Biology, life table and host specificity of the mushroom pest, Brennandania lambi (Acari: Pygmephoroidea)

Biology and life table parameters of Brennandania lambi (Krczal) were studied at different temperatures while feeding on white mushroom (Agaricus bisporus) mycelium cultured on mushroom compost. The duration of egg and larva development, preoviposition and oviposition period, female longevity, and the time to 50% mortality declined as temperature increased from 16 to 28°C. The threshold temperature of development (female) was 9°C and the thermal constant for completion of development (female) was 195 day-degrees. At 16, 20, 24 and 28°C, the total fecundity (eggs/female) was 71, 67, 66 and 57, respectively and the daily fecundity rate (eggs/female/day) was 5.6, 8.7, 8.7 and 9.1, respectively. The sex ratio (female/male) ranged from 1.9 to 2.1 at 16–28°C. At 16, 20, 24 and 28°C, the intrinsic rate of natural increase (r _m) was 0.11, 0.18, 0.22 and 0.27, respectively, and the population doubling time was 6.1, 3.9, 3.2 and 2.5 days, respectively. All life stages of the mite died when exposed to 35°C constant temperature for 24h, or to 32°C constant temperature for 12 days or to 31–35°C (average 32.9°C) ambient temperature for 4 days. Brennandania lambi completed development only when fed on Ag. bisporus mycelium growing on mushroom compost. It could not survive on mushroom mycelia of Auricularia auricula, Au. polytricha, Ganoderma lucidum, Hericium erinaceus, Lentinus edodes, Pleurotus ostreatus, P. sajor-caju and Tremella fuciformis.     

Life history and demography of the predatory mite,Amblyseius longispinosus Evans

The life history and demography of Amblyseius longispinosus Evans were studied under laboratory conditions 25–28°C and 65–85% RH using the red form of the twospotted spider mite, Tetranychus urticae Koch complex as prey. The entire development from egg to adult averaged 102.5 h for both sexes with a survival rate of 90.0%. Immature females consumed more prey eggs than males, averaging 7.9 and 6.4 eggs/day, respectively. The embryonic development time for male eggs was longer (average 45.2 h) than for female eggs (average 42.6 h). The average duration of succeeding stages did not differ between the sexes: larva, 15.7; protonymph, 21.1; and deutonymph, 23.0 h. Larvae were not observed to feed. Statistically significant differences between sexes were noted with respect to consumption of the deutonymphs (female, average 4.0±0.2; male, average 2.9±0.3 eggs/day) but not of the protonymphs. Mating occurred on the same day as adult emergence and was repeated several times during the reproductive life. Egg laying started on the second day after emergence. Parameters relating to oviposition were: fecundity, gross (50.7) and net (43.3) eggs/female; hatchability, 99.6% (gross) and 99.5% (net); the oviposition lasted at most 28 days and showed a peak in the first week. Net reproductive rate (R₀) was 36.7 female offspring/female/generation time (T) of 9.0 days. The sex ratio was biased toward the females (71.9%), while the intrinsic rate of increase (r_m) was 0.4 with a doubling time (DT) of 1.7 days. Males lived longer (36 days) than females (30 days) with a LT₅₀ for males of 26 days and for females of 22 days.     

Influence of diet on life table parameters of <Emphasis Type="Italic">Iphiseius degenerans</Emphasis> (Acari: Phytoseiidae)

Studies on the reproduction, longevity and life table parameters of Iphiseius degenerans (Berlese) were carried out under laboratory conditions of 25 ± 1 °C, 75 ± 5% RH and 16L:8D h. As food sources for the predatory mite, Ricinus communis L. pollen, all stages of the spider mite Tetranychus urticae Koch, Frankliniella occidentalis (Pergande) larvae, and Ephestia kuehniella Zeller eggs were selected. All diets were accepted as food by the adult mites. Female longevity ranged from 29.5 to 42.4 days, the highest value was recorded on a diet of Ephestia eggs. The highest percentage of females escaping the experimental arena was observed on the diet consisting of thrips larvae. The highest oviposition rate (1.9 eggs/female.day) was recorded when the predator was fed on spider mites on an artificial substrate. For other diets, oviposition rates ranged from 1.0 to 1.3 eggs/female.day. The intrinsic rate of natural increase (r _m) of I. degenerans varied between 0.015 and 0.142 females/female.day. The diet consisting of castor bean pollen resulted in the highest population growth whereas the diet on spider mites brushed off onto a bean leaf arena resulted in the slowest population growth. This can be explained by the inability of the predator to cope with the webbing of T. urticae, and the high escape rate of the progeny when reared on spider mites. The percentage of females in the offspring ranged from 40 to 73%.This revised version was published online in May 2005 with a corrected cover date.     

