 1.2 eggs/day due to egg limitation. Cephalonomia stephanoderis females responded to increasing host availability with a linear increase in host feeding. Overall, parasitoids killed more coffee berry borers by feeding and paralysis than by parasitism. At low host density, the pre-oviposition phase was extended, oogenesis was delayed, more males were produced, and host feeding occasionally occurred concurrently with oviposition. We suggest that the efficacy of C. stephanoderis as a biological control agent depends on seasonal variations in host density. Inoculative vs inundative releases in coffee plantations are discussed in relation to the abundance of the coffee berry borer during the fructification and interharvest periods.     

Life history studies of <Emphasis Type="Italic">Prorops nasuta</Emphasis>, a parasitoid of the coffee berry borer

Life history studies were conducted in the laboratory on the African parasitoid Prorops nasuta Waterston (Hymenoptera: Bethylidae), a parasitoid of the coffee berry borer, Hypothenemus hampei (Ferrari) (Coleoptera: Scolytidae). The female wasp enters an infested coffee berry, kills the adult borer and seals the entrance of the berry with the body of the borer, impeding the entry of other organisms into the berry. The preoviposition period ranges from 3 to 14 days (mean 5.42 ± 0.37 SE). During this time females feed on the immature stages and paralyse fully grown larvae and pupae of the CBB. P. nasuta is an idiobiont solitary ectoparasitoid. Eggs are laid externally on the last instar larvae and pupae. Mean development time (egg to adult) for males and females was 27.7 (±0.37 SE) and 30 (±0.12 SE) days, respectively. Median survival for wasps fed on final instar CBB larvae was 27.7 days, significantly longer than any other treatment, while for females without food it was 2.5 days. In culture, females produced an average of 4.3 (±0.39 SE) progeny during their lifetimes. Adults began emerging at 30.6 days (±0.28 SE) after cultures were started and peak production was reached at 36 days, declining thereafter. Males normally emerged from coffee beans 2–3 days before females. Males usually emerged from 07:00 to 09:00h and females from 10:00 to 14:00 h. The culture sex ratio (proportion of males) was 0.21. Virgin females produced only male offspring.     

Comparison of Host-Searching and Ovipositional Behavior of Cardiochiles nigriceps Viereck (Hymenoptera: Braconidae), a Parasitoid of Heliothis virescens (Fabricius) (Lepidoptera: Noctuidae), in Tobacco and Cotton

To compare host-searching and ovipositional behavior of Cardiochiles nigriceps Viereck (CN) in tobacco and cotton, behavior of CN females was observed on Hellothis virescens (Febricius)-infested plants and recorded using a Psion Organizer II with Observer software. Hover, search, oviposit, preen, rest, walk, agony-search, drop-search, and miss-search were the nine behaviors observed for each plant species. Agony-search was a searching behavior exhibited when CN females were prevented from ovipositing in a host because they were unable to reach the host. Drop-search and miss-search were the searching behaviors that occurred by CN females after a host dropped from the plant and when CN females missed ovipositing in a host, respectively. Hovering, searching, oviposition, miss-search, resting, and walking were equally common for CN on tobacco and cotton. Agony-search was exhibited more on tobacco than cotton, while drop-search and preening occurred more on cotton than tobacco. Female CN spent more time hovering in tobacco than in cotton, indicating that these females preferred tobacco to cotton. Female parasitoids also spent more time in the main ovipositional sequence, hover–search–oviposit, in tobacco than in cotton, mainly because females hovered more in tobacco than in cotton. Frequency and time of behavioral sequences associated with experiencing frustration in ovipositing due to difficulty in reaching or finding a host and subsequent level of success in ovipositing in a new host were determined. Overall, the rate of successful oviposition by CN females was significantly higher in tobacco (87.7%) than in cotton (79.5%). For cotton and tobacco, the rate of successful oviposition by CN females was significantly reduced (ca. 40.0%) when they experienced frustration in ovipositing compared to the rate of ovipositional success (100%) for CN females when hosts were present and easily accessible. Encountering ovipositional frustration also increased the duration of time for successful oviposition in comparison to the time spent for problem-free oviposition in cotton and tobacco. Interestingly, with successful oviposition, the amount of time spent in a behavioral sequence was not significantly different for tobacco and cotton regardless of whether females experienced ovipositional frustration or not. However, female CN invested more time in tobacco than in cotton when they were unsuccessful in ovipositing due to the inaccessibility or disappearance of hosts. CN females' preference for tobacco over cotton probably gave them more motivation to continue searching for hosts, especially those not easily attained, on tobacco than on cotton. Nevertheless, CN females readily searched in cotton for HV even in the presence of host-infested tabacco. CN females possibly could be successful in maintaining levels of HV below an economic threshold when using a tobacco trap crop to protect cotton as the main crop.     

