Temperature-dependent development of diapausing larvae of Chilo partellus (Swinhoe) (Lepidoptera: Crambidae)

Temperature-dependent development rate, percent diapause induction (hibernation at low temperature and aestivation at high temperature), and survival of diapausing larvae of Chilo partellus (Swinhoe, 1885) were examined on 13 constant temperatures ranging from 8 to 40 °C. Development of hibernating and aestivating larvae occurred from 10 to 25 °C and 27–38 °C, respectively. However, no development occurred at 8 °C and 40 °C. To determine actual thermal conditions that affect development and trigger both kind of diapause (hibernation and aestivation), various thermal parameters were estimated by fitting the development rate data to two linear (Ordinary equation and Ikemoto & Takai) models and thirteen non-linear models. The lower thermal thresholds (T_min) for development of diapausing larvae of C. partellus were calculated as 9.60 °C and 10.29 °C using the ordinary linear model and Ikemoto & Takai model, respectively. Similarly, the thermal constants (K) estimated using the ordinary linear model was 333.33 degree-days and that estimated with Ikemoto & Takai model was 338.92 degree-days. Among the non-linear models, Lactin-2 followed by Lactin-1 were found to be the best as these models estimated the critical temperatures (T_min, T_max and T_opt) similar to those of observed values. Conclusively, the Ikemoto & Takai linear model and Lactin-2 followed by Lactin-1 non-linear models are useful and efficient for describing temperature-dependent development and estimating the temperature thresholds of diapausing larvae of C. partellus. Our findings provided fundamental information for estimation of thermal requirement and temperature based development models for diapausing larvae of C. partellus. This information will be highly useful for predicting the occurrence, seasonal emergence, number of generations and population dynamics of C. partellus.     

Yield loss due to the stemborer Chilo partellus (Swinhoe) (Lepidoptera: Crambidae) at different nitrogen application rates to maize

Field trials were conducted at Kibaha and Morogoro in eastern Tanzania during two seasons to evaluate the effect of nitrogen fertilization (0, 50, 75, 100 kg [N]/ha) on pest abundance, plant damage and yield loss of maize due to stemborers. In general, ear and grain weights increased linearly with nitrogen level. In the infested plot, grain weight increased 2.5 and 1.8 fold from 0 to 100 kg [N]/ha in the short and long rainy season, respectively, at Kibaha, and 1.4 and 1.6 times at Morogoro. Yield loss decreased with an increase in nitrogen application and the effect was stronger under high than low borer infestation levels. The results show the beneficial effect of nitrogen on the plant’s ability to compensate for borer damage. Analysis of economic benefits of applying fertilizer and insecticide treatment indicated that using insecticides is not profitable under high-pest-low-soil fertility conditions.     

Chilo partellus (Swinhoe) (Lepidoptera: Pyralidae) a new invasive species in Israel

Chilo partellus, the spotted stem borer (SSB), is reported for the first time from the fields of sorghum and corn in Israel. Spotted stem borer larvae were first discovered in July 2010 on sorghum and corn plants in the Western Galilee. The new pest had spread all over northern Israel by the end of 2011. The identity of the new pest was confirmed using the morphology of the dissected male genitalia and its attraction to SSB sex pheromone.     

Effect of learning on the oviposition preference of field-collected and laboratory-reared Chilo partellus (Lepidoptera: Crambidae) populations

Recent studies show that Vetiver grass, (Vetiveria zizanioides (L.) Nash), may have potential as a dead-end trap crop in an overall habitat management strategy for the spotted stem borer, Chilo partellus (Swinhoe) (Lepidoptera: Crambidae). Vetiver grass is highly preferred for oviposition, in spite of the fact that larval survival is extremely low on this grass. The oviposition behaviour of female Chilo partellus moths was investigated by determining the amount and size of egg batches allocated to maize and Vetiver plants and studying the effect of rearing conditions and oviposition experience on host plant selection. Two-choice preference tests were used to examine the effect of experience of maize (a suitable host plant) and Vetiver plants on the oviposition choice of C. partellus. For both field-collected and laboratory-reared moths, no significant differences were found in the preference distributions between the experienced groups. It is concluded that females do not learn, i.e. that they do not change their preference for Vetiver grass after having experienced oviposition on either maize or this grass, which supports the idea that trap cropping could have potential as a control method for C. partellus. Differences observed between field-collected and laboratory-reared moths in the amount and size of egg batches laid on maize and Vetiver grass indicate that data obtained from experiments with laboratory-reared insects should be treated with caution.     