Effect of temperature on reproductive parameters and longevity of <Emphasis Type="Italic">Tyrophagus putrescentiae</Emphasis> (Acari: Acaridae)

The effect of temperature on reproductive parameters and longevity of the mold mite, Tyrophagus putrescentiae (Schrank) was examined at seven constant temperatures, ranging from 10 to 34 °C, and a relative humidity of 90±5%. Preoviposition period and fecundity were adversely affected by extreme temperatures and the oviposition period increased as temperature was reduced. Different patterns were observed for longevity data for males and females, with greater longevities for males at intermediate temperatures and more similar values for both sexes at extreme temperatures. Polynomial and non-linear models provided a good fit of the relationship of reproductive and longevity parameters with temperature. The effect of temperature on the intrinsic rate of increase of T. putrescentiae populations was established by the non-linear Lactin model. The optimum temperature for development was obtained at 30 °C. At this temperature, the population doubling time is 1.75 days. The lower and upper thresholds for T. putrescentiae populations were established at 10.4 and 34.8 °C, respectively. Altogether, these data provide basic information to develop sound physical control strategies of the mold mite.     

The predatory mite Hypoaspis miles: temperature dependent life table characteristics on a diet of sciarid larvae, Bradysia paupera and B. tritici

Life table characteristics of Hypoaspis miles Berlese (Acarina: Hypoaspidae) fed on a mixture of Bradysia paupera Tuomikoski (Diptera: Sciaridae) and B. tritici Coquillet larvae were investigated in laboratory experiments at 4 temperatures (15, 20, 25, 30 °C) for development time, juvenile mortality, sex ratio, preoviposition period, oviposition period, postoviposition period, age-specific fecundity, and adult longevity. Juvenile development time decreased with increasing temperature from 46 days at 15 °C to 10 days at 30 °C. The lower temperature threshold was 9.9 °C and development required 205 °D. Juvenile mortality decreased from 52% at 15 °C to 3% at 25 °C and then increased to 24% at 30 °C. Preoviposition period varied with temperature from 12 days at 15 °C to 3 days at 25 °C and then increased to about 4 days at 30 °C. Oviposition period decreased with increasing temperature from 58 days at 15 °C to 25 days at 30 °C. The mean number of eggs per female per day increased from 0.4 at 15 °C to 2.3 at 25 °C and decreased to 1.3 at 30 °C. Age-specific fecundity was described by a temperature dependent model from which the maximum daily fecundity rate could be estimated to be attained at 25.6 °C. Female longevity was significantly shorter than for males, and decreased from 90 days at 15 °C to 34 days at 30 °C. Sex ratio was female-biased at all 4 temperatures and increased with temperature up to 25 °C, decreasing at 30 °C. Estimates of net reproductive rate, intrinsic rate of increase, finite rate of increase, mean generation time and doubling time were obtained. The r _m -value increased with temperature from 0.031 day^-1 at 15 °C to 0.133 day^-1 at 25 °C, after which it decreased to 0.112 day^-1 at 30 °C. The study showed that H. miles can develop and reproduce at temperatures between 15 and 30 °C. H. miles and sciarids have approximately the same optimum temperature and thresholds for development and reproduction and H. miles can be used for biological control of sciarids within the temperature range where the pest occurs.     

Life history study of Thrips setosus

Development, reproduction and population growth of Thrips setosus Moulton (Thysanoptera, Thripidae), reared on a leaf of kidney bean, was studied under six different constant temperatures, and the effect on reproduction of short photoperiod during immature stages was examined. Survival rates from hatch to adult were more than 67.5% at temperatures between 17.5 and 27.5 °C, but less than 55% at 30 °C. Developmental rates increased linearly as rearing temperature increased. A total of 181.1 degree-days, above a developmental zero of 12.5 °C, were required to complete development from egg to adult oviposition. These data were related to records of field temperatures in Kurashiki in western Japan, and an estimate produced that, under outdoor conditions, a maximum of between seven and 12 generations could have developed annually between 1990 and 1999. There were no significant differences in mean adult longevity and mean fecundity among three temperatures (20, 22.5 and 25 °C). The intrinsic rate of natural increase (r _m) was 0.1997 at 25 °C. Reproductive diapause was induced by a photoperiod less than 12 h at 20 °C.     

