The Effects of Clofentezine on Life-table Parameters in Two-spotted Spider Mite Tetranychus urticae

Sublethal effects of the growth inhibitor, clofentezine, on life-table parameters of Tetranychus urticae Koch females treated at different developmental stages with a concentration causing ≥90% mortality were investigated. Females which survived treatment as ‘early’ (0–24 h old) eggs produced 12% more offspring than the untreated females during the first five days of oviposition. This resulted in a significant rise in the intrinsic rate of increase (r _j ): 0.324, compared to 0.299 in the untreated females. This effect may be interpreted as hormoligosis. Clofentezine treatment at any other developmental stage of T. urticae significantly decreased both longevity and fertility of female survivors. Females which survived treatment either as ‘late’ (72–96 h old) eggs or larvae had 2.6 times lower net reproductive rate (R ₀) than the untreated females, and the r _j values were significantly lower: 0.242 and 0.215, respectively (0.285 in the untreated females). Females which survived treatment either as protonymphs or deutonymphs had 3.9 times and 6 times lower R ₀, respectively. Corresponding r _j values were 0.178 and 0.146, respectively (0.247 in the untreated females). The clofentezine treatment at all stages influenced the age distribution of survivors. The sublethal effects of clofentezine and their impact on T. urticae management are discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Sublethal Effects of Tebufenpyrad on the Eggs and Immatures of Two-Spotted Spider Mite, <Emphasis Type="Italic">Tetranychus urticae</Emphasis>

Sublethal effects of the mitochondrial electron transport inhibitor (METI) tebufenpyrad on Tetranychus urticae Koch females surviving treatment as eggs or immatures (≥90% mortality) were investigated in life-table assay. The developmental time of females that had survived treatment as eggs (2 mg/l) or larvae (2.5 mg/l) was 1 day longer, while that of protonymphs (2.5 mg/l) or deutonymphs (4 mg/l) was 2 days longer, in addition to reduced longevity and fertility, compared to control. The treatment significantly reduced the intrinsic rate of increase in female survivors: corresponding values were 0.258, 0.278, 0.207 and 0.209, respectively (0.307 in untreated females). The offspring age distribution was significantly affected by the treatment. Sublethal effect of tebufenpyrad and its impact on T. urticae management are discussed.     

Potential of an indigenous strain of the entomopathogenic fungus Beauveria bassiana as a biological control agent against the Red Palm Weevil, Rhynchophorus ferrugineus

The potential of a strain of Beauveria bassiana (Ascomycota: Clavicipitaceae) obtained from a naturally infected Rhynchophorus ferrugineus (Coleoptera: Curculionidae) pupa as a biological control agent against this weevil was evaluated both in the laboratory and in semi-field assays. Laboratory results indicate that this strain of B. bassiana can infect eggs, larvae and adults of R. ferrugineus (LC₅₀ from 6.3 × 10⁷ to 3.0 × 10⁹ conidia per ml). However, mortality was not the only indicator of treatment efficacy because adults of either sex inoculated with the fungus efficiently transmitted the disease to untreated adults of the opposite sex, with male-to-female and female-to-male rates of transmission of 55% and 60%, respectively. In addition, treatment with B. bassiana significantly reduced fecundity (up to 62.6%) and egg hatching (32.8%) in pairing combinations with fungus-challenged males, females or both sexes. Likewise, 30-35% increase in larval mortality was observed in larvae obtained from eggs from fungus-challenged females or from untreated females coupled with inoculated males, resulting in an overall 78% progeny reduction. Semi-field preventive assays on potted 5-year old Phoenix canariensis palms, with efficacies up to 85.7%, confirmed the potential of this strain as a biological control agent against R. ferrugineus.     

Reproductive biology of Fidiobia dominica (Hymenoptera: Platygastridae), an egg parasitoid of Diaprepes abbreviatus (Coleoptera: Curculionidae)