Dominance of spotted stemborer Chilo partellus Swinhoe (Lepidoptera: Crambidae) over indigenous stemborer species in Africa's changing climates: ecological and thermal biology perspectives

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Invasive maize spotted stem borer Chilo partellus Swinhoe 1885 (Lepidoptera: Crambidae) is the dominant pest in maize agro-system in the Mediterranean Region of Turkey

The maize spotted stem borer Chilo partellus Swinhoe 1885 (Lepidoptera: Crambidae) was recorded for the first time in Turkey in 2014. This pest is described as invasive, especially in the current climate change dispensation. It is important to investigate the population dynamics of C. partellus for better management practices. We studied the species abundance distribution (SAD) models and some biodiversity indices of three major stem bores, C. partellus, Ostrinia nubilalis and Sesamia nonagrioides as these are the most common and major stem borers of maize in the Mediterranean Region. With the help of light traps, weekly catches of the three major maize stem borers were collected during two maize growing seasons (first cropping and second cropping seasons) in 2018 and 2019. The results showed that C. partellus emerged earlier (April-May) and was very dominant throughout the first cropping season of maize. Ostrinia nubilalis and S. nonagrioides emerged late in June-July, and its population remained low throughout the first cropping season. The population structure of all three stem borers was similar throughout the second cropping season of maize. The relative abundance (RA) of C. partellus was significantly higher than that of O. nubilalis and S. nonagrioides in the first cropping (p <.0001) when compared to the second cropping season (p >.05) for both years. In the first cropping season for both years, no SAD model fitted the observed species distribution, although Fisher’s log series was relatively similar. In the second cropping for both years, the geometric series distribution model fitted the observed SAD. The Berger-Parker dominance index was greater for the first cropping season than that of the second, whereas the opposite was true for the Simpson’s evenness and evenness of Pielou. High RA of C. partellus and the geometric series distribution observed in the second cropping are indicative of a possible dominant status of the C. partellus in the Mediterranean Region of Turkey. Our findings indicated that seasonality is a fundamental driving factor influencing the distribution of the three major stem borers of maize in the Mediterranean Region. In addition, the long and warmer winters could be the reason for the dominance of C. partellus as indicated by the diversity indices.     

Release and establishment of Cotesia flavipes Cameron (Hymenoptera: Braconidae) an exotic parasitoid of Chilo partellus (Swinhoe) (Lepidoptera: Crambidae) in East and Southern Africa

Cotesia flavipes Cameron (Hymenoptera: Braconidae) was first imported into Kenya in 1991 from Pakistan for control of Chilo partellus (Swinhoe) (Lepidoptera: Crambidae). First releases were made at the Kenya coast in 1993 and a few recoveries of the parasitoid were made the following year. Additional foreign exploration for C. flavipes was conducted in central India in 1996 and 1998, which resulted in additional importation of the parasitoid for subsequent releases in eastern and southern Africa. Region-wide releases commenced with releases in Mozambique in 1996, Somalia in 1997 and Uganda in 1998. By 2005, many releases had been made in 10 countries in East and Southern Africa with establishment being reported in all of them except Eritrea but including Ethiopia where releases had never been made. This paper describes the progress made in the release and establishment of C. flavipes in East and Southern Africa and quantifies the rate of spread from the initial release sites in Kenya.     