Life history of the phoretic predatory mite Arctoseius semiscissus (Acari: Ascidae) on a diet of sciarid fly eggs

A life table was constructed for the phoretic predatory mite Arctoseius semiscissus when it was offered sciarid eggs as food. The entire development from egg to adult lasted 7.9 days on average for both sexes with a survival rate of 68% and a sex ratio of 0.62. Parameters relating to oviposition were total fecundity (58.5 eggs per female), egg hatchability (79%) and oviposition (maximally 35 days). The net reproductive rate (Ro) was 24.49, the generation time (T) 13.85, the intrinsic rate of increase (rm) 0.23 and the finite rate of increase () 1.26. Exp Appl Acarol 22: 643–648 © 1998 Kluwer Academic Publishers     

High Temperatures Eliminate Wolbachia, a Cytoplasmic Incompatibility Inducing Endosymbiont, From the Two-spotted Spider Mite

Wolbachia can induce cytoplasmic incompatibility (CI) in the arrhenotokous two-spotted spider mite between uninfected females and infected males. Cytoplasmic incompatibility is expressed through a male-biased sex ratio and a low hatchability, and can be suppressed by removing Wolbachia from spider mites reared on a diet with antibiotics. Here we investigated whether heat-treatment can elimate Wolbachia from infected mites. Using a PCR assay with a Wolbachia-specific primer pair (ftsZ), and by standard crosses, we were able to show that 71 per cent of the mites had lost the Wolbachia infection after rearing the infected population at 32 ± 0.5 °C for four generations. The infection could be completely removed when mites were reared at 32 ± 0.5 °C for six generations. Curing through high temperatures could be one of the reasons why mixed infected/uninfected populations occur in the field. An additional consequence of rearing mites at 32 ± 0.5 °C was the shortened development time. The effect of environmental temperature on the abundance of Wolbachia and possible behavioural consequences for the spider mite are discussed.     

Bionomics and population growth statistics of apterous virginoparae of woolly apple aphid, Eriosoma lanigerum, at constant temperatures

Development, survival, reproduction and population growth statistics of apterous virginoparae of woolly apple aphid, Eriosoma lanigerum (Hausmann) (Hemiptera: Aphididae) at constant temperatures of 10, 13, 15, 20, 25, 30 and 32°C are reported. The developmental times of all life stages were inversely related to temperature ranging from 10 to 30°C. Span of total development (time from birth to adulthood) decreased from 57.8 days at 10°C to 11.7 days at 30°C and increased to 16.8 days at 32°C. A good linear model fit (R²>0.96) between developmental rate and temperature in the range 10–25°C was observed for all life stages. The lower developmental threshold was estimated at 5.8°C for instar I, 4.8°C for instar II, 4.9° for instar III and 4.4°C for instar IV. The lower temperature threshold for total development was estimated at 5.2°C. The upper developmental limit was found to be 32°C. Mean degree-day accumulations required for completion of instars I, II, III, IV and total development were: 125.6, 51.0, 47.7, 50.7 and 267.6, respectively. Fecundity, larviposition period and adult longevity were reduced with increasing temperature. Net reproductive rate was greatest at 15°C whereas intrinsic rate of increase peaked at 25°C. Optimal performance, as measured by fecundity, survival and intrinsic rate of increase, ocurred in the range 13–25°C.     

Reproduction,longevity and life table parameters of <Emphasis Type="Italic">Tyrophagus neiswanderi</Emphasis> (Acari: Acaridae) at constant temperatures

