Trade-offs underlying polyphagy in a facultative ant-tended florivorous butterfly: the role of host plant quality and enemy-free space

The underlying mechanisms mediating the use of multiple host plants were investigated in Parrhasius polibetes (Lycaenidae), a florivorous and facultative myrmecophilous butterfly. Plant traits such as presence of ant–treehopper associations as a source of enemy-free space, flower bud dimensions, toughness, thickness, trichomes, and the corresponding performance and wear of P. polibetes mandibles were examined for three natural hosts: Schefflera vinosa (Araliaceae), Pyrostegia venusta (Bignoniaceae) and Luehea grandiflora (Malvaceae). Parasitism levels of larvae found on the three hosts were also determined. Almost all Luehea had ant–treehopper associations, and all larvae found on this host were non-parasitized. Parasitism was low in larvae found on Schefflera, half of which hosted ant–treehopper associations. No ant–treehopper association was found on Pyrostegia, where parasitism was significantly higher compared to other hosts. In the laboratory, P. polibetes performed well on Schefflera, followed by Pyrostegia. No larvae survived when fed with Luehea. Flower buds of Luehea were thicker and tougher than those of Schefflera and Pyrostegia. Indeed, mandibles of larvae reared on Luehea showed substantial wear, whereas those reared either on Schefflera or Pyrostegia presented no significant damage. Additionally, we suggest that co-occurrence with ant–treehopper associations on a plant provides parasitoid-free space for P. polibetes larvae. Our results support the hypothesis that ecological trade-offs among host plants (i.e., food quality and enemy-free space) promote polyphagy in natural populations of P. polibetes. Host morphological traits seem to play a relevant role in P. polibetes performance. To our knowledge, this is the first report showing the costs of polyphagy in a myrmecophilous butterfly.     

Biosteres longicaudatus: Developmental dependence on host (Anastrepha suspensa) physiology

Larvae of Anastrepha suspensa that were in the first day of the third instar were parasitized by females of the solitary endoparasitoid, Biosteres longicaudatus. At the end of the 6-hr oviposition period, larvae were ligated posterior to the ring gland so that some larvae had parasitoids anterior to the ligature while in others, the parasitoids were in the abdomen, posterior to the ligature. Ninety-two percent of the parasitoids anterior to the ligature hatched to the first through third instars. Parasitoids posterior to the ligature had a 75% egg hatch to the first instar only. No larval molts to the second or subsequent instars occurred in these parasitoids. Upon parabiosis to 3-day-old, unparasitized host pupae, the ligated larvae pupated and 97% of the first-instar parasitoids in these parabiosed larval abdomens molted to the second instar. Newly laid parasitoid eggs transplanted to 3-day-old pupal hosts had less than one-third of the egg hatch of those transplanted to first-day third-instar hosts. The data implicate the physiological state of the host (vis-a-vis pupation and associated events) as being an important factor in the development of the endoparasitoid.     

Protection of potato crop against Phthorimaea operculella (Zeller) infestation using frass extract of two noctuid insect pests under laboratory and storage simulation conditions

The oviposition deterrent effect of water extract of Spodoptera littoralis and Agrotis ipsilon larval frass on Phthorimaea operculella adult females was studied using two types of larval food “Natural host and Semi-artificial diet” under laboratory and storage simulation (semi-field) conditions. Extracted frass of fed larvae on semi-artificial diet showed complete oviposition deterrent effect at treatments with 4th, 5th and 6th instars of S. littoralis, also at treatments with 1st–3rd and 6th instars of A. ipsilon, while the same effect was observed when the larvae fed on castor oil leaves as a natural host only at treatment with frass extract of A. ipsilon 6th instar larvae. Presence of low amounts of phenols and flavonoids in water extract of A. ipsilon larval frass resulted in relatively more effect as oviposition deterrent to fertile adult females on treated oviposition sites, while the opposite effect was obtained in S. littoralis larval frass experiments. At semi-field experiments, the percentage reduction of laid eggs reached 100% after two?days at treatments with frass extracts of 4th and 5th S. littoralis larval instars and A. ipsilon 6th instar larvae fed on semi-artificial diet and/or castor oil leaves. Percentage reduction of laid eggs for untreated sacks reached 93.24 and 48.95% after 2 and 30?days, respectively, when placed between treated sacks, in comparison with the mean number of laid eggs for isolated control.     