Evaluation of Beauveria bassiana and Metarhizium anisopliae for Controlling Chilo partellus (Lepidoptera: Crambidae) in Maize

Maize (variety Katumani) was planted in a greenhouse and plants were infested with 20 Chilo partellus second instar larvae 3 and 4 weeks after plant emergence. One isolate of Beauveria bassiana (BB-01) and four isolates of Metarhizium anisopliae (PPRC-4, PPRC-19, PPRC-61 and EE-01) were sprayed onto the leaf whorl at 2×10⁸ conidia/mL 24 h after infestation. Leaf damage by the larvae was greatly reduced by the treatments. The mean daily temperature and relative humidity in the greenhouse ranged from 10 to 35°C and 30 to 90%, respectively. The growth of infested unsprayed plants was less than that of fungi treated plants. Stem tunneling (1-5%), deadheart (0-33%), number of attacked nodes (0.3-2.5) and holes (0.2-3.3) were also reduced in plants sprayed with conidial suspensions. Isolates PPRC-4, PPRC-19 and PPRC-61 seemed to be the best candidates for further development.     

Consequences of diapause on post‐diapause development,reproductive physiology and population growth of Chilo partellus (Swinhoe)

We investigate the effects of diapause on post‐diapause development, reproductive physiology and population growth of Chilo partellus (Swinhoe) (Crambidae: Lepidoptera). Aestivating and hibernating larvae of C. partellus are exposed to diapause terminating conditions (consisting of an LD 12 : 12 h photocycle at 27 ± 1 °C and 65 ± 5% relative humidity with a fresh diet) to terminate the diapause and observations are made on percentage pupation, pupal duration and weight, adult reproductive performance and population growth parameters. We find that the diapause in C. partellus significantly reduces the percentage pupation and weights of pupae, ultimately lowering the weight and reproductive performance of adults. Reduced weights of adult females are found to be directly associated with a lower number of egg cells in ovaries. Nevertheless, the reproductive performance of C. partellus males is also found to be greatly affected in the diapause (hibernation and aestivation) experiencing population in terms of the deposition of a lower number of spermatophores and eupyrene sperm in the reproductive tracts of females compared with the nondiapausing population. The results of the present study clearly indicate that a reduction in longevity, fecundity and egg viability, as well as a reduced rate of deposition of spermatophores and eupyrene sperm, in a diapause experiencing population of C. partellus ultimately leads to a reduction in population growth parameters, thus having implications for bio‐ecology and population dynamics under a changing climatic scenario.     

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.     

Complete mitochondrial genome of Chilo suppressalis (Walker) (Lepidoptera: Crambidae)

The complete sequence of the mitochondrial genome (mitogenome) of the rice stem borer Chilo suppressalis (Walker) (Lepidoptera: Crambidae) was determined to be 15,465 bp. It contains 13 protein-coding genes (PCGs), 22 tRNA genes, the large and small rRNA genes, and an A+T-rich region. The nucleotide composition of the mitogenome of C. suppressalis is highly A+T biased, accounting for 79.70% in whole mitogenome, 77.74% in PCGs, 84.70% in tRNAs, 81.20% in rRNAs and 94.19% in A+T-rich region, respectively. The PCGs have typical ATN start codons, except for cox1, which contains the unusual CGA. The C. suppressalis A+T-rich region contains a conserved structure combining the motif ATAGA and a 19-bp poly-T stretch, but absence of the 9-bp poly-A element upstream trnM.     

Survival and development of maize stem borer Chilo Partellus (Swinhoe) Lepidoptera: Crambidae on artificial diet

The Life cycle of maize stem borer, Chilo partellus (Swinhoe) was studied in in vitro conditions. Development of stem borer undergoes following stages like egg, larvae, pupa and moth. The egg incubation period ranged from 3 to 6 days, larval stage was observed in five instars. The mean value of I, II, III, IV and V instars showed 3.8 ± 0.16, 5.2 ± 0.02, 6.1 ± 0.06, 7.35 ± 1.5, and 10.12 ± 0.29 days, respectively and complete larvae period ranged from 42 to 49 days. Pupae stage was observed in 8–9 days. The pre-mating and mating period was found at 9.10 ± 1.20 and 5.14 ± 1.08 h while egg laying period in 4.1 ± 1.32 days respectively. Fecundity rate of stem borer is from 262 to 657 eggs. The life span of adult male (3-7) and female (3-8) days was observed with a mean of 6.30 ± 0.85 and 5.10 ± 0.69 days respectively. Life cycle of stem borer gets completed in 47 to 51 days. Development of quality insects in required quantities at different developmental stages and their timely supply plays an inevitable role particularly for insect-breeding resistant programs. Hence to meet these challenges we had tried to standardize an artificial diet with cost effective to rear Chilo partellus under in vitro conditions.     