Oviposition behavior of Phymastichus coffea LaSalle (Hymenoptera: Eulophidae) on its natural host

Phymastichus coffea LaSalle is an African endoparasitoid of adults of the coffee berry borer, Hypothenemus hampei (Ferrari) that was introduced into Mexico for biological control purposes. The objective of this study was to investigate the oviposition behavior of P. coffea on its host under laboratory conditions. The results indicate that P. coffea females exhibited 12 behavioral acts during and after oviposition, such as, walking, cleaning, flying, surface landing, resting, host landing, host antennation, touching the host with forelegs, host recognition, walking on the host, oviposition and cleaning after oviposition. The sequence of each behavior is described in an ethogram.     

Behavioural responses of bethylid parasitoid species of the coffee Berry borer to chemicals cues from host and non-host dust/frass

The bethylid wasps Cephalonomia stephanoderis Betrem and Prorops nasuta Waterston are parasitoids of the coffee berry borer, Hypothenemus hampei (Ferrari), the most serious insect pest of coffee worldwide. Recent studies on the host location behaviour of these parasitoids have shown that females are attracted to volatile compounds released by immature stages and dust/frass of H. hampei. Also, these studies have reported that the locomotory behaviour of parasitoids is influenced by contact chemical cues from host dust/frass. In this study, we investigated the responses of females of both species to volatiles and contact cues from dust/frass from different sources. Volatiles from dust/frass from H. hampei-infested coffee and maize infested with Sitophilus sp were attractive to P. nasuta females. In contrast, C. stephanoderis females were only attracted to the volatiles from dust/frass from H. hampei-infested coffee. Volatiles from an artificial diet used for rearing H. hampei were not attractive to females of both species. The effect of methanolic extracts from dust/frass from different sources on the patch-searching time of parasitoids was analysed using the Ethovision software. P. nasuta females spent more time in the patches treated with extracts of dust/frass from H. hampei-infested berries, artificial diet, and from maize infested with Sitophilus sp than on patches treated with dry coffee extract or solvent control. C. stephanoderis females spent more time on patches treated with dust/frass extracts from H. hampei-infested berries and artificial diet than on patches treated with an extract of dust/frass from maize infested with Sitophilus sp, an extract of dry coffee, and methanol control.     

Possible Age-Dependent Variation in Egg-Loaded Host Selectivity of the Pierid Butterfly, Anthocharis scolymus (Lepidoptera: Pieridae): A Field Observation

To understand better the effects of age on host selection through conspecific egg detection by Anthocharis scolymus females, field observations were performed at a graveyard where the insect population was isolated and the host plant was limited to the cruciferous plant, Turritis glabra. We chased females and recorded their oviposition behaviors and the conditions of plants which females approached. Older females tended to avoid ovipositing on egg-loaded host plants and selectively laid eggs on nonloaded host plants. This result was not confounded by other factors such as seasonality, air temperature, plant height, plant phenological stage, surrounding vegetation, host plant density, and extent of plant damage. We discuss the possibility that females make an oviposition decision on how to lay their limited number of eggs during their remaining lifetime.     

Discrimination by Coptera haywardi (Hymenoptera: Diapriidae) of hosts previously attacked by conspecifics or by the larval parasitoid Diachasmimorpha longicaudata (Hymenoptera: Braconidae)

Coptera haywardi (Oglobin) is an endoparasitoid of fruit fly pupae that could find itself in competition with other parasitoids, both con- and heterospecific, already resident inside hosts. In choice bioassays, ovipositing C. haywardi females strongly discriminated against conspecifically parasitised Anastrepha ludens (Loew) pupal hosts. They also avoided pupae previously attacked by Diachasmimorpha longicaudata (Ashmead), a larval–prepupal koinobiont endoparasitoid, and the degree of larval-parasitoid superparasitism had no effect on this avoidance. There was no difference in the number of ovipositor insertions when hosts previously parasitised by a conspecific and D. longicaudata were exposed simultaneously. As females aged the degree of host discrimination declined. An ability to discriminate against pupae previously attacked as larvae suggests low levels of both conspecific and heterospecific competition in the field.     