Oviposition decisions in an endoparasitoid under self-superparasitism conditions

Superparasitism is a widespread phenomenon. Having accepted superparasitism, mated female parasitoids must decide on the sex of each egg they subsequently lay into the same host. Theory predicts that this decision is either based on host quality, when more male eggs are laid in hosts that are already parasitized because they are perceived to be of poorer quality; or more eggs are laid of the sex that is most likely to be a strong larval competitor, i.e. generally females.Anastatus disparis is a facultative endoparasitic egg parasitoid. We used ‘artificial’ hosts to explore outcomes of decision making by A. disparis during superparasitism under a manipulated absence of larval competition. When only one egg was laid it was always female. As the number of eggs laid increased, so more of them were male. This supports the theory that oviposition decisions are based on host quality; more male eggs were laid in hosts that were already parasitized and thus of poorer quality.In a second experiment, eggs were exposed to parasitoids for different periods of time. Half the eggs were dissected to determine the number of parasitoid eggs that had been laid. The remaining eggs were incubated and the number and sex of offspring that ultimately emerged, following larval competition, were recorded. Under superparasitism conditions fierce larval competition ensued; only one offspring survived and they were predominantly female.In conclusion, oviposition decisions by female A. disparis accepting self-superparasitism were made based on host quality.     

萧氏松茎象的生活史、产卵和取食习性

萧氏松茎象Hylobitelus xiaoi是严重蛀害国外松的新害虫。在江西赣南地区两年发生一代,以幼虫在蛀道、成虫在蛹室或土中越冬。卵和蛹的发育起点温度分别为8.4℃和7.5℃,有效积温分别为215.9和345.3日·度。各虫态历期：13～28℃间,卵为52.9～11.4天;13～30℃间,蛹为60.1～16.4天; 25℃恒温下,幼虫128.9天。成虫靠爬行活动,极少飞翔。成虫具夜出性活动节律：即傍晚上树行取食、交配和扩散等活动,早晨回到树干基部或土缝中。成虫需取食松枝作为补充营养,产卵前期46.3天,产卵期 105.3天,卵产于近表土的寄主树皮内,每雌产卵35.7粒。幼虫5～7龄,以幼树危害最烈。     

Performance of Microplitis tuberculifer (Hymenoptera: Braconidae) parasitizing Mythimna separata (Lepidoptera: Noctuidae) in different larval instars

The solitary parasitoid Microplitis tuberculifer (Wesmael) is an important biological control agent of various lepidopteran pests in Asia. We examined the preference of M. tuberculifer for different instars of its common host, Mythimna separata (Walker), host instar effects on parasitoid development, and the consequences of parasitism in different stages for growth and consumption of host larvae. The wasp successfully parasitized the first four larval instars of M. separata, but not the fifth, which appeared to be behaviorally resistant. First and second instars were parasitized at higher rates compared to thirds and fourths in no-choice situations, ostensibly due to longer handling times for the latter, but second instars were most preferred in a choice test that presented all stages simultaneously. Although later instar hosts yielded heavier cocoons, the fastest parasitoid development was obtained in second instars. Lower sex ratios were obtained from first instars as females appeared to lay a smaller proportion of fertilized eggs in small hosts. Both weight gain and food consumption of parasitized larvae were reduced significantly within 24 h of parasitism, regardless of the stage parasitized, and final body weights were less than 10% those of unparasitized larvae. Thus, M. tuberculifer has good potential as a biological control agent of M. separata, successfully parasitizing the first four larval instars and dramatically reducing plant consumption by the host in all cases.     

