Temperature related effects on embryonic development of the Mediterranean locust, Dociostaurus maroccanus

Laboratory studies were conducted to assess the effect of temperature on the development of the eggs of Dociostaurus maroccanus (Thunberg) (Orthoptera, Acrididae) during anatrepsis (stages I–XIV) and during catatrepsis (stages XV–XX). The developmental rates of anatrepsis were studied at five constant temperatures ranging from 10 to 30°C. Egg development occurred over the entire range but at 10°C the embryos were unable to complete anatrepsis. The relationship between temperature and developmental times for completing anatrepsis was analysed by the non‐linear Logan type III model. The optimal temperature estimated for the development of eggs during anatrepsis was 24.7°C; the lower and upper thermal thresholds were 9°C and 31°C, respectively. Once the embryos completed anatrepsis, only those incubated at 15°C continued morphogenesis beyond stage XIV (diapause stage) without a low‐temperature exposure period. The developmental rate of catatrepsis was studied at four constant temperatures ranging from 15°C to 30°C after exposure to low‐temperature, 10°C, for 30, 60 or 90 days. For catatrepsis, temperature and developmental time were linearly and inversely related. Linear regression was used to estimate the lower developmental threshold and the degree days requirements for catatrepsis. Both decreased with longer exposure to the low temperature; the former from 13.8°C to 10.5°C and the latter from 212.8 to 171.5 degree days, following 30 and 90 days at 10°C, respectively. Our results improve the ability of decision support systems for Mediterranean locust pest management by providing better forecasts to land managers and pest advisors.     

Development of microsporidia-infected Muscidifurax raptor (Hymenoptera: Pteromalidae) at different temperatures

Muscididfurax raptor, a pupal parasitoid of house flies and other filth flies, is commonly infected with the microsporidium Nosema muscidifuracis. To determine the effects of infection on developmental time, uninfected and infected adult M. raptor were allowed to parasitize pupae of the house fly (Musca domestica) for 24 h. Exposed pupae of the two groups (infected and uninfected) were held at 15, 20, 25, 30, 32, and 34 °C with 75–80% relative humidity. Development of infected M. raptor was significantly longer at all temperatures than that of uninfected parasitoids, resulting in approximately 7% extensions of developmental times. Uninfected females completed development in 14.6, 19.6, and 30.4 days at 30, 25, and 20 °C, respectively, compared with 15.8, 20.7, and 32.3 days for infected females at these temperatures. The differences in developmental times provided narrow windows for isolating large proportions of uninfected M. raptor females for disease management programs. This window was greatest at 20 °C; 61% of the uninfected females emerged by day 30, at which time only 10% of the infected females had emerged.     

INFLUENCE OF TEMPERATURE ON CERTAIN BIOLOGICAL ATTRIBUTES OF A LADYBEETLE COCCINELLA SEPTEMPUNCTATA LINNAEUS

Influence of temperature on certain biological attributes of an aphidophagous ladybeetle, Coccinella septempunctata Linnaeus, feeding on mustard aphid, Lipaphis erysimi (Kaltenbach), at five different temperatures, viz. 20, 25, 27, 30 and 35°C was investigated. Its developmental period was shortest (11.7 ± 0.09 days) at 35°C and longest (20.6 > 0.35 days) at 20°C. Developmental rate increased with increase in temperature. Hatching percent, larval survival, adult emergence and growth index were maximum at 30°C and minimum at 20°C. Oviposition period and fecundity were highest at 30°C and lowest at 20°C. A positive linear relationship exists between temperature and developmental rate and negative correlation between the duration of immature life stages and temperature. The proportion of developmental period allocated to each immature stage was found to be similar at each temperature regime. Thus, 30°C was found as the most suitable for C. septempunctata amongst the five temperatures tested.     

Defect in translation of poliovirus RNA at the restrictive temperature.