The reproductive biology of Fidiobia dominica Evans (Hymenoptera: Platygastridae) was studied in the laboratory (25.6 +/- 1 degrees C) using host eggs of Diaprepes abbreviatus L. (Coleoptera: Curculionidae). F. dominica readily parasitized D. abbreviatus eggs on both host plant and wax paper substrates. The number of egg masses parasitized and the number of offspring produced were higher when females were offered more than one host egg mass but did not differ when either two or three egg masses were offered. Female parasitoids that were provided with host eggs and a honey food source lived significantly longer than those that were not provided a food source; however, they did not parasitize more D. abbreviatus eggs. Oviposition occurred in host eggs from 0 to 7 d old, and host mortality was relatively consistent for eggs 0-5 d old and lower for eggs 6-7 d old. Successful parasitoid emergence seldom occurred after host eggs were 4 d old, and by 7 d, no adults successfully emerged. Developmental time from egg to adult was 19.3 +/- 0.2 d for males, significantly more rapid than the females (20.4 +/- 0.1 d). The mean longevity of adult females was 8.0 +/- 0.4 d, with a mean oviposition period of 2.7 +/- 0.3 d; males survived 8.1 +/- 0.4 d. The demographic parameters including intrinsic rate of increase (r(m)), generation time (T), and net reproduction (R(o)) were 0.142/d, 22 d, and 22.4 female eggs/d, respectively.     

Clofentezine and fenbutatin oxide resistance in the two-spotted spider mite, Tetranychus urticae Koch (Acari: Tetranychidae) from deciduous fruit tree orchards in Western Australia

The resistance status of Tetranychus urticae Koch to the miticides clofentezine and fenbutatin oxide was investigated in deciduous fruit tree orchards in Western Australia. Mites were collected from 30 orchards across three fruit growing regions and subjected to discriminating–dose assays. There was a significant relationship between clofentezine (=;0.64) and fenbutatin oxide (V=;0.44) use and the proportion of resistant mites in each sample. Improved clofentezine models were obtained when the Perth Hills region (V=;0.90) was modelled separately from Donnybrook and Manjimup (V=;0.80). Clofentezine could be expected to fail in the latter regions after four sprays and in the former region after only one spray. The relatively low pest status of T. urticae in Western Australia did not delay the clofentezine resistance. For any given number of spray applications the response of T. urticae to fenbutatin oxide was more variable than it was to clofentezine.     

Temperature and diet effects on immature development of predatory mite Typhlodromus athenas Swirski and Ragusa (Acari: Phyotseiidae)

Temperature and food quality both can influence growth rates and developmental time of herbivorous insects and mites. Typhlodromus athenas Swirski and Ragusa is an indigenous mite in the Mediterranean region and data on its temperature dependent development are lacking. In the current study, temperature-dependent development and survival of T. athenas immature stages were evaluated on eggs and all stages of Tetranychus urticae Koch, as well as on almond (Prunus amygdalis Batsch) pollen, under seven constant temperatures ranging from 15 to 35°C, 65% RH, and a photoperiod of 16:8 (L:D) h. On both diets survival was considerably high at all temperatures. The longest developmental period of immature stages was recorded at 15°C, whereas the shortest was at 30°C. Female immatures on almond pollen had shorter developmental time compared to that on twospotted spider mites. Food had a significant effect on female total developmental time at temperatures lower than 25°C. The lower developmental thresholds, estimated by a linear model, for egg-to-adult of females and males fed on pollen were 8.60 and 8.77°C, respectively, whereas on T. urticae they were 10.15 and 10.62°C, respectively. Higher values of tmin for total development were estimated by a nonlinear model (Lactin-2), and ranged from 10.21°C for both females and males on almond, to 11.07 for females and 10.78°C for males on prey. Moreover, this model estimated optimal and lethal temperatures accurately. The results of this study indicate that T. athenas appears better adapted to higher temperatures that occur in the Mediterranean region and may be a useful biological control agent.     

Euseiusfinlandicus (Acari: Phytoseiidae) as a potential biocontrol agent against Tetranychus urticae (Acari: Tetranychidae): life history and feeding habits on three different types of food

Life history and reproductive parameters of the generalist predatory mite Euseius (Amblyseius) finlandicus (Oudemans) were studied in the laboratory at 25 +/- 1 degrees C, with a 16L:8D photoperiod and 60 +/- 15% RH, to investigate its response to different food sources: an eriophyid mite Aceria sp., tulip pollen Tulipa gesnerana L., and two-spotted spider mite Tetranychus urticae Koch. Total developmental time of the immature stages was the shortest on eriophyid mites, followed by pollen, and then spider mites. Fecundity was highest on pollen (43.69 eggs; 1.63 eggs/female/day), then eriophyid mites (39.73 eggs; 1.37 eggs/female/day) and lowest on spider mites (18.16 eggs; 0.80 eggs/female/day). Intrinsic rate of increase (Rm), net reproductive rate (Ro) and finite rate of increase (lambda) followed the same pattern [pollen (0.168, 27.96 and 1.183, respectively), eriophyid mites (0.153, 20.81 and 1.167), spider mites (0.110, 9.44 and 1.119)]. Mean generation time (days) was the shortest on pollen (19.90), followed by eriophyid mites (20.02), and then spider mites (20.59). Average spider mite larvae consumed by E. finlandicus during immature stages were 9.18 for males and 11.85 for females. Adult E. finlandicus females consumed an average of 166.38 spider mite protonymphs during adult stage compared to an average of 66.55 by males. The number of prey protonymphs consumed per day by females was highest in the oviposition period, lower in the pre-oviposition period and the lowest in the post-oviposition period. The eriophyid mite as a prey recorded the shortest developmental time, while pollen as food recorded the highest oviposition rate in E. finlandicus. The potential of this predator as a biocontrol agent against T. urticae is discussed.     