Does superparasitism improve host suitability for parasitoid development? A case study in the Microplitis rufiventris-Spodoptera littoralis system

Superparasitism refers to the oviposition behavior of parasitoid females who lay their eggs in an already parasitized host. Recent studies have shown that allocation of additional eggs to an already parasitized host may be beneficial under certain conditions. In the present work, mortality of Microplitis rufiventris wasps was significantly influenced by both host instar of Spodoptera littoralis larvae at parasitism and level of parasitism. In single parasitization, all host instars (first through sixth) were not equally suitable. Percentage of emergence success of wasp larvae was very high in parasitized first through third (highly suitable hosts), fell to 60% in the fourth instar (moderate suitable) and sharply decreased in the penultimate (5th) instars (marginally suitable). Singly parasitized sixth (last) instar hosts produced no wasp larvae (entirely unsuitable), pupated and eclosed to apparently normal adult moths. The scenario was different under superparasitism, whereas supernumerary individuals in the highly suitable hosts were almost always killed as first instars, superparasitization in unsuitable hosts (4th through 6th) had significant increase in number of emergence success of wasp larvae. Also, significantly greater number of parasitoid larvae successfully developed in unsuitable hosts containing three wasp eggs than counterparts containing two wasp eggs. Moreover, the development of surplus wasp larvae was siblicidal in earlier instars and nonsiblicidal gregarious one in the penultimate and last “sixth” instars. It is suggested that the optimal way for M. rufiventris to deal with high quality hosts (early instars) is to lay a single egg, while the optimal way to deal with low quality hosts (late instars) might be to superparasitize these hosts.     

Isolation of corn stunt spiroplasma in Jamaica, and probable dual infection with maize mosaic virus

Females of Cydia ptychora oviposited on cowpea, Vigna unguiculata ssp. unguiculata in the field between 18.00 and 19.00 h commencing about 30 days after planting with a few eggs laid on the leafbuds and the undersides of leaves. Oviposition increased as the host plant developed the preferred sites, floral sepals and their remains on pods, and peaked at about 58 days after planting regardless of planting date. First instar larvae were found wandering on the plant between 06.45 and 11.15 h but other larval instars were rarely encountered outside the pod except when changing pods or leaving pods to pupate in the soil. Contrary to previously reported observations, larval development occurred in leafbuds and inside flowers both in the field and in laboratory rearing. Development was optimal on pods, and mortality was high in leafbuds.     

The effect of a foliar disease (rust) on the development of Gastrophysa viridula (Coleoptera: Chrysomelidae)

Abstract.

• 1 Gastrophysa viridula Degeer (Coleoptera: Chrysomelidae) and the pathogenic rust fungus Uromyces rumicis (Schum.) Wint. both occur on leaves of Rumex crispus L. and R.obtusifolius L. Individual stages of beetle development, and egg laying, were compared on healthy and infected leaves of each plant species in the laboratory. Oviposition choice was investigated in the field and laboratory.
• 2 Beetles reared on infected leaves of each species had greater larval mortality and slower development than those reared on healthy leaves. Although larvae feeding on infected leaves consumed up to 2.5 times more dry weight than those reared on healthy leaves, they had a lower relative growth rate and pupated at a lower weight. These changes were consistent with the reduced nutritive quality of rust-infected Rumex leaves.
• 3 Fecundity of beetles reared on infected leaves of both species was considerably reduced. Eggs laid by beetles feeding on infected R.crispus leaves also had a reduced viability.
• 4 The beetle developed consistently poorer on healthy R.crispus than on healthy R.obtusifolius throughout its life-cycle. Differences in larval performance were greater between host species than between infected and healthy leaves.
• 5 Oviposition was similar on infected and healthy R.crispus in both the laboratory and field. However, adults consumed less, and laid fewer eggs on infected than on healthy R.obtusifolius. The pattern of egg laying on different aged leaves was affected by rust infection: a greater proportion of eggs was laid on the older, infected leaves, than on the equivalent aged leaves on the healthy plants. Few larvae survived from eggs laid on rusted leaves in the field.