Laboratory evaluation of the virulence of Beauveria bassiana isolates to the sorghum shoot borer Chilo partellus Swinhoe (Lepidoptera: Pyralidae) and their characterization by RAPD-PCR

Beauveria bassiana has long been used as a mycopesticide. It has a wide host range; isolates have been reported to differ in host range and virulence to a given insect species. Identification of a molecular marker linked to a virulent phenotype to a target pest would be useful in screening for isolates effective against it. Twenty B. bassiana isolates were tested for their virulence to the second instar larvae of Chilo partellus Swinhoe in laboratory bioassays and their DNA fingerprints were generated by RAPD-PCR. Three arbitrary categories of aggressiveness were chosen; isolates that caused >70%, between 70 and 40% and <40% larval mortality were grouped as highly, medium and less aggressive types, respectively. In the random amplified polymorphic DNA (RAPD) analysis a 30% variability was observed among the isolates; which clustered into three major groups. The groups based on virulence rating did not match with the RAPD clusters. One of the highly aggressive isolates clustered with less aggressive isolates in one cluster and the other grouped along with the medium aggressive isolates in a different cluster. The B. bassiana isolates were classified phenotypically based on the taxonomic order of the original insect host and the climatic zone (tropical/temperate) from which they were isolated. No correlation between the aggressiveness of the isolate and the relatedness of the original insect host to the tested insect was observed; both the highly aggressive isolates were from coleopteran insects. A correlation was found between the RAPD grouping and the phenotypic classification of the isolates. All the lepidopteran isolates grouped into one major cluster, most sub clusters were constituted by isolates from the same climatic zone.     

Comparative assessment of the thermal tolerance of spotted stemborer,Chilo partellus (Lepidoptera: Crambidae) and its larval parasitoid,Cotesia sesamiae (Hymenoptera: Braconidae)

Under stressful thermal environments, insects adjust their behavior and physiology to maintain key life‐history activities and improve survival. For interacting species, mutual or antagonistic, thermal stress may affect the participants in differing ways, which may then affect the outcome of the ecological relationship. In agroecosystems, this may be the fate of relationships between insect pests and their antagonistic parasitoids under acute and chronic thermal variability. Against this background, we investigated the thermal tolerance of different developmental stages of Chilo partellus Swinhoe (Lepidoptera: Crambidae) and its larval parasitoid, Cotesia sesamiae Cameron (Hymenoptera: Braconidae) using both dynamic and static protocols. When exposed for 2 h to a static temperature, lower lethal temperatures ranged from ?9 to 6 °C, ?14 to ?2 °C, and ?1 to 4 °C while upper lethal temperatures ranged from 37 to 48 °C, 41 to 49 °C, and 36 to 39 °C for C. partellus eggs, larvae, and C. sesamiae adults, respectively. Faster heating rates improved critical thermal maxima (CT_max) in C. partellus larvae and adult C. partellus and C. sesamiae. Lower cooling rates improved critical thermal minima (CT_min) in C. partellus and C. sesamiae adults while compromising CT_min in C. partellus larvae. The mean supercooling points (SCPs) for C. partellus larvae, pupae, and adults were ?11.82 ± 1.78, ?10.43 ± 1.73 and ?15.75 ± 2.47, respectively. Heat knock‐down time (HKDT) and chill‐coma recovery time (CCRT) varied significantly between C. partellus larvae and adults. Larvae had higher HKDT than adults, while the latter recovered significantly faster following chill‐coma. Current results suggest developmental stage differences in C. partellus thermal tolerance (with respect to lethal temperatures and critical thermal limits) and a compromised temperature tolerance of parasitoid C. sesamiae relative to its host, suggesting potential asynchrony between host–parasitoid population phenology and consequently biocontrol efficacy under global change. These results have broad implications to biological pest management insect–natural enemy interactions under rapidly changing thermal environments.     