Aphanogmus sp. (Hymenoptera: Ceraphronidae): a hyperparasitoid of the coffee berry borer parasitoid Prorops nasuta (Hymenoptera: Bethylidae) in Kenya

This is the first report of a hyperparasitoid of Prorops nasuta, a primary parasitoid of the coffee berry borer. Aphanogmus sp. is a gregarious ectoparasitoid of larval and pupal stages of P. nasuta, which was found in coffee berry samples collected on the ground of an organic coffee plantation in Western Kenya. The hyperparasitoid shows a clear pattern of emergence from year to year, following its host. Aphanogmus sp. parasitizes around 10% of P. nasuta immature stages under field conditions.     

Costs and benefits of host feeding in the parasitoid wasp Trichogramma turkestanica

Females of the parasitoid wasp Trichogramma turkestanica Meyer (Hymenoptera: Trichogrammatidae) generally host feed after ovipositing on the first egg of Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) they encounter. We measured the impact of host feeding on the fecundity and longevity of females, in absence of host or food, and on the fitness of their progeny. We also determined if the frequency of host feeding is influenced by the humidity level at which T. turkestanica females developed. Host feeding increased egg production by 70% but decreased female longevity. This impact of host feeding on the longevity of females is probably due to the allocation of carbohydrates to egg production at the expense of somatic maintenance. Humidity did not influence the occurrence or duration of host feeding. The size of individuals developing in eggs on which females host fed was smaller, indicating that their fitness was affected.     

Foraging Behavior and Patch Time Allocation by Fopius arisanus (Hymenoptera: Braconidae), an Egg-Larval Parasitoid of Tephritid Fruit Flies

This study quantitatively describes the host-searching behavior of Fopius arisanus (Sonan) (Hymenoptera: Braconidae), an important egg-larval parasitoid of tephritid fruit fly pests, on coffee berries infested with host eggs of the Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae). We also investigate the parasitoid's response to local variation in host patch quality. The temporal pattern of behavioral organisation was examined by constructing an ethogram. The parasitoid spent over 90% of its foraging time in detecting and locating hosts after arriving on a host-infested fruit, and displayed a relatively fixed behavioral pattern leading to oviposition. Patch residence time increased in the presence of host-associated cues, following successful ovipositions, and with increasing size of host clutches per fruit, but decreased with each successive visit to the same host patch and with increasing availability of alternative host patches. The parasitoid females discriminated against previously parasitized hosts and spent significantly less time and searching effort on patches previously exploited by herself or by conspecific females. The effective host-searching behavior, perfect host discrimination ability, and success-motivated searching strategy shown by F. arisanus ensured a thorough exploitation of host resources by this parasitoid.     

Polyphagy and primary host plants: oviposition preference versus larval performance in the lepidopteran pest Helicoverpa armigera

Oviposition preference and several measures of offspring performance of Helicoverpa armigera (Hübner) were investigated on a subset of its host plants that were selected for their reputed importance in the field in Australia. They included cotton, pigeon pea, sweet corn, mungbean, bean and common sowthistle. Plants were at their flowering stage when presented to gravid female moths. Flowering pigeon pea evoked far more oviposition than did the other plant species and was the most preferred plant for neonate larval feeding. It also supported development of the most robust larvae and pupae, and these produced the most fecund moths. Common sowthistle and cotton were equally suitable to pigeon pea for larval development, but these two species received far fewer H. armigera eggs than did pigeon pea. Mungbean also received relatively few eggs, but it did support intermediate measures of larval growth and survival. Fewest eggs were laid on bean and it was also the least beneficial in terms of larval growth. Among the host plant species tested, only flowering pigeon pea supported a good relationship between oviposition preference of H. armigera and its subsequent offspring performance. Australian H. armigera moths are thus consistent with Indian H. armigera moths in their ovipositional behaviour and larval performance relative to pigeon pea. The results suggest that the host recognition and acceptance behaviour of this species is fixed across its geographical distribution and they support the theory that pigeon pea might be one of the primary host plants of this insect. These insights, together with published results on the sensory responses of the females to volatiles derived from the different host plant species tested here, help to explain why some plant species are primary targets for the ovipositing moths whereas others are only secondary targets of this polyphagous pest, which has a notoriously broad host range. Handling Editor: Joseph Dickens     

Vetiver grass (Vetiveria zizanioides (L.) Nash) as trap plant for Chilo partellus (Swinhoe) (Lepidoptera: Pyralidae) and Busseola fusca (Fuller) (Lepidoptera: Noctuidae)