     

Reciprocal influences and costs of parasitism on the development of Corcyra cephalonica and its endoparasitoid Venturia canescens

Many endoparasitoids develop successfully within a range of host instars. Parasitoid survival is highest when parasitism is initiated in earlier host instars, due to age-related changes in internal (physiological) host defences. Most studies examining fitness-related costs associated with differences in host instar have concentrated on the parasitoid, ignoring the effects of parasitism on the development of surviving hosts that have encapsulated parasitoid eggs. A laboratory experiment was undertaken examining fitness-related costs associated with encapsulation of Venturia canescens (Hymenoptera: Ichneumonidae) eggs by fifth (L5) instar larvae of Corcyra cephalonica (Lepidoptera: Pyralidae). Growth and development of both host and parasitoid were monitored in C. cephalonica larvae containing 0, 1, 2, or 4 parasitoid eggs. Adult size and fecundity of C. cephalonica did not vary with the number of eggs per host. However, there was a distinct increase in host mortality with egg number, although most parasitoids emerged from hosts containing a single egg. The most dramatic effect on the host was a highly significant increase in development time from parasitism to adult eclosion, with hosts containing 4 parasitoid eggs taking over 2.5 days longer to complete development than unparasitized larvae. The egg-to-adult development time and size of adult V. canescens did not vary with egg number per host, as demonstrated in a previous experiment using a different host (Plodia interpunctella). The results described here show that there are fitness-related costs to the host associated with resistance to parasitism.     

Host plant, oviposition behavior and larval ecology of a sawfly leafminer, Profenusa japonica (Hymenoptera: Tenthredinidae)

The host plant, oviposition behavior and larval ecology of Profenusa japonica Togashi are reported for the first time. Adults of P. japonica mated and oviposited on a polyantha rose, Rosa multiflora (Rosaceae), in April. Each female adult laid an egg on the edge of a leaflet. Hatched larvae consumed the parenchymatous layer of leaflets and in so doing created a mine. The larval stage comprised five instars. On average, 70.6% of the total area of a leaflet was consumed by one larva. Female adults of P. japonica laid eggs singly, probably to avoid larval competition for food.     

Host plant affects pollen beetle (Meligethes aeneus) egg size

Abstract. In some herbivorous insect species, egg size is larger on low‐quality hosts than on high‐quality hosts and may be related to the prospect that larger offspring are more likely to survive on a poor host. Sizes of eggs laid by pollen beetles [Meligethes aeneus Fab. (Coleoptera: Nitidulidae)] were examined with insects confined on one of two different host plants that had previously shown differences in adult preference and larval performance. Individual females were also exposed sequentially to both the low‐quality host (Sinapis alba L.) and the high‐quality host (Brassica napus L.) and the size of their eggs was determined. Pollen beetles laid shorter eggs on low‐quality hosts both for different females on different host plants and for the same individuals on different host plants, in contrast to the prediction that low‐quality hosts would receive larger eggs than high‐quality hosts. Previously, egg production rate was shown to be reduced when pollen beetles are exposed to low‐quality hosts and it is suggested that oogenesis is incomplete, resulting in shorter eggs. The possibility that this is related to antibiosis on S. alba is discussed.     

Larval host plant affects fitness consequences of egg size variation in the seed beetle Stator limbatus

Egg size variation often has large effects on the fitness of progeny in insects. However, many studies have been unable to detect an advantage of developing from large eggs, suggesting that egg size variation has implications for offspring performance only under adverse conditions, such as during larval competition, periods of starvation, desiccation, or when larvae feed on low-quality resources. We test this hypothesis by examining the consequences of egg size variation for survivorship and development of a seed-feeding insect, Stator limbatus, on both a low-quality (Cercidium floridum) and a high-quality (Acacia greggii) host plant. Our results are consistent with the hypothesis. S. limbatus larval performance was affected by egg size only when developing on the poor-quality host (C. floridum); larvae from large eggs survived better on C. floridum than those from small eggs, while there was no evidence of an effect of egg size on progeny development time, body weight, or survivorship when larvae developed on A. greggii. These results indicate intense selection for large eggs within C. floridum-associated populations, but not in A. greggii-associated populations, so that egg size is predicted to vary among populations associated with different hosts. Our results also support this hypothesis; females from a C. floridum-associated population (Scottsdale) laid larger eggs than females from an A. greggii-associated population (Black Canyon City).     

