Selection of Bemisia nymphal stages for oviposition or feeding,and host-handling times of arrhenotokous and thelytokous <Emphasis Type="Italic">Eretmocerus mundus</Emphasis> and arrhenotokous <Emphasis Type="Italic">E. eremicus</Emphasis>

Host-handling behavior is an important aspect of parasitoid foraging behavior. When a parasitoid encounters a potential host, the handling behavior starts with the evaluation of the host and continues if the host has been judged acceptable. Host handling is usually terminated after egg laying or host feeding and host marking. Host-handling behavior of an arrhenotokous population of two Eretmocerus species, E. mundus Mercet and E. eremicus Rose and Zolnerowich, along with a thelytokous population of E. mundus were compared under laboratory conditions. Several elements of host-handling behavior, including encountering, ascending, turning on host, descending, preening, egg laying, and host feeding were recorded. There were no correlations among the durations of these phases across parasitoid populations/species or host nymphal instars. Duration of different phases of host-handling behavior showed only slight and sometimes significant differences between different Eretmoceruspopulations/species. The actual laying of the egg had the longest duration of all host-handling behaviors, and was longer on third nymphal instars than on younger ones. Females of the three populations/species accepted the first three nymphal stages either for egg laying or for host feeding. Females spent a lot of time to make wounds in the host when preparing for host feeding, and eventually killed the host. The implications of these findings for the use of the different Eretmoceruspopulations/species in biological control are discussed.     

Oviposition site selection in Cactoblastis cactorum (Lepidoptera): constraints and compromises

Summary Oviposition by Cactoblastis cactorum on Opuntia ficus-indica and O. aurantiaca was assessed from the positioning of egg sticks on plants in the field. The number of egg sticks laid on O. ficus-indica plants was affected by: (1) plant size; (2) moth emergence near the plant; (3) cladode condition; and (4) plant conspicuousness. These factors contributed towards the clumping of egg sticks on plants. There was no apparent oviposition preference for one of the two host plant species despite the fact that egg predation was higher and fecundity lower on O. aurantiaca. The selection of a site for oviposition on the host plants was influenced by: (1) cladode condition; (2) height above ground; and (3) shelter from wind during oviposition. Succulent cladodes were the favoured sites for oviposition. The evidence suggests that in C. cactorum, oviposition site selection is largely the net result of a compromise between oviposition behaviour selected for increasing the probability of juvenile survival and oviposition behaviour selected for increasing the probability of laying the full complement of eggs. In addition, environmental and physiological factors such as wind and wing-loading, are thought to place constraints on the range of sites available for oviposition.     

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

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

Genetic variation for rate of development in natural populations of Drosophila melanogaster

We have sampled wild chromosomes from two natural populations of Drosophila melanogaster and obtained flies fully homozygous for the second chromosome, the third chromosome, or both, as well as flies heterozygous for one or both wild chromosomes and balancer chromosomes. Rate of embryogenesis (egg laying to larval hatching) and rate of development from egg to adult are measured, by classifying the individuals into fast, intermediate, and slow developmental classes. The experiments indicate that variation for rate of embryogenesis and for rate of egg-to-adult development is plentiful in the natural populations. Various hypotheses are enunciated to account for the small range of phenotypic variation observed in wild-type individuals with respect to the two parameters (embryogenesis and egg-to-adult development) and for the difficulty in changing the mean rates by artificial selection. Appropriate experiments may decide among the hypotheses, helping us to understand the genetic control of rate of ontogenesis, which is an important fitness component.     