Biology and field performance of gryon clavigrallae (Hymenoptera: Scelionidae), an egg parasitoid of Clavigralla spp. (Hemiptera: Coreidae) in India

The biology and impact of Gryon clavigrallae Mineo, an egg parasitoid of Clavigralla scutellaris Spinola and C. gibbosa (Westwood), was investigated. The calculated developmental threshold temperatures for females and males were 15.6 degrees C and 15.8 degrees C, respectively. Emergence exceeded 94% at temperatures between 22 and 30 degrees C. Adult females lived on average 28-96 days when fed with honey. Without food, adults lived < 6 days. Mean fecundity was 56.4 eggs per female. A significant trend of lower fecundity after longer periods of host deprivation was observed. Gryon clavigrallae females successfully oviposited in host eggs of all ages though eggs < 4 days old were preferred. Total host handling times were significantly longer on C. gibbosa eggs (23.5 min) than on C. scutellaris eggs (12.0 min). Females readily distinguished parasitized from non-parasitized host eggs. Superparasitism was observed when few or no unparasitized eggs were available. Eggs of the two Clavigralla species can be separated by surface structure and condition after eclosion. Clavigralla scutellaris laid significantly larger egg clusters than C. gibbosa (19.9 versus 10.5 eggs per cluster). Gryon clavigrallae was present as soon as the first Clavigralla spp. egg clusters were found on pigeonpea. The percentage of egg clusters parasitized increased early in the season with egg cluster density and remained high (up to 83%) despite fluctuations in host density. Overall, G. clavigrallae parasitized 40 and 58% of C. gibbosa and C. scutellaris eggs. The percentage of egg clusters parasitized and the number of eggs parasitized per cluster increased significantly with egg cluster size. The overall parasitoid sex ratio was highly female biased but varied with the number of eggs parasitized per cluster.     

Influence of temperature on cyhexatin resistance inTetranychus urticae (Acari: Tetranychidae)

The effect of post-treatment temperature on the toxicity of cyhexatin to cyhexatin-susceptible (S) and resistant (R) strains ofTetranychus urticae Koch was investigated. Females and developmental stages of both strains showed a positive temperature-toxicity coefficient with wettable powder (WP) and emulsifiable concentrate (EC) formulations of cyhexatin. Females of the S strain were 3.0× and 2.9× less susceptible at 15°C than at 28°C with the EC and WP formulations respectively, while females of the R strain were 5.2× and 23.6× less susceptible at 15°C with these formulations.     

Biological, Chemical and Integrated Control of Two-spotted Spider Mite Tetranychus urticae on Dwarf Hops

Chemical, biological and integrated programmes for the control of two-spotted spider mite, Tetranychus urticae, were compared on dwarf hops in 1997 and 1998. In both years integrated control, which consisted of an application of the ovicidal acaricide clofentezine followed by a release of the predatory mite Phytoseiulus persimilis at 10 individuals per plant, was the most effective treatment. Similar numbers of spider mites were recorded on plots that were treated with P. persimilis only or with a single application of the acaricide tebufenpyrad, and plots with either of these treatments had higher levels of infestation than the plots under integrated control. The highest numbers of spider mites were found on the untreated plots. This study indicates that integrated control of T. urticae using clofentezine in conjunction with P. persimilis is likely to be more effective than an approach based on chemical or biological measures only. It is suggested that an integrated system would have the added benefit over a pesticide-only programme of reducing pressure on the pest for the selection of strains resistant to acaricides.     