Host status of wheat and corn for Chilo suppressalis (Lepidoptera: Crambidae)

Host plant specificity depends on recognition of the host and the ability to discriminate it from nonhost plants. Chilo suppressalis (Walker) (Lepidoptera: Crambidae), an important insect pest of rice, is considered to be polyphagous, although few papers have ever reported infestation of C. suppressalis on most of the recorded hosts. The present investigation was designed to test whether two important gramineous crops, wheat and corn, are host plants of C. suppressalis based on the host-plant finding process in cage and Y-tube olfactometer tests, oviposition and egg hatching, and larval feeding and survival. In the cage tests, gravid C. suppressalis females did not differentiate rice plants from wheat or corn plants when only visual cues were involved, but were more attracted to rice plants when only olfactory cues or both vision and olfaction were present. The Y-tube olfactometer tests further confirmed that the females did not prefer wheat or corn plants, and revealed that they responded equally to clean air and odors from wheat or corn plants. Under no-choice and choice condition alike, the females laid eggs on a lower proportion of wheat and corn plants and egg number and hatching rate were significantly reduced on wheat and/or corn plants than on rice plants. Larval feeding was not observed in wheat and lower in corn than in rice plants, and no pupae or surviving larvae were collected from wheat and corn plants. The results suggest that wheat and corn are not host plants of C. suppressalis. These findings are discussed in context of host-finding process in C. suppressalis and management of resistance to transgenic Bt rice.     

Diapause induction and photoperiodic clock in Chilo suppressalis (Walker) (Lepidoptera: Crambidae)

The mature larvae of the rice stem borer, Chilo suppressalis Walker (Lepidoptera: Crambidae) enters facultative diapause in response to short‐day conditions in the autumn (August–September). Diapause induction and photoperiodic clock mechanism were investigated in C. suppressalis larvae reared on an artificial diet in the present study. The critical night length for diapause induction was about 9 h 53 min to 10 h 39 min at 22 to 28°C. The third‐instar larvae were found to be relatively sensitive to diapause induction. Photoperiodic response under non‐24‐h light–dark cycles showed that scotophase length played an essential role in the induction of larval diapause in C. suppressalis, and consecutive exposure to long‐night cycles was necessary for a high diapause incidence. In the Nanda–Hamner experiment, diapause incidence peaked at scotophase of 12 h and dropped rapidly at scotophases > 24 h. In the Bünsow experiment, diapause incidence was clearly suppressed, especially at the light pulse located 8 h in the scotophase. Both the Nanda–Hamner and Bünsow experiments showed no rhythmic fluctuations with a period of about 24 h; thus the photoperiodic clock in C. suppressalis is a non‐oscillatory hourglass timer or a rapidly damping circadian oscillator.     

Proteolytic profile in the larval midgut of Chilo suppressalis Walker (Lepidoptera: Crambidae)

Chilo suppressalis is a key constraint on production of rice. The current research was conducted to study the types of digestive proteases in the larval midgut of C. suppressalis. It was found that activity of total digestive proteases increased from the first to the fifth larval instars, which showed different nutritional requirements. Four types of proteinases and two types of exopeptidase were identified so that their activities from the highest to the lowest activities is trypsin‐like, chymotrypsin‐like and elastase for proteinases, and amino and carboxypeptidases for exopeptidases. Meanwhile, just one type of cysteine protease, cathepsin D, was determined in the fourth and fifth instar larvae. The optimal pH for activity of total protease was found to be pH 9–10 and optimal temperature was observed to be 35–40°C, where there was the highest proteolytic activity. Some specific inhibitors of proteases including PMSF, TLCK, TPCK, DTT, E‐64, cystatin, phenanthroline and EDTA were used to confirm the types of proteases in the midgut of C. suppressalis.     