Identifying reproductive events using archival tags: egg‐laying behaviour of the small spotted catshark Scyliorhinus canicula

The use of archival depth telemetry as a means of remotely assessing the reproductive rates of free‐ranging fishes is explored. This is achieved by electronically tracking the vertical movements of individual female small spotted catsharks Scyliorhinus canicula in the natural environment, whilst simultaneously evaluating the temporal and vertical distributions of egg‐laying in this species. Distinctive patterns of short‐term (0·3–3·7 h), shallow‐water activity are documented in the time–depth profiles of female S. canicula that occur at an appropriate depth (1·0–2·3 m) and periodicity (every 10–12 days) to represent egg‐laying behaviour. Putative egg‐laying behaviour was exhibited simultaneously by two individually tracked female S. canicula during late‐spring and early‐summer. The results highlight that, provided species behaviour is suitable and complementary methods such as previous data, laboratory experiments and field surveys can be used to validate the patterns observed, archival depth telemetry offers an unobtrusive means by which egg production and egg‐laying behaviour of free‐living fishes can be estimated. As precise information regarding life‐history parameters is difficult to obtain for free‐ranging fish species, this technique could be used to improve the parameterization of species demographic models that are relevant to the management of wild fish populations.     

共生菌通过氧气浓度调控果蝇产卵偏嗜性

【目的】研究东方醋杆菌(Acetobacter orientalis)和短小乳杆菌(Lactobacillus brevis)通过氧气浓度调控黑腹果蝇(Drosophila melanogaster)产卵的偏嗜性,并研究其对后代存活率和发育历期的生物学意义。【方法】利用双相选择装置解析果蝇产卵行为。利用分光光度仪和pH计测量细菌的OD值与pH值。通过改变果蝇的视觉、味觉和嗅觉感官以检测介导果蝇产卵行为的感觉系统。用后代的发育历期与存活率评价其生物学意义。【结果】果蝇产卵对东方醋杆菌具有趋避性,产卵指数为–0.76,对短小乳杆菌具有趋向性,产卵指数为0.5,并且二者接种比例决定了果蝇的产卵选择。氧气作为一个调节因子,可改变果蝇对东方醋杆菌和短小乳杆菌的产卵偏嗜性。氧气是东方醋杆菌生长必需的条件,有氧培养的OD值为1.3,而无氧培养的OD值为0.4。氧气是短小乳杆菌生长非必需的条件,有氧和无氧培养的OD值均为2.2。破坏嗅觉可严重地影响果蝇对细菌的产卵偏嗜性。东方醋杆菌和短小乳杆菌均促进了果蝇的生长,分别提高存活率1倍和1.5倍,分别缩短果蝇发育历期1 d和2 d。【结论】东方醋杆菌和短小...     

Experimental comparison of methods for simultaneous selection of two correlated traits in Tribolium

Summary Two lines of Tribolium castaneum were selected in each of three replicates for egg laying between 7 and 11 days after adult emergency and for adult weight at 12 days, using theoretical (IT) and empirical (IP) index selection methods. Index coefficients were given empirically in the IP line and they were adjusted in the successive generations of selection according to the results obtained in the previous ones. Highly repeatable selection responses in all replicates occurred in both lines for the aggregate genotype (egg laying plus adult weight) and for each individual trait. The IP line tended to increase slightly more than the IT line for aggregate genotype and egg laying, while the highest response in adult weight was obtained with the IT method. The two methods gave consistently different responses in each replicate. The expected results were that IT selection should not exceed IP selection for the aggregate genotype and egg laying while theoretically the IT method should have been superior for increase adult weight. Theoretical expectations for adult weight have been fulfilled in practice. The IP method would be preferred in a practical sense because of its simplicity and freedom from need of parameter estimation.     