Translation of the RNA of LSc type 1 poliovirus was examined in vivo at the restrictive temperature (39 °C). During the first two hours of infection at 39 °C the levels of viral polyribosomes were 50% lower than at 35 °C (permissive temperature). During the third hour of infection at 39 °C, only 4 to 10% of the control levels of polyribosomes were observed. Three experiments indicate that the elongation of viral peptides was not occurring properly at 39 °C. First, cultures incubated at 39 °C during the third hour of infection with both [³⁵S]methionine and [³H]uridine exhibit a fourfold increase in the ratio of viral protein/viral RNA in the polyribosome region of sucrose gradients in comparison to controls kept at 35 °C. However, at both temperatures the relative size distribution of polyribosomes was similar. Second, the ratios of released protein/nascent protein after 90-second and 5-minute pulses with [³⁵S]methionine indicate that elongation of peptide chains was inhibited at 39 °C. Third, when initiation of synthesis of viral protein was blocked with 150 mM-NaCl, the polyribosomes disaggregated four to five times more rapidly at 35 °C than at 39 °C. The data indicate that translation of viral RNA is inhibited at the restrictive temperature because of a reduced rate of elongation of viral proteins. The reduced rate of peptide chain elongation at 39 °C was fully reversible when cultures were shifted to 35 °C in the presence of 150 mm-NaCl. The latter finding indicates a conformational change in viral protein at 39 °C.     

Development and reproduction of a potential biological control agent, Aphelinus varipes (Hymenoptera: Aphelinidae), at different temperatures

The development and reproductive potential of an indigenous parasitoid, Aphelinus varipes (Förster), was studied at 15, 17, 20, 25, and 30 °C. Developmental durations decreased with increasing temperatures. The emergence rate was higher than 90 % at 15, 17, and 20 °C. Offspring sex ratios were 0.69, 0.54, and 0.70 at 17, 20, and 25 °C, respectively, but were 0.14 at 15 °C and 0.38 at 30 °C. Developmental zeros of females and males were calculated as 9.9 and 9.6 °C, respectively. The effective accumulative temperature (K) was 204.1 degree-days in both sexes. Fecundity peaked in early age after emergence, then gradually decreased in a fluctuating manner at 20 and 25 °C. Host feeding continued constantly during the life of female adults at two temperatures. Single female parasitoids produced 218.5 and 203.1 mummies at 20 and 25 °C, respectively, during their lifespans. Aphids killed by parasitoid host feeding numbered 79.1 at 20 °C and 63.8 at 25 °C. Longevities were 27.0 days at 20 °C and 20.6 days at 25 °C. Moreover, intrinsic rates of natural increase (r _m) were estimated as 0.151 at 20 °C and 0.227 at 25 °C. We discuss the potential of A. varipes as biological control agents by comparing them with Aphidius colemani Viereck, which has been introduced to horticultural crops in greenhouses in Japan.     

Developmental rates of the aphid Aphis pomi (Aphidoidea: Aphididae) and its parasitoid Aphidius ervi (Hymenoptera: Aphidiidae)

Developmental rates of the aphid Aphis pomi and its parasitoid Aphidius ervi were compared at four constant temperatures (15, 20, 25, 30°C). Aphis pomi required 159.07 degree-days (DD) above lover developmental threshold (LDT) of 0.3°C. Aphidius ervi required 229.73 DD above LDT of 5.98°C. Developmental rate of A. pomi is higher than the developmental rate of A. ervi at the same conditions     

The combined effects of temperature and diet on development and survival of a crab spider,Misumenops tricuspidatus (Fabricius) (Araneae: Thomisidae)

In the present study, the combined effects of temperature and diet on development and survival of a crab spider, Misumenops tricuspidatus (Fabricius) (Araneae: Thomisidae) in laboratory conditions were investigated. The experiments were carried out at five constant temperatures ranging from 15°C to 35°C on two kinds of diets, fruit flies (Drosophila melanogaster) and a mixed diet of fruit flies and dung flies. It was found that development rate of eggs increased with successive temperature increments, reaching a maximum at 30°C, then declined at 32°C and that eggs survived well between 20°C and 30°C (>70%), but no eggs survived to hatching at 35°C regardless of whether the spiders were fed on single or mixed diet. Juveniles completed development on both diets at all constant temperatures tested, but survival was low at the extreme temperatures. Juvenile development times decreased over successive temperature increments up to 30°C, then increased at 32°C. Females developed faster than males. Diet also influenced development time, survival and number of moults to reach maturity. Juveniles raised on the mixed diet composed of fruit flies and dung flies developed faster, survived better, and required fewer moults to reach maturity than on a diet composed of only fruit flies. Plots of development rates (reciprocal of mean development times) and survival rates (expressed as percentages) against constant temperatures indicated that M. tricuspidatus is well adapted to low temperatures, but detrimentally affected by high temperatures. Using linear regression, the lower development threshold (LDT) and the sum of effective temperatures (SET) for all life stages of M. tricuspidatus on each diet were estimated. LDT and SET varied among developmental stages and between diets.     