Effect of leaf toughness and temperature on development in the lilac pyralid,Palpita nigropunctalis (Bremer) (Lepidoptera: Crambidae)

This study focuses on three factors that affect the survival of the lilac pyralid, Palpita nigropunctalis (Lepidoptera:Crambidae): (1) the effect of leaf toughness on survival rate to clarify the availability of leaves as food, (2) the effect of temperature on immature development to determine the lower thermal threshold, and (3) the effect of temperature on head capsule width to clarify whether head capsule width can be used to discriminate among field-collected larval instars. Larvae could develop on Osmanthus fragrans var. aurantiacus leaves collected in April, but not on leaves collected in June or September which were too tough to eat. More than 80% of the larvae on the leaves of Ligustrum lucidum, Ligustrum japonicum, Ligustrum obtusifolium and Syringa vulgaris completed development, regardless of the collection time. P. nigropunctalis completed development on L. lucidum at temperatures from 15 to 27.5 °C with a photoperiod of either 15 L:9D or 16 L:8D, but not at 30 °C, at which temperature no eggs hatched. The lower thermal threshold and thermal constant for total development from egg to adult were estimated at about 7 °C and 450–460 degree-days. Most of the larvae were 5-instar type larvae (passed through 5 instars) regardless of the temperature, but a few 6-instar type larvae (4 of 355) were noted at temperatures of 22.5 °C and higher. No overlap of the ranges of head capsule widths was detected for the 5-instar type larvae, indicating that head capsule width can be used to discriminate among field-collected larval instars.     

Treatment of millet crop plant (Sorghum bicolor) with the entomopathogenic fungus (Beauveria bassiana) to combat infestation by the stem borer,Chilo partellus Swinhoe (Lepidoptera: Pyralidae)

Experiments were done to test if Beauveria bassiana can become an endophyte in sorghum and confer protection from stem borer. Four-week-old sorghum seedlings were treated with B. bassiana. The plants were examined for endophytic presence of B. bassiana, 30 and 60 days after treatment. Stem cultures from treated plants showed growth of B. bassiana. PCR amplification using fungal specific primers for a conserved region of β tubulin gene yielded identical 360 bp products from both B. bassiana and treated sorghum plants. In a subsequent experiment, B. bassiana treated and untreated (control) sorghum plants were artificially infested with stem borer (Chilo partellus) larvae 15 days post treatment and the extent of damage was compared. About 40% of the control plants developed dead heart while no plant in the B. bassiana treated plot did. In the surviving control plants, stem tunneling by shoot borer was significantly higher compared to B. bassiana treated sorghum plants.     

Effects of Hexaflumuron and Pyriproxyfen on the purified Phenoloxidase of Chilo suppressalis Walker (Lepidoptera: Crambidae)

Effects of two insect growth regulators (IGRs), hexaflumuron and pyriproxyfen, were studied on the purified phenoloxidase (PO) of Chilo suppressalis. Purification procedure revealed two isozymes of PO, namely POI and POII. IC₅₀ concentrations of hexaflumuron and pyriproxyfen on POI were 0.36, 0.23?μg/ml and on POII were 0.105, 0.42?μg/ml, respectively. Determination of optimal pH and temperature revealed pH 5 and temperature 40?°C as the optimal values for the enzymatic activity. Treating POs with IC₅₀ concentrations of two IGRs was pH and temperature dependent. Effects of these IGRs on POI caused significant increase of K_m value versus control suggesting competitive inhibition. Hexaflumuron and pyriproxyfen cause reduction in V_max value of POII versus control suggesting non-competitive inhibition. The current study shows direct effects of two IGRs on purified PO of C. suppressalis for the first time. These findings could be helpful to develop safe compounds with inhibitory mechanism on PO to neutralise insect immune responses against entomopathogenic agents.