Female Mate Choice in a Paternal Brooding Blenny: The Process and Benefits of Mating with Males Tending Young Eggs

Female mate preference for males tending young offspring has been demonstrated in many fishes; however, not much is known about the choice process. Using the barred chin blenny Rhabdoblennius ellipes, a fish with male uniparental care, field experiments were conducted to investigate the female preference for males tending young eggs and then whether females choose the males with young eggs by discriminating young eggs from old eggs in the nests. Males tending young eggs (0‐ to 2‐d old) acquired new eggs nine times more frequently than those tending old eggs (3‐ to 5‐d old) regardless of other traits in males and nests. In the two egg‐switching field experiments (old to young and young to old), contrary to our expectation, male mating success was neither enhanced when given young eggs nor inhibited when given old eggs. These results suggested that females choose males with young eggs not by discriminating the developmental stage of eggs in the nests but by using other choice processes. By choosing males with young eggs, females may benefit from the dilution effects of egg predation and filial cannibalism risks and avoid male parental care failure.     

Effect of multiple mating on the reproductive performance of the rice leaffolder moth,Cnaphalocrocis medinalis (Lepidoptera: Crambidae)

To clarify whether multiple mating of females and males affects the reproductive performance of the rice leaffolder moth, Cnaphalocrocis medinalis (Guenée), we examined the effect of the number of matings (once, twice, or three times) for females (female treatment) and males (male treatment) on the incidence of moth mating, number of eggs laid, egg hatchability, and adult longevity. We also compared the effect of multiple mating imposed on males or females separately with the effect of that imposed on both sexes simultaneously (both sexes treatment). The incidence of mating of females and males that mated three times (3-mated females and males) was significantly lower than for females and males that mated twice or once (2-mated or 1-mated females and males). The incidence of mating of 1-mated moths (both sexes) was significantly higher than for 2-mated or 3-mated moths (both sexes). Two-mated or 3-mated females laid significantly more eggs with significantly higher hatchability than 1-mated females. Females that mated with 1-mated males (second male mating) or 2-mated males (third male mating) laid significantly fewer eggs than those that mated with virgin males (first male mating). Females laid significantly more eggs after the second and third matings for moths of both sexes than after the first mating for moths of both sexes. The mechanisms of improvement and decline of female reproductive performance when multiple mating was imposed on males or females are also discussed in relation to the reproductive biology of C. medinalis.     

Effects of mating rates on oviposition,sex ratio and longevity in a predatory mite <Emphasis Type="Italic">Neoseiulus cucumeris</Emphasis> (Acari: Phytoseiidae)

Two aspects of mating effects on the fecundity, sex ratio and longevity of Neoseiulus cucumeris (Acari: Phytoseiidae) were examined in laboratory experiments: (1) females mated by one, two or three different males (unmated and 3 days old) at 5-day intervals, and (2) females mated by males with different age/mating status (number of females mated previously by the male). Females allowed to mate with a second or third male at 5-day intervals produced 39 eggs on average, but those mated with a single male produced 28 eggs on average. Matings with additional males 5 or 10 days after the first male increased the duration of the oviposition period of these females by 5–7 days and at the same time reduced the post-oviposition period by about 10 days. Overall, females with additional matings by one or two different males at 5-day intervals survived a few days shorter than females without additional males. Mating with a different female each day, a male of N. cucumeris could mate with 5–8 females, which produced a total of 85–116 eggs: females mated with a male during days 1 and 2 in its adulthood and with a male of the last 2 days of life (days 7 and 8) produced about half as many eggs as females mated with a male during 3–6 days of its adulthood. Females mated with males that are too young or too old had a shorter oviposition period and a longer post-oviposition period and longevity than females mated with middle-aged males. In both experiments, rates of oviposition remained similar in females with high or low fecundity. This indicates that in both cases, the increased fecundity is due to the extension of the oviposition period through additional sperm supplied by the second male and or third male (in experiment 1) or more sperm by males not too young nor too old (experiment 2).     

Maternal age and egg weight affect offspring performance in the predatory stink bug Podisus nigrispinus

To investigate influences of maternal age and egg weight on developmental time and body weight of offspring in the Neotropical pentatomid Podisus nigrispinus, a study was carried out using large (0.36 mg) and small (0.32 mg) eggs of young (two-week- old) females and large eggs of old (seven-week-old) females. Incubation period of eggs was influenced only by egg weight, i.e. small eggs took a relatively shorter time to hatch. Both maternal age and egg weight had significant effects on total nymphal and egg-to- adult developmental times. Overall, offspring of young parents had shorter developmental times. None of the developmental periods were significantly affected by sex. No significant differences were found among various offspring groups for percentages of egg hatch, nymphal survival, and sex ratio. Variation of egg weight in relation to the number of eggs per day observed within the first month of life suggests a trade-off between weight of individual eggs and oviposition rate. It is concluded that for mass production of the predators in view of their use in augmentative biocontrol programmes, young parents (2–4 weeks old) should be preferred.     