Effect of high temperature on the development of nuclear polyhedrosis virus in the silkworm,Bombyx mori

Effect of a high temperature on the development of nuclear polyhedrosis and nuclear polyhedrosis virus (NPV) was studied employing pupae and isolated pupal abdomens of the silkworm, Bombyx mori. It was shown that pupae inoculated with an NPV and incubated at 35°C survived longer than those incubated at 25°C. At lower dosages of virus, pupae at 35°C escaped death from NPV. When inoculated pupae were incubated at 35°C for varying periods and then transferred to 25°C, the longer the pupae had been kept at 35°C the longer they survived. In contrast, when inoculated pupae were transferred from 25° to 35°C, the longer the pupae had been kept at 25°C the sooner after inoculation they died. Essentially the same results were obtained in isolated abdomens which were in an arrested state of development, excluding the possibility that observed thermal inhibition of viral diseases is dependent upon the altered developmental processes at high temperatures. Virus titration experiments showed that, under experimental conditions utilized, no detectable accumulation of infectious NPV was present in abdomens inoculated with an NPV and incubated at 35°C. When inoculated abdomens were shifted up from 25° to 35°C at 3 days postinoculation, NPV accumulation was inhibited almost immediately, and when inoculated abdomens were shifted down from 35° to 25°C, infectious NPV started to accumulate as early as 1 day after the shift. It was also shown that the pattern of infectious NPV accumulation and that of nucleic acid increase in infected abdomens gave a rough correlation. These results indicate that the thermal inhibition of viral diseases is attributed, at least in part, to the restricted accumulation of infectious progeny and suggest that the virus replication mechanism itself is more sensitive to high temperatures than that related to other events necessary for viral replication to be initiated.     

Temperature-dependent development of <Emphasis Type="Italic">Macrolophus pygmaeus</Emphasis> and its applicability to biological control

A linear model and three nonlinear models (Logan type III, Lactin and Brière) were applied to Macrolophus pygmaeus (Rambur) (Hemiptera: Miridae) at constant temperatures and validated under diel temperature variation, and field conditions. Complete development from egg to adult, with >80% survivorship, occurred at nine constant temperatures between 15 and 32 °C. Total developmental time decreased from a maximum at 15 °C (68.48 days) to a minimum at 30 °C (18.69 days) and then increased at 32 °C (23.44 days). Optimal survival and the highest developmental rate occurred within the range of 27–30 °C. The adjusted determination coefficients were high for linear and nonlinear models (>0.89). Field validation showed high levels of accuracy in all models (≥93.4%). These valid mathematical models contribute to optimal application, field management, and mass rearing of M. pygmaeus for its applicability to biological control.     

Development ofDiatraea Saccharalis [Lep.: Pyralidae] at constant temperatures

At constant temperatures between 15.6 and 32°C the incubation time of eggs ofDiatraea saccharalis (F.) was reduced by each increase in temperature. At 34°C the time decreased. Highest (98.6%) and lowest (9.9%) egg hatch occurred at 26 and 34°C, respectively. Larvae completed development at temperatures ranging from 22 to 34°C; however, only 4.4% of the larvae pupated at 34°C. Duration of the larval stage at 30°C (♂=18.1 days; ♀=19.1 days) was ca. 14 days shorter than at 22°C. Maximum rate of development in the pupal stage occurred at 28°C (ca. 6.8 days), and a higher temperature increased developmental time and mortality. Adult longevity and egg production generally were reduced with increasing temperatures and egg production was highest at 24°C (729.8 eggs/ moth). As many as 7 larval stages occurred; but most larvae completed development in 5 stages, and none completed development in less than 5 stages. The female larval stage was ca. 1 day longer than that of males, and this difference occurred primarily in the 5th stage.     