The preference of lepidopterous stem borer moths to oviposit on certain wild host plants can be exploited in habitat management systems by using those hosts as trap crops. Vetiver grass (Vetiveria zizanioides (L.) Nash) was evaluated for its attractiveness and suitability to the pyralid Chilo partellus (Swinhoe) (Lepidoptera: Pyralidae) and the noctuid Busseola fusca (Fuller) (Lepidoptera: Noctuidae). Two choice tests were conducted in the laboratory and in the greenhouse to determine oviposition choice of C. partellus for maize, Vetiver and rice (Oryza sativa L.), and of B. fusca for Vetiver and maize. C. partellus larval survival was evaluated in green house studies. Results indicated that C. partellus chose Vetiver grass over maize though larval survival on Vetiver was extremely low. B. fusca did not show any host preference.     

Acceptance and suitability of four lepidopteran stemborers for the development of the pupal parasitoid Xanthopimpla stemmator (Hymenoptera: Ichneumonidae)

Xanthopimpla stemmator (Thunberg), a solitary endoparasitoid of lepidopteran stemborer pupae, was recently imported into East Africa as a candidate biological control agent of gramineous stemborers. Suitability of Busseola fusca Fuller, Chilo partellus (Swinhoe), Eldana saccharina (Walker) and Sesamia calamistis Hampson, for the development of X. stemmator was studied in the laboratory. One- to 6-day-old laboratory reared pupae of the four stemborer species were exposed to naïve X. stemmator females. All host pupae and ages were acceptable for oviposition. The parasitoids inflicted multiple probe wounds on 67.8% of pupae exposed. B. fusca, C. partellus and S. calamistis were equally suitable with 56.4, 59.4 and 52.3%, respectively, of probed pupae leading to emergence of adult parasitoids. E. saccharina was less suitable with only 22.6% of probed pupae producing parasitoids. Emergence of parasitoids did not differ significantly across the six pupal ages for B. fusca and S. calamistis, but varied for C. partellus and E. saccharina. No parasitoids emerged from 6-day-old E. saccharina pupae. Realized fecundity of females reared on the four stemborer pupae showed that fewer progeny were produced by females emerging from E. saccharina than females reared on the other three stemborer species. Eldana saccharina may be a poor host for X. stemmator in Kenya, but this parasitoid is a potential candidate for biological control of B. fusca, C. partellus and S. calamistis.     

Host plant selection by a monophagous herbivore is not mediated by quantitative changes in unique plant chemistry: Agonopterix alstroemeriana and Conium maculatum

Host plant selection by ovipositing females is a key process determining the success of phytophagous insects. In oligophagous lepidopterans, host-specific plant secondary chemicals are expected to be dominant factors governing oviposition behavior; distinctive compounds can serve as high-contrast signals that clearly differentiate confamilial hosts from non-hosts increasing the accuracy of host quality evaluation. Agonopterix alstroemeriana (Clerk) (Lepidoptera: Oecophoridae) and Conium maculatum L. (Apiaceae) form an extremely specialized plant-herbivore system, with A. alstroemeriana monophagous on C. maculatum, a plant with few other insect herbivores at least in part due to its virtually unique capacity among plants to produce piperidine alkaloids. Here we have studied the response of A. alstroemeriana oviposition to unique host plant secondary metabolites, piperidine alkaloids, and widespread compounds, mono- and sesquiterpenes, in a concentration-dependent fashion. Rates of oviposition were negatively correlated with Z-ocimene concentrations. To confirm the deterrent properties of this monoterpene for A. alstroemeriana oviposition, we conducted a choice experiment using artificially damaged C. maculatum plants, with higher emission of volatiles, and undamaged control plants. Damaged plants were less preferred as oviposition sites compared to the controls. The lack of association between oviposition and piperidine alkaloids, defenses unique to Conium species, suggests that quantitative changes of these species-specific chemicals do not play a predominant role in host selection by the monophagous A. alstroemeriana.     