The astigmatid mite Tyrophagus neiswanderi Johnston and Bruce is mainly considered a pest of ornamental and horticultural crops. However, this mite has been found infesting Cabrales cheese in Spain, though its population density is low compared to Acarus farris, the prevalent species of astigmatid mite encountered in Cabrales cheese maturing caves. One of the factors that might be influencing this differential abundance is temperature. In the present study the effect of temperature on reproductive parameters and longevity of T. neiswanderi was examined at six constant temperatures, ranging from 10 to 31°C, and a relative humidity of 90 ± 5%. Preoviposition period, fecundity and daily fecundity were adversely affected by extreme temperatures while the oviposition period increased as temperature was reduced. Male and female longevity increased as temperature decreased, but males showed significantly greater longevity than females. Additionally, this difference was greater as temperature decreased. The effect of temperature on the intrinsic rate of natural increase of T. neiswanderi populations was described by the non-linear Lactin model. The optimum temperature for development was predicted at 26.6°C. At this temperature, the population doubling time is 2.8 days. The lower and upper thresholds for T. neiswanderi populations were calculated at 7.4 and 31.7°C, respectively. According to these results, the influence of temperature on the low population density of this mite found in Cabrales maturing caves compared with A. farris is discussed.     

不同三叶草对二斑叶螨生长发育和繁殖的影响

【目的】明确红三叶、白三叶和杂三叶3种寄主对二斑叶螨Tetranychus urticae Koch生长发育和繁殖的影响,以及3种寄主间二斑叶螨的生物学差异。【方法】在室内(25±1)℃条件下,采用离体叶片法研究了3种三叶草对二斑叶螨种群参数的影响。【结果】3种三叶草对二斑叶螨生长发育、雌成螨寿命以及繁殖力有显著影响(P<0.05)。在杂三叶上,二斑叶螨发育历期、产卵前期明显比红三叶和白三叶上要长,而且雌成螨寿命和产卵期也明显缩短。每雌产卵量在红三叶上最高(150.87粒),白三叶次之(139.43粒),而杂三叶上最低(86.95粒)。二斑叶螨在3种三叶草上的存活曲线均为Ⅰ型,存活率高低依次为红三叶、白三叶和杂三叶。净增殖率(R0)、内禀增长率(rm)和周限增长率(λ)在红三叶上最高、杂三叶上最低,而平均世代周期(T)和种群加倍时间(Dt)在杂三叶上最长、红三叶上最短。【结论】3种三叶草对二斑叶螨的适合度具有一定差异,红三叶和白三叶对二斑叶螨具有更高的适合度。     

Effect of temperature,humidity and photoperiod on mortality of Mononychellus tanajoa (Acari: Tetranychidae) infected by Neozygites cf. floridana (Zygomycetes: Entomophthorales)

The effect of temperature, humidity and photoperiod on the development of Neozygites cf. floridana (Weiser and Muma) in the cassava green mite, Mononychellus tanajoa (Bondar) was studied in the laboratory. Dead infected mites began to appear 2.5 days after inoculation. At 33 and 28°C peak mortalities were higher and occurred earlier (after 2.5 days), than at 23 and 18°C. Mean LT₅₀ (time for half the infected mites to die) decreased with increasing temperature as follows: 3.9, 3.0, 2.9 and 2.5 days at 18, 23, 28 and 33°C, respectively. When placed under conditions of high relative humidity for a period of 24 h, the percentage of dead infected mites from which the fungus sporulated was highest at 28°C (51.4%) and lowest at 33°C (6.5%). The development of the fungus inside the mite was not significantly affected by ambient humidity or photoperiod. No significant interactions between tested factors were found.     

Coexistence of Tetranychus urticae (Acarina: Tetranychidae) with different capacities for diapause: comparative life-history traits

Summary Individuals of the two-spotted spider mite, Tetranychus urticae Koch (Acarina: Tetranychidae), with and without a capacity for diapause, coexist in central Japan. Diapause appears to be adaptive because females in diapause suffered less mortality than non-diapausing individuals when frozen at –24°C for more than 4 h. However non-diapausing females showed good survival up to 4 h of freezing. Active non-diapausing mites survived on rose leaves in Kyoto (35°N) throughout the winters of 1988–1989 and 1989–1990. Cultures of mites with low (LD) and high (HD) diapause capacities at 18°C and 9L-15D photoperiod were successfully selected from the rose population and from a population on chrysanthemum in Nara (34.4°N). Their life-history traits at 15, 20, 25 and 30°C were characterized. HD and LD mites from both populations were of similar ages at first reproduction at 15–30° C. However, at temperatures 20° C, HD individuals produced more eggs than LD individuals, resulting in higher fecundity and intrinsic rate of natural increase. These traits allow HD individuals, which stop breeding between October and April, to increase faster in summer than LD individuals. This provides a mechanism, together with climatic fluctuations, in maintaining the coexistence of diapausing and non-diapausing T. urticae in Kyoto where winter conditions are rarely lethal to the non-diapausing individuals.     