Life table and predation of Neoseiulus longispinosus (Acari: Phytoseiidae) on Oligonychus coffeae (Acari: Tetranychidae) infesting tea

Life table and predation of the predatory mite Neoseiulus longispinosus (Evans) on the red spider mite (RSM), Oligonychus coffeae (Nietner), a major pest of tea in India, were studied in the laboratory. Developmental time from egg to adult varied from 4 to 14 days at 30 to 15 °C, respectively; at 35 °C no larva survived. Survival of immature stages was more than 94 % at all temperatures. Threshold temperature for development of immature stages of females and males was 10 and 9.9 °C, respectively, and thermal constant was 84.03 degree-days for females and 80 for males. Sex ratio was female biased and temperature (20–30 °C) had no clear effect on sex determination. Egg hatchability was 73 % at 35 °C and >97 % at lower temperatures. Average number of eggs laid per female/day was higher at 30 °C than at 20 or 25 °C. The highest net reproductive rate (R ₀) was 40.7, at 20 °C. Mean generation time (T) decreased from 28 to 13 days with temperature increasing from 20 to 30 °C. Weekly multiplication (6.5) and intrinsic rate of natural increase (r _m ) (0.268) were highest at 30 °C. Males lived longer than females at every temperature tested. Longevity was highest at 20 °C (50 days for females and 55 for males). Survival and longevity were adversely affected by temperature above 30 °C. Daily consumption of prey increased with the advancement of predator’s life stages; adult females consumed the highest numbers of prey items, preferably larvae and nymphs.     

Development times and fecundity of three important arthropod pests of strawberry in the United Kingdom

The development rates and fecundity of three important pests of strawberry in the UK were determined over a range of temperatures. Development time of the strawberry tarsonemid mite, Phytonemus pallidus, from egg lay to adult, ranged from a mean of 28.4 days at 12.5°C to 8.8 days at 25°C. No nymphs developed to adult at 10°C. Females lived for up to 45 days and laid a mean of 24.3 and 28.5 eggs at 20°C and 25°C respectively. Total development time from egg lay to adult for the strawberry blossom weevil, Anthonomus rubi, ranged from a mean of 95.7 days at 10°C to 18.2 days at 25°C. Mean fecundity at 20°C was 157.6 eggs, and the oviposition period averaged 71.6 days. When nymphs were reared on strawberry, development of the European tarnished plant bug, Lygus rugulipennis, from egg lay to adult, ranged from 83.8 days at 15°C to 28.8 days at 25°C. Development times on groundsel were shorter and ranged from 65.6 to 22.2 days at 15°C and 25°C. Only two nymphs developed to adults at 10°C; no eggs hatched at that temperature. Mean fecundity at 20°C was 75.4 eggs, but ranged from 23 to 179. Under a fluctuating temperature regime of 10°C for 12 h:20°C for 12 h, nymphs of L. rugulipennis took 40.3 days to become adult on strawberry, and 33.4 days on groundsel. Simple linear models fitted the developmental rate ‐ constant temperature relationship well for all species, accounting for 95–98% of the total variation in observed developmental rates. Development under fluctuating temperatures illustrated the potential problem of extrapolating linear models beyond the conditions of the experiment.     

Effects of temperature during early life history on embryonic and larval development and growth in haddock

The effect of incubation temperature (2, 4, 6, 8 and 10° C) on haddock Melanogrammus aeglefinus development and growth during the embryonic period and in subsequent ontogeny in a common post‐hatch thermal environment (6° C) was investigated. Hatching times were inversely proportional to incubation temperature and ranged from 20·3 days at 2° C to 9·1 days at 10° C. Growth rates were directly proportional to incubation temperature during both the embryonic and larval periods. There was a significant decline in growth rates following hatch in all temperature groups. Compared to the endogenously feeding embryos, growth rates in the exogenous period declined by 4·4‐fold at 4° C to 3·9‐fold at 8° C, indicative of the demarcation between the endogenous and exogenous feeding periods. Yolk utilization varied from 17 days at 2° C to 6 days at 10° C and followed a three‐stage sigmoidal pattern with the initial lag period inversely proportional to incubation temperature. Time to 50% yolk depletion varied inversely with temperature but occurred 1–1·5 days post‐hatch at all temperatures. Additionally, the period between 10 and 90% yolk depletion also decreased with increased temperature. Overall developmental rate was sequential with and directly proportional (2·3‐fold increase) to incubation temperature while the time spent in each developmental stage was inversely proportional to temperature. Larger embryos tended to be produced at lower temperatures but this pattern reversed following hatch, as larvae from higher temperature groups grew more rapidly than those from other temperature groups. Larvae from all temperatures achieved a similar length (c.total length 4·5 mm) upon complete yolk absorption. The study demonstrated the significant impact that temperature has upon developmental and growth rates in both endogenous and exogenous feeding periods. It also illustrated that temperature changes during embryogenesis had significant and persistent effects on growth in subsequent ontogeny.     