Predation and oviposition by predatory Stethorus japonicus, Oligota kashmirica benefica, and Scolothrips takahashii in egg patches of various spider mite species

Predation and oviposition by three predacious insects [ Scolothrips takahashii Priesner (Thysanoptera: Thripidae), Stethorus japonicus H. Kamiya (Coleoptera: Coccinellidae), and Oligota kashmirica benefica Naomi (Coleoptera: Staphylinidae)] were examined using egg patches of three spider mite species [ Amphitetranychus viennensis (Zacher), Tetranychus urticae Koch, and Panonychus mori Yokoyama (all Acari: Tetranychidae)] that are pest species on Japanese pear [ Pyrus serotina Rehder (Rosaceae)]. Scolothrips takahashii females consumed more prey and laid more eggs in A. viennensis and T. urticae egg patches than in P. mori egg patches. Females also left P. mori egg patches sooner than they left other egg patches even at high prey density. Predation and oviposition of S. takahashii in a prey patch were greatly enhanced by the complicated webs produced by A. viennensis . Stethorus japonicus females consumed significantly more prey and laid more eggs in A. viennensis egg patches than in P. mori egg patches, with intermediate values in T. urticae egg patches, reflecting the quality of each spider mite species as prey. Oligota kashmirica benefica females consumed more prey and laid more eggs in A. viennensis and T. urticae egg patches than in P. mori egg patches, particularly at high prey density. However, predation and oviposition by O. kashmirica benefica increased greatly with increasing prey density, even in P. mori egg patches, indicating that prey density was the most important factor in predation and oviposition by this species. These results are discussed in relation to the potential effect of each predator on the suppression of different spider mite species on Japanese pear.     

Repeated copulation in the brown planthopper, Nilaparvata lugens Stál (Homoptera; Delphacidae)

Abstract. 1. Mating of the brown planthopper, Nilaparvata lugens , was investigated in relation to oviposition. Females became unreceptive immediately after mating and showed various types of repelling behaviour to courting males. As a result, females usually did not mate repeatedly in quick succession, but after ceasing to lay fertilized eggs they behaved as virgins and mated again before producing more fertilized eggs.
2. Copulation, followed by deposition of fertilized eggs, occurred twice, or in a few cases three times, throughout the adult stage. Copulation lasted about 2 min at the first mating and about 1 min at the second or third mating.
3. When the number of fertilized eggs began to decrease rapidly, oviposition rate also decreased, but it increased again immediately after re-mating. Repeated copulation was also related to the potential rate of population increase. In the present experiments, the estimates of net reproductive rate ( R ₀) in brachypterous females was 287.7 (first mating), 88.3 (second mating) and 7.2 (third mating) and in macropterous females 286.8, 120.6 and 3.6 respectively.
4. In a patchy environment, repeated copulation following an invasion may contribute to the increase of a population.     

Effects of neem seed derivatives on behavioral and physiological responses of the Cosmopolites sordidus (Coleoptera: Curculionidae)

Both in a choice and multi-choice laboratory tests, fewer adults of the banana root borer, Cosmopolites sordidus (Germar), settled under the corms of the susceptible banana "Nakyetengu" treated with 5% aqueous extract of neem seed powder or cake or 2.5 and 5% emulsified neem oil than on water-treated corms. Feeding damage by larvae on banana pseudostem discs treated with 5% extract of powdered neem seed, kernel, or cake, or 5% emulsified neem oil was significantly less than on untreated discs. The larvae took much longer to locate feeding sites, initiate feeding and bore into pseudostem discs treated with extract of powdered neem seed or kernel. Few larvae survived when confined for 14 d on neem-treated banana pseudostems; the survivors weighed two to four times less than the larvae developing on untreated pseudostems. Females deposited up to 75% fewer eggs on neem-treated corms. In addition, egg hatching was reduced on neem-treated corms. The higher the concentration of neem materials the more severe the effect.     