Reproductive and developmental biology of Gonatocerus ashmeadi (Hymenoptera: Mymaridae), an egg parasitoid of Homalodisca coagulata (Hemiptera: Cicadellidae)

The reproductive and developmental biology of Gonatocerus ashmeadi Girault, a parasitoid of the glassy-winged sharpshooter Homalodisca coagulata (Say), was determined at five constant temperatures in the laboratory: 15; 20; 25; 30; 33 °C. At 30 °C, G. ashmeadi maintained the highest successful parasitism rates with 46.1% of parasitoid larvae surviving to adulthood. Lifetime fecundity was greatest at 25 °C and fell sharply as temperature either increased or decreased around 25 °C. Temperature had no effect on sex ratio of parasitoid offspring. Mean adult longevity was inversely related to temperature with a maximum of 20 days at 15 °C to a minimum of eight days at 33 °C. Developmental rates increased nonlinearly with increasing temperatures. Developmental rate data were fitted with the modified Logan model for oviposition to adult development times across each of the five experimental temperatures to determine optimal and upper lethal temperature thresholds. The lower developmental threshold estimated by the Logan model and linear regression were 1.10 and 7.16 °C, respectively. Linear regression of developmental rate for temperatures 15–30 °C indicated that 222 degree-days were required above a minimum threshold of 7.16 °C to complete development. A temperature of 37.6 °C was determined to be the upper development threshold with optimal development occurring at 30.5 °C. Demographic parameters were calculated and pseudo-replicates for intrinsic rate of increase (r_m), net reproductive rates (R_o), generation time (T_c), population doubling time (T_d), and finite rate of increase (λ) were generated using the bootstrap method. Mean bootstrap estimates of demographic parameters were compared across temperatures using ANOVA and nonlinear regression.     

Effect of mating frequency on fecundity and longevity of the predatory mite <Emphasis Type="Italic">Kampimodromus aberrans</Emphasis> (Acari: Phytoseiidae)

The effect of single versus multiple mating on longevity and fecundity as well as the number of matings required to maximize a female’s reproductive success of the predatory mite Kampimodromus aberrans Oudemans were studied under laboratory conditions. Newly emerged adult females of the stock colony of K. aberrans were placed individually on a bean leaf disc, and maintained at 25°C and 16:8 LD. A young male remained with a female for limited periods or continuously. Mating was a requisite for oocyte maturation and oviposition. Females which mated three to four times during their life and females in continuous presence of males, laid significantly and considerably more eggs than single-mated females. Virgin females lived the longest, and those in continuous presence of males the shortest. In all cases and irrespective of the number of matings, the sex ratio of the offspring was male-biased in the first three to four days of oviposition period, and female-biased in later days.     

Effect of temperature and sperm supply on the reproductive potential ofTetranychus evansi (Acari: Tetranychidae)

The biology ofTetranychus evansi Baker and Pritchard was studied at five different temperatures on excised leaves ofSolanum douglasii Dunal. The theoretical lower threshold temperature of development was ca. 13°C. The combined immature stages required ca. 148 day-degrees for completion. Progressively later initiation of mating resulted in higher peak proportions of females produced, up to an 8-day period between emergence and mating at 30°C. The proportion of male offspring always increased towards the end of the oviposition period, apparently because of sperm depletion. The calculated intrinsic rate of increase was highest (0.432) at 35°C, the highest temperature tested. At that same temperature, daily oviposition rate and net reproductive rate were maximum (9.5 and 111.77, respectively), and mean generation time and doubling time were minimum (10.92 and 1.6 days, respectively).     

Environmental influences on fecundity, egg viability and egg cannibalism in the scale insect predator, Chilocorus nigritus

The effects of temperature on age-specific fecundity, egg viability and egg cannibalism were examined in the coccidophagous coccinellid, Chilocorus nigritus being fed on the cyanophyllum scale, Abgrallaspis cyanophylli. There were no significant differences in lifetime fecundity or oviposition rate at constant temperatures in the range of 20 to 30°C although there was a trend for both parameters to increase with increasing temperatures. Total fecundity ranged from five to 1890 eggs per female whilst the oviposition rate varied from 0.2–12.1 eggs day^-1. Egg viability was significantly greater at 26 and 30°C (63 and 74%, respectively) than at lower temperatures (18 to 30%) whilst egg cannibalism was highest at 24 and 26°C (being the mid range of the temperatures tested). Mean pre-oviposition periods varied from 54 days at 20°C to 8.2 days at 30°C. The effect of relative humidities in the range of 33% to 75% on oviposition rate, egg viability and egg cannibalism was measured and found to have no significant effect on either parameter.     