Reproductive Biology of Celatoria compressa (Diptera: Tachinidae), a Parasitoid of Diabrotica virgifera virgifera (Coleoptera: Chrysomelidae)

The tachinid Celatoria compressa, a parasitoid of adult Diabrotica species in North America has been studied as a candidate classical biological control agent for the western corn rootworm, Diabrotica virgifera virgifera, in Europe. Prior to its potential importation, a thorough understanding of the parasitoid's reproductive biology is essential, and is an important component in the evaluation of a species as a biological control agent. In this study it has been clarified that C. compressa belongs to a group of a few tachinid species characterised by having eggs that contain fully developed larvae which are laid directly into the host. After mating, the egg load of females increased steadily from day 1 to a maximum egg load on day 4. Thereafter eggs containing fully developed first instar larvae reached a maximum of 31 in 69 eggs. At the first day of larviposition, females laid on average only five eggs into multiple hosts, which is in contrast to the availability of 18 eggs containing fully developed first instars in the uterus per female at that time. During a mean female's larviposition period of 23 days, a total of 33 first instars were larviposited into the hosts, which is only half of the female's egg load. Lifetime fecundity of C. compressa was significantly correlated with longevity. However no relationship was found between body size and either lifetime fecundity or longevity. In this study, an inverse host density-dependent pattern of percent parasitism was shown for C. compressa under 24-h fixed-time laboratory conditions, reflecting a Holling type II response. The number of host parasitized per C. compressa female reached an upper limit of 10 hosts with an increasing host density, which can be explained by the long host handling time of C. compressa.     

Evaluation of tebufenozide carry-over and residual effects on spruce budworm (Lepidoptera: Tortricidae)

Laboratory and field studies investigated carry-over effects of tebufenozide on spruce budworm, Choristoneura fumiferana (Clemens). In the laboratory, third and fourth instars were fed either sublethal doses of tebufenozide (10 ppm) or water onAbies balsamea (L.) Mill. needles, reared to adulthood and allowed to oviposit on laying surfaces 1 or 14 d after being sprayed with water or tebufenozide concentrations of 17.5, 35.0, and 70.0 g/liter. Percentage adult emergence and sex ratio were not affected by larval ingestion of the tebufenozide. Also, the mean number of eggs laid on untreated wax paper by moths reared from tebufenozide-treated larvae was similar to the controls. Hence, tebufenozide did not inhibit carry-over effects on treated larvae. Oviposition on tebufenozide-treated wax paper by moths reared from untreated larvae was affected by both the substrate concentration and the age of the treatment residue. When offered treated and untreated laying surfaces simultaneously, C. fumiferana did not show a preference. However, significantly fewer eggs were laid on both laying surfaces by fewer females than when tebufenozide was absent. Residual tebufenozide on wax paper did not affect egg hatch but topical applications were toxic to eggs. Field studies appear to corroborate laboratory results and suggest that although the ingestion of tebufenozide by larval spruce budworm might not impair adult reproduction, the insecticide's presence in the environment could inhibit oviposition. This inhibition was considered to be a primary factor in tebufenozide's multi-year effects against spruce budworm populations.     

Life history parameters of the microtype parasitoid fly Pales pavida (Diptera: Tachinidae), and effects of host age and number of eggs ingested by host on parasitism success

With the aim of developing better procedures for rearing the microtype tachinid fly Pales pavida (Meigen), we performed ecological studies in the laboratory using the natural host Mythimna separata (Walker), investigating larval development, mating behaviour, individual oviposition patterns and relationships between parasitisation and number of eggs ingested (NEI) per host. The host instar at the time of parasitoid egg ingestion significantly affected the development time of the immature parasitoid: development took longer when the hosts ingested eggs when at the fifth instar than at the sixth (last) instar. There was no difficulty obtaining mated females in the laboratory when day 0–1 female flies were kept with day 2–4 males. Mean lifetime fecundity was 5805?±?568 eggs per female. Daily rates of oviposition by individual females varied widely; the greatest number of eggs laid in a day was exactly 1700. When the NEI by day 1 fifth instars or day 0 or 3 last instars was 1, 3, 6 or 10, the parasitisation percentage tended to increase with increasing NEI, although it did not differ significantly between NEI 6 and 10. Therefore, the percentage adult emergence per egg ingested decreased from NEI 6 to 10, particularly in the case of last instars. Using day 0 last instars, with six eggs ingested per host, should increase parasitisation rates and shorten the development time of the parasitoid for rearing.     