The effect of temperature,darkness, starvation and various food types on growth,survival and reproduction of Helisoma duryi,Biomphalaria alexandrina and Bulinus truncatus (Gastropoda: Planorbidae)

Helisoma duryi has been proposed as a biological control agent in schistosomiasis due to its superiority in laboratory competition experiments with various species of the intermediate host snails. Therefore it was considered important to evaluate the response of this snail species and the intermediate host species, Biomphalaria alexandrina and Bulinus truncatus, to various physical, chemical and biological factors under laboratory conditions in order to obtain information on the similarities in the ecological niches of these species. The factors considered in the present paper are: temperature, darkness, starvation and food. All three species had optimal growth and egg laying at 26–28 °C. Only H. duryi survived for a longer period at 33°C and it was capable of starting egg laying at this temperature although the onset was delayed. However, low temperature (18°C) caused a relatively larger decrease in egg laying of H. duryi than in the other two species. Growth and egg laying was reduced for H. duryi and B. truncatus kept under darkness and B. alexandrina could not tolerate maintenance under darkness. A few days of starvation of juvenile snails had no effect on later growth and egg laying capacity of the survivors, although mortality in B. truncatus was increased. B. alexandrina had a lower tolerance to starvation than the other two species. Egg laying of snails fed only one of the three laboratory food types decreased for all three species in the order: Vov-vov (dog food in dry pellets), Tetramin (fish food) and lettuce. Combinations of lettuce and one or more proteinaceous food types gave optimal growth and egg laying for all three species.     

Diet, fecundity and egg size in some polyphagous predatory carabid beetles

Egg size was measured at different rates of egg laying in three polyphagous carabid species, known to be useful predators of cereal aphids; the small Bembidion lampros Herbst and the medium-sized Pterostichus cupreus L. and P. melanarius Illiger. Variations in fecundity, as well as the ability of the medium-sized species to also build up fat reserves, were obtained when beetles were subjected to different dietary regimes consisting of aphids, or foods with a lower or higher protein content. Egg size was found to be dependent on the rate of egg laying within a species. A diet of cereal aphids appeared to be adequate for egg production in these polyphagous carabids, but female P. cupreus were unable to build up fat reserves when they ingested aphids contaminated with the aphicide pirimicarb. Beetles were able to devote resources to more and larger eggs (B. lampros), or to larger eggs and/or fat reserves (P. melanarius/P. cupreus) when given access to a carbohydrate-rich food with low protein content. The highest rate of egg laying was obtained when female P. cupreus and P. melanarius were given a more varied diet at frequent intervals; including regular shifts between unsprayed aphids, carbohydrate-rich food and protein-rich maggots. Within the varied diet treatment a negative relationship was obtained between egg size and egg number among similar-sized individuals of P. cupreus and P. melanarius; females producing the largest number also laid the smallest eggs. Egg size affected larval survival, since first instars hatching from large eggs were found to survive longer than those hatching from small eggs. The influence of differences in food intake on reproduction, maintenance metabolism, and survival of fieldinhabiting carabids is discussed.     

Neuronal input contributes to sequence of Aplysia egg laying behaviors

Egg laying in Aplysia is controlled by the bag cell neuroendocrine system, which releases multiple peptides during a long-lasting electrical discharge. Following the discharge, a fixed sequence of head and neck movements is performed in which two phases can be distinguished: an appetitive or preparatory phase, in which the substrate is prepared, and a consummatory phase, when the egg string is deposited. During egg laying, feeding responses are suppressed. In this study, Aplysia fasciata was used. When the movement of the egg string through the genital groove was prevented by ligation, lesions of the nerve innervating the genital pore completely abolished the consummatory egg-laying behaviors. This shows that a nervous connection between the genital pore area and the central nervous system is important for the consummatory egg-laying behaviors.We found that suppression of feeding responses to seaweed occurred only during the consummatory phase of egg laying in controls, but animals with ligated genital grooves continued to show normal responses to food. It is hypothesized that a neuronal feedback, possibly together with the bag cell peptides, is critical for the temporal organization of egg-laying behavior in Aplysia.Abbreviations CNS central nervous system - ELH egg laying hormone     

Fluctuating Selection,Egg Banks and Population Genetic Structure in Cyclically Parthenogenetic Species