Cross-mating experiments with geographically different populations of <Emphasis Type="Italic">Amblyomma cajennense</Emphasis> (Acari: Ixodidae)

The effect of temperature on the development and fecundity of Sancassania polyphyllae fed on tissues of Polyphylla fullo larvae was studied at 15, 20, 25, 30, and 35 ± 1°C and 65 ± 10% RH in a dark incubator. Mean developmental period of immature stages decreased significantly with increasing temperatures from 15 to 30°C. Developmental periods at 30–35°C were not significantly different. The estimated lower developmental thresholds of the various immature stages ranged between 10.1 and 11.5°C. The thermal constant for the egg-to-female adult was 93.5 degree-days. The pre-oviposition, oviposition, and post-oviposition periods and female longevity were significantly longer at 15°C than at higher temperatures. Mean total and daily fecundity were the highest at 25°C, which were significantly different from those obtained at 15, 20 and 30°C. The net reproductive rate (R ₀) was the highest at 25°C (588.3 ♀/♀). The longest mean generation time (T ₀) occurred at 15°C (36 days) and the shortest occurred at 30°C (9.2 days). The highest intrinsic rate of increase (r _m) for S. polyphyllae was observed at 25 (0.61 ♀/♀/day) and 30°C (0.62 ♀/♀/day).     

The thermal tolerance of crown-of-thorns (Acanthaster planci) embryos and bipinnaria larvae: implications for spatial and temporal variation in adult populations

To understand the role of sea temperature on the population biology of the crown-of-thorns sea star Acanthaster planci, the thermal window for embryonic and larval development was investigated. In two experiments, the response of embryos and larvae across 12 temperatures from 19.4 to 36.5 °C was quantified as the percentage of individuals reaching cleavage stage embryos, blastula, gastrula, early-bipinnaria, late-bipinnaria larvae or abnormal. Measurements were made at 7 times up to 72 h post-fertilisation, with the morphometrics of larvae measured in the 72-h sample. Acanthaster planci developed at temperatures between 19.4 and 33.2 °C, with a thermal window for development to the late-bipinnaria stage between 25.6 and 31.6 °C. Development rate, normal development and larval size were optimal at 28.7 °C, with development rates remaining relatively constant up to 31.6 °C. Rates of abnormality increased steadily (early embryonic stages) above 28.7 °C and was 100 % at temperatures approaching 33 °C. These experiments provide a more detailed insight into the response of A. planci developmental stages to temperature. The present day distribution of the species in eastern Australia overlap with the optimal thermal window for development to the late-bipinnaria stage (≈25–32 °C), implying a role of temperature in controlling population distributions and abundances. Despite this, short- or long-term temperature increases may not be a major modulator of the crown-of-thorns recruitment success, population dynamics and distribution in the future as no significant change in development rates, larval survival and growth occurred within this thermal window. Therefore, moderate (1–2 °C) increases in sea temperatures caused by El Niño or near-future ocean warming may not drive an increase in developmental and settlement success. Indeed, without any acclimation to warmer temperatures expected under near-future warming (+2 to 4 °C), climate change could ultimately reduce larval survival due to elevated mortality above the optimal development temperature.     

Impact of fluctuating temperatures on development of the koinobiont endoparasitoid Venturia canescens

The effect of temperature on the biology of Venturia canescens (Gravenhorst) (Hymenoptera: Ichneumonidae) is well understood under constant temperature conditions, but less so under more natural, fluctuating conditions. Herein we studied the influence of fluctuating temperatures on biological parameters of V. canescens. Parasitized fifth-instar larvae of Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) were reared individually in incubators at six fluctuating temperature regimes (15–19.5 °C with a mean of 17.6 °C, 17.5–22.5 °C with a mean of 19.8 °C, 20–30 °C with a mean of 22.7 °C, 22.5–27.5 °C with a mean of 25 °C, 25.5-32.5 °C with a mean of 28.3 °C and 28.5–33 °C with a mean of 30 °C) until emergence and death of V. canescens adults. Developmental time from parasitism to adult eclosion, adult longevity and survival were recorded at each fluctuating temperature regime. In principle, developmental time decreased with an increase of the mean temperature of the fluctuating temperature regime. Upper and lower threshold temperatures for total development were estimated at 34.9 and 6.7 °C, respectively. Optimum temperature for development and thermal constant were 28.6 °C and 526.3 degree days, respectively. Adult longevity was also affected by fluctuating temperature, as it was significantly reduced at the highest mean temperature (7.0 days at 30 °C) compared to the lowest one (29.4 days at 17.6 °C). Survival was low at all tested fluctuating temperatures, apart from mean fluctuating temperature of 25 °C (37%). Understanding the thermal biology of V. canescens under more natural conditions is of critical importance in applied contexts. Thus, predictions of biological responses to fluctuating temperatures may be used in population forecasting models which potentially influence decision-making in IPM programs.     