Larvae of the Japanese termitophilous predator Isoscelipteron okamotonis (Neuroptera,Berothidae) use their mandibles and silk web to prey on termites

Several species of Berothidae are known to be specialist predators of termites during their larval stages. A female of the Japanese berothid Isoscelipteron okamotonis (Nakahara, 1914) was captured in the field and laid eggs in captivity. Of 67 eggs, 20 hatched into larvae, 3 of 9 which pupated and 1 of which reached the adult stage. The first and third instars preyed on workers of the termite Reticulitermes speratus, although the second instars did not. Larvae did not show the specific predatory behaviors observed in Northern American relatives. Instead, they bound paralyzed termites with silk that was apparently used for hunting. This is the first record of an Old World berothid raised from egg to adult in the laboratory.     

Biology, rearing and field release on Guam of Euplectrus maternus, a parasitoid of the fruit-piercing moth, Eudocima fullonia

The eulophid parasitoid, Euplectrus maternus, is a gregarious ectoparasitoid of the larvae of thefruit-piercing moth, Eudocima (Othreis) fullonia. This parasitoid is indigenous to India and was released into Guam in 1998–99 to aid in the biological control of E. fullonia, an important pest of ripeguava, banana, mango, pomegranate, litchi,papaya, tomato, orange and other fruit crops.In the laboratory, biology and rearing of E. maternus on the larvae of E. fulloniawere studied. In no-choice tests, ovipositionby E. maternus was significantly greateron first and second instar larvae than on thirdinstar larvae. However, in free-choice tests,oviposition by E. maternus wassignificantly greater on second instar larvae,followed by first and third instar larvae. Thisparasitoid did not lay eggs on later instars,either in no-choice or free-choice tests.Females laid a significantly greater number ofeggs (>80%) on the dorsal surface of thelarvae than on either lateral or ventralsurfaces (1 to 8%). In general, more eggs werelaid on abdominal segments 1–3 on larvalinstars one to three. The survival of theimmature stages (from egg to adult) of E. maternus on second instar E. fullonialarvae was 61%, which was significantlygreater than the survival rates on first andthird instar larvae (32% and 26%), indicatingthat second instar host larvae are ideal formass rearing of the parasitoid. Mated femaleE. maternus continuously laid eggs on thesecond instar larvae of E. fullonia forup to 30 days, but the greatest number of eggswere laid during the first week after exposure.When E. maternus laid more than two eggson host larva, more female progeny wereproduced, indicating female-biasedreproduction. Adult parasitoids lived longerwhen fed with a honey: water (50% w/v)solution than with pure honey. Similarly, thefecundity of females increased significantlywhen fed with the honey-water solution whencompared to feeding with pure honey. To date,E. maternus has not established in Guam.     

Effect of host diet on acceptance of eastern spruce budworm eggs byTrichogramma minutum

The acceptance behaviour of the parasitic wasp, Trichogramma minutum Riley (Hymenoptera: Trichogrammatidae), was studied on two types of egg masses of eastern spruce budworm (SBW), Choristoneura fumiferana Clemens. Host eggs were obtained from moths that had been fed balsam fir or artificial diet during their larval stages. Parasitoids were reared from both types of hosts. Host acceptance assays were conducted where females were provided with a single egg mass and with a choice between egg masses. Regardless of the parasitoid's rearing host, females parasitised hosts reared from artificial diet more often than those from balsam fir. Response was consistent in both no-choice and choice assays. Rejection of hosts tended to occur during the examining stage of behaviour, suggesting that acceptability is judged externally. If a host was accepted, the same number of eggs were laid per egg mass regardless of treatment.