Life-history traits of the acarophagous lady beetle,<Emphasis Type="Italic">Stethorus japonicus</Emphasis> at three constant temperatures

Stethorus japonicusKamiya (Coleoptera: Coccinellidae) is an indigenous ladybird beetle in Japan, which feeds on many spider mite species. We evaluated the development, survivorship and life-history parameters of this lady beetle on a diet of eggs of the two-spotted spider mite, Tetranychus urticae Koch (red form) (Acari: Tetranychidae). In addition, the effect of short photoperiod on its reproduction was assessed. Survival rates from egg to adult were more than 71% at temperatures between 17.5 and 30 °C. The highest immature mortality was 100% at 35 °C followed by 76% at 15 °C and 52% at 32.5 °C. The lower threshold temperature for development from egg to egg-laying adult was 13.0 °C and the thermal constant was calculated as 238.7° days. Based on these data, the maximum number of generations that could complete development in a year under field conditions in Ibaraki, central Japan, would be between five and seven. The intrinsic rates of natural increase (r_m) were 0.093 at 20 °C, 0.156 at 25 °C and 0.241 at 30 °C. Reproductive diapause was induced at photoperiods with light phases shorter than 13 h at 18 °C.     

A comparison of the reproductive ability of <Emphasis Type="Italic">Varroa destructor</Emphasis> (Mesostigmata:Varroidae) in worker and drone brood of Africanized honey bees (<Emphasis Type="Italic">Apis mellifera</Emphasis>)

Colony infestation by the parasitic mite, Varroa destructor is one of the most serious problems for beekeeping worldwide. In order to reproduce varroa females, enter worker or drone brood shortly before the cell is sealed. To test the hypothesis that, due to the preference of mites to invade drone brood to reproduce, a high proportion of the mite reproduction should occur in drone cells, a comparative study of mite reproductive rate in worker and drone brood of Africanized honey bees (AHB) was done for 370 mites. After determining the number, developmental stage and sex of the offspring in worker cells, the foundress female mite was immediately transferred into an uninfested drone cell. Mite fertility in single infested worker and drone brood cells was 76.5 and 79.3%, respectively. There was no difference between the groups (X ² = 0.78, P = 0.37). However, one of the most significant differences in mite reproduction was the higher percentage of mites producing viable offspring (cells that contain one live adult male and at least one adult female mite) in drone cells (38.1%) compared to worker cells (13.8%) (X ² = 55.4, P < 0.01). Furthermore, a high level of immature offspring occurred in worker cells and not in drone cells (X ² = 69, P < 0.01). Although no differences were found in the percentage of non-reproducing mites, more than 74% (n = 85) of the mites that did not reproduce in worker brood, produced offspring when they were transferred to drone brood.     

The influence of temperature and resource level on the fecundity of a predatory planktonic mite,Piona exigua Viets

Summary Piona exigua Viets is a predaceous freshwater mite that can potentially affect the population densities of its cladoceran prey. As part of a study of the effect of Piona exigua on its prey populations we measured the effects of water temperature and prey density on the lifespan, age at first reproduction and per capita egg production of adult female mites. Mites were raised in the laboratory at five prey (Ceriodaphnia, Daphnia) densities (5, 15, 30, 60, 120/l) at 15° C and at four temperatures (10, 15, 18, 22° C) at c. 60 prey/l. In response to increased food level, mites increased the number of eggs laid to reach a maximum at 60 prey/l, the rate at which they were laid increased and the pre-reproductive period was shorter. Low temperatures prolonged the pre-reproductive period. At temperatures above 10° C, food level and temperature had more influence on the timing of reproductive events (growth rate, oviposition rate, age at first reproduction) than on the size of females at first reproduction. When temperatures exceed 10° C and food levels exceed 5 prey/l the major scope for reproductive plasticity in Piona lies in the timing and duration of egg production.