Temperature dependent development and temperature thresholds of Rhyncaphytoptus ficifoliae Keifer (Diptilomiopidae)

The effects of temperature on developmental rate of Rhyncaphytoptus ficifoliae Keifer (Diptilomiopidae) were determined at six constant temperatures (17, 20, 25, 30, 33 and 36 °C) on fig leaves. The total developmental time of females decreased as temperature increased from 17 (21.62 days) to 33 °C (6.02 days), and then increased at 36 °C (6.47 days). Using the ordinary and Ikemoto and Takai (2000) linear models the estimated lower temperature thresholds (T_min) for total developmental time of females were 10.78 and 10.37 °C and the constant temperatures (k) were 140.25 and 144.78°-days (DD), respectively. Data also were fitted to SSI nonlinear temperature-dependent model. The estimated T_L, intrinsic optimum temperature (T_Ф) and T_h for total immature stages of females by SSI model were 11.11, 23.72 and 37.98 °C, respectively. With use of the obtained data from rearing R. ficifoliae under constant temperatures in laboratory and temperature data in Khorramabad region in 2017, the real developmental rate of this mite in natural conditions was described. The highest (100%) and lowest values (51.67%) of survival rate for immature stages were found at 25 and 36 °C, respectively. The presented information could be used to predict the population dynamics of main pest R. ficifoliae for an effective management.     

Parameter estimation for a temperature-dependent development model of Thrips palmi Karny (Thysanoptera: Thripidae)

Development of immature Thrips palmi Karny was investigated at 12.5, 15, 17.5, 20, 22.5, 25, 27.5, 30, 32.5, and 35 °C, 20–40% RH and a photoperiod of 14:10 (L:D) h. Developmental time decreased with increasing temperature up to 32.5 °C in all stages. The total developmental time was longest at 12.5 °C (64.2 days) and shortest at 32.5 °C (9.2 days). The lower developmental threshold was 10.6, 10.6, 9.1, and 10.7 °C for egg, larva, prepupa, and pupa, respectively. The thermal constant required to complete the respective stage was 71.7, 59.2, 18.1, and 36.8DD. The lower threshold temperature and thermal constant were 10.6 °C and 183.3DD, respectively, for total immature development. The nonlinear relationship between developmental rate and temperature was well described by the modified Sharpe and DeMichele biophysical model (r² = 0.905–0.998). The distribution of developmental completion of each stage was described by the 3-parameter Weibull function (r² = 0.855–0.927). The temperature-dependent developmental models of T. palmi developed in this study could be used to predict its seasonal phenology in field and greenhouse vegetable crops.     

Preimaginal development,adult longevity and fecundity ofEncarsia inaron (Hym.: Aphelinidae) parasitizingSiphoninus phillyreae (Hom.: Aleyrodidae) in California

A laboratory study of preimaginal development, adult longevity and fecundity ofEncarsia inaron (Walker) was conducted. Preimaginal developmental times varied with temperature, from 55–60 d at 15±1°C to 14–17 d at 30±1°C. No development took place at 10°C, which was approximately the developmental minimum estimated from regression analysis of developmental rates vs. temperature in the range 15–30°C. Development was slowed and survival was reduced at 32°C. Females lived an average of 18.6 days and laid a average of 159 eggs/female at 25°C. At 25°C, average preimaginal survival was 59.3%, and the sex ratio was 73.5% female. The net reproductive rate (R₀) forE. inaron calculated from these studies was 69.3, while the intrinsic rate of natural increase was 0.1686 individuals per individual per day. Oviposition was concentrated slightly in third instar nymphs of the host.