Laboratory Evaluation of a Laboratory-selected Organophosphate-resistant Strain of Amblyseius finlandicus (Acari: Phytoseiidae) for Possible Use in Finnish Apple Orchards

The survival of adults and immatures, and reproduction at rates of 150, 300 and 900 ppm a.i. of azinphosmethyl were compared among four types of Amblyseius finlandicus females and their offspring; a laboratory-selected colony (SEL16) resistant to azinphosmethyl, an unselected base colony (Tuorla), reciprocal F1 females from crosses between SEL16 and a very susceptible colony (Hiekkaharju). The SEL16 and reciprocal F1 females had a higher survivorship than the unselected Tuorla base colony after 2 and 4 days on leaves treated with the three concentrations of insecticide. Azinphosmethyl decreased oviposition by the surviving females of all types, but the decrease was smaller in the SEL16 strain than in the base colony. The survivorship of immatures of SEL16 on fresh azinphosmethyl residues after 6 days was low at all three concentrations, but no survivors were found in the base colony at any of the concentrations. Females deposited significantly more eggs on the untreated areas of the leaves. The importance of this behavior for biological control is discussed. There were no significant differences in developmental rate, ovipositional rate, sex ratio and ovipositional period between the SEL16 and unselected base colony on untreated leaf substrates, suggesting that resistance was not associated with decreased fitness. A mode of inheritance study indicated that azinphosmethyl resistance was incompletely dominant. This was also suggested by the results of bioassays, where the reciprocal F1 females performed nearly as well as the SEL16 strain on leaves treated with field rates of azinphosmethyl.     

Predation and life table of Feltiella acarisuga (Diptera: Cecidomyiidae) preying on eggs of Tetranychus urticae (Acari: Tetranychidae)

Feltiella acarisuga (Vallot) is a common gall midge that feeds on many species of spider mites. Predation and all major biological and life table parameters of F. acarisuga were determined using eggs of the twospotted spider mite, Tetranychus urticae (Koch), as prey under laboratory conditions (26.7 degrees C, 85% RH, and 14:10 [L:D] h). First-, second-, and third-instar F. acarisuga larvae consumed an average of 37.8, 60.4, and 87.2 T. urticae eggs per day, respectively. Larvae consumed a total of 185.4 eggs each from first to third instars, and female larvae consumed 201.2 spider mite eggs, 18.6% more than male larvae. Developmental time of F. acarisuga was 2.6, 7.8, and 7.0 d for eggs, larvae, and pupae, respectively, with an average of 17.4 d from oviposition to adult emergence. Female immatures developed slightly slower ( approximately 1 d) than males. Adult longevities of F. acarisuga averaged 13.0 d. After an average of 1.1-d preoviposition period, each female laid an average of 33.3 eggs during her lifespan. The intrinsic rate of natural population increase (r), net reproductive rates (R(o)), gross reproductive rate (Sigmam(x)), generation time (T), doubling time (DT), and the finite rate of increase (lambda) of F. acarisuga were estimated by life table analysis at 0.1155/d, 15.25 eggs per female, 21.69 eggs per female, 23.6 d, 6.0 d, and 1.1225/d, respectively. Because the r value of F. acarisuga is relatively lower than that of T. urticae, frequent release of F. acarisuga may be necessary to suppress populations of T. urticae.     

Reproductive behavior and spawning success of female Amblyglyphidodon leucogaster (Pisces: Pomacentridae) from the Red Sea

The reproductive behavior of female whitebelly damselfish, Amblyglyphidodon leucogaster, was investigated in the Gulf of Aqaba, Red Sea over two breeding seasons. Females were promiscuous, mating with 7–10 different males throughout the season. Females lay eggs in distinct batches, defined as the total number of eggs laid in a day. Generally females deposit a batch of eggs with one male (87.2%) and are capable of laying a new batch every other day. Egg batch size averaged 4009 eggs and females laid from 2 to 22 egg batches per season. The variation in spawning success was not correlated to body size. Females preferred to deposit eggs in nests that already contained early stage eggs (0–2 days old). Within a nest, females chose to lay eggs contiguous to the youngest egg batch, regardless if the nest contained either a single batch or multiple batches of different ages. Female within-nest spawning patterns appear to be a consequence of between nest preferences for nests with young eggs. It is proposed that the strong within-nest preference is a consequence of mate selection where females may use new egg batches as a visual cue as part of a copying style. Such a style may reduce the risk of predation and increase feeding opportunities, because less time is expended in mate selection, which would provide additional resources for egg production and ultimately increase female spawning success over the breeding season. This revised version was published online in July 2006 with corrections to the Cover Date.