Influence of host age and host deprivation on longevity, egg load dynamics and oviposition rate in Microplitis rufiventris

Adult size, longevity, egg load dynamics and oviposition ofMicroplitis rufiventris Kok. which began their development in the first, second, third （preferred hosts） or fourth （non-preferred hosts） instar larvae of Spodoptera littoralis （Boisd.） were studied. The parasitoid size was largely determined by the initial host size at parasitism. Non-ovipositing females derived from older hosts lived for longer periods than those derived from younger ones. However, the ovipositing females, irrespective of their size, lived for almost the same periods. At emergence, the oviducts of adult females contain a significant amount of mature eggs available for oviposition for a few hours on eclosion day. Egg load increases during the early phase of adult life. The amount of additional mature eggs and rate of egg maturation per hour was greater for wasps derived from preferred hosts compared with those in females derived from non-preferred hosts. The pattern of egg production in M. rufiventris females depended on the availability of hosts for parasitization. Host-deprived females depleted the egg complement with aging; the longer the host deprivation, the lower the oviduct egg load. Marked reduction in both realized or potential fecundity of host-deprived females was observed following host availability. Host privation for more than 3 days induced a marked deficit fecundity pattern through the female＇ s life. The realized fecundity was determined by the interaction among host availability, the number of eggs that are matured over the female＇ s life span, oviposition rate and host size from which the female was derived. These results suggest that： （i） M. rufiventris wasp is a weak synovigenic species; （ii） the maturation of additional eggs is inhibited once the maximum oviduct egg load is reached; （iii） the egg load of the newly emerged female is significantly less than the realized fecundity; and （iv） because M. rufiventris females oviposit fewer eggs when they begin depleting their egg supply at 3 days, augmentative releases will require release immediately following emergence to ensure the highest parasitization rate in the field.     

蝶蛹金小蜂在杭州的年生活史

蝶蛹金小蜂是菜粉蝶蛹期的主要寄生蜂。在杭州,绝大多数以老熟幼虫在寄主蛹内越冬。田间养虫室内饲养结果:1979年发生11—12代;1981年早、中、迟三组分别为12—13代、10—11代以及9—10代,三组合计,非越冬代成虫自5月上旬至12月初始终不断,一年之内,前后(连续或不连续)两代成虫重叠发生的天数至少175天,前后(连续或不连续)三代成虫重叠发生的天数至少49天。 2、3月间小蜂即可羽化,远远早于田阔第一代菜粉蝶蛹的出现,惟不能寄生越冬后的越冬代菜粉蝶蛹,但越冬代蜂的羽化盛期仍处于田间第一代菜粉蝶蛹初见至化蛹始盛期之间,且喂食后寿命很长,故越冬代蜂与第一代菜粉蝶蛹之间的发生期基本上是吻合的。其后,两者代数既多(菜粉蝶一年发生9代),发生又都不整齐,小蜂成虫与菜粉蝶蛹均连绵不断,时间上彼此也是吻合的。这是此蜂自然寄生率高并成为其重要天敌的一个主要原因。 对发生代数多、世代重叠的重要天敌建议分期分批饲养,以深入掌握其生活史及与寄主发生的关系。     

The effects of feeding damage in ragweed Ambrosia artemisiifolia (Asteraceae) on populations of Zygogramma suturalis (Coleoptera,Chrysomelidae)

Summary Field sampling indicated that the number of eggs laid by a Zygogramma suturalis female within a sampling plot correlated inversely with the mean degree of ragweed damage. Feeding on extensively damaged ragweed in the laboratory caused a drop in oviposition intensity, and a considerable proportion of females completely stopped oviposition. Feeding on slightly damaged ragweed had no significant effect on oviposition intensity. Ovipositing females preferred to feed on the intact ragweed and lay their eggs close to it. The locomotory activity of ovipositing females was significantly higher on highly damaged ragweed whereas non-ovipositing (diapausing) females and males were behaviourally indifferent to the extent of ragweed damage. Under natural conditions, ovipositing females more frequently left damaged host plants for less damaged one. If the degree of ragweed damage is high over a large area, the insects that were unable to find undamaged plants for several days oviposited less and some females entered diapause. The adaptive effect of these reactions is a decrease of population density in advance, before it might drop as a result of starvation. These results are in agreement with the second model of insect reaction to the damage-induced changes in a host plant (Edwards and Wratten 1987).     

Natural parasitism of Metaparasitylenchus hypothenemi (Tylenchida: Allantonematidae) on the coffee berry borer in Chiapas,Mexico

We assessed the parasitism of Metaparasitylenchus hypothenemi on its host, the coffee berry borer, in 20 coffee plantations of Mexico. A total of 23,568 adult borers were dissected, with 179 of these infected with nematodes (0.76% infection rate). Although the level of parasitism is not encouraging, the nematode is another natural enemy, which limits the population growth of this pest in Mexico.