Associations between neutral genetic markers and genes under selection have been suggested to explain the population genetic structure of neutral genes in cyclically parthenogenetic freshwater invertebrates. A simulation model was constructed in order to analyse the extrapolated consequences of observed fluctuations in genotype frequencies in Daphnia, in the presence of egg banks. When of sufficient depth and magnitude, egg banks in combination with fluctuating selection were shown to maintain genetic variation within populations indefinitely. The level of equilibrium diversity increased with the depth and magnitude of the banks, and with intensity of selection. The same threshold was responsible for genetic differentiation between populations, which was independent of migration rate, and which was attained very rapidly following initial Hardy–Weinberg equilibrium. In the absence of selection, egg banks increased the effective size of local populations, thereby decreasing genetic differentiation at migration-drift equilibrium. These results suggest that egg banks are crucial to the genetic structure in the presence of fluctuating selective pressures, but more data are needed if this knowledge is to be used in an improved general understanding of the genetic structure of cyclically parthenogenetic species.     

Thermal acclimation in the fresh water planarians,Dugesia tigrina and D. dorotocephala

The fecundity of two British populations of the freshwater crayfish Austropotamobius pallipes was examined. A number of maternal-size-related variables were correlated with counts of pleopodal eggs per female. Egg numbers did not decrease with increasing time from initial laying. Ovarian egg counts were considerably higher than expected pleopodal egg numbers. The influence of locational parameters is discussed and regional comparisons made.     

Synchronization of laying by great spotted cuckoos and recognition ability of magpies

Brood parasites rely entirely on the parental care of host species to raise the parasitic nestlings until independence. The reproductive success of avian brood parasites depends on finding host nests at a suitable stage (i.e. during egg laying) for parasitism and weakly defensive (i.e. non‐ejector) hosts. Finding appropriate nests for parasitism may, however, vary depending on ecological conditions, including parasite abundance in the area, which also varies from one year to another and therefore may influence coevolutionary relationships between brood parasites and their hosts. In this scenario, we explored: 1) the degree of laying synchronization between great spotted cuckoos Clamator glandarius and magpies Pica pica during two breeding seasons, which varied in the level of selection pressure due to brood parasitism (i.e. parasitism rate); 2) magpie responses to natural parasitism in the pre‐laying period and successfulness of parasitic eggs laid at this stage; and 3) magpie responses to experimental parasitism performed at different breeding stages. We found that, during the year of higher parasitism rate, there was an increase in the percentage of parasitic eggs laid before magpies started laying. However, the synchronization of laying was poor both years regardless of the differences in the parasitism rate. The ejection rate was significantly higher during the pre‐egg‐laying and the post‐hatching stages than during the laying stage, and hatching success of parasitic eggs laid during the pre‐egg‐laying stage was zero. Thus, non‐synchronized parasitic eggs are wasted and therefore poor synchronization should be penalized by natural selection. We discuss four different hypotheses explaining poor synchronization.     

Attractive blue‐green egg coloration and cuckoo−host coevolution

Recent evidence suggests that blue‐green coloration of bird eggshells may be related to female and/or egg phenotypic quality, and that such colour may affect parental effort and therefore the nutritional environment of developing nestlings. Here we suggest that these relationships and the signal function of eggshell coloration would affect the outcome of coevolution between avian brood parasites and their hosts in at least three different non‐exclusive evolutionary pathways. First, by laying blue‐green coloured eggs, cuckoo females may exploit possible sensory biases of their hosts, constraining the evolution of parasitic egg recognition, and thus avoid rejection. Second, because of the relatively high costs of laying blue eggs, cuckoo females may be limited in their ability to mimic costly blue‐green eggs of their hosts because cuckoo females lay many more eggs than their hosts. Furthermore, costs associated with foreign egg recognition errors would be relatively higher for hosts laying blue eggs. Third, cuckoos may use coloration of host eggs for selecting individuals or specific hosts of appropriate phenotypic quality (i.e. parental abilities). We here explored some predictions emerging from the above scenarios and found partial support for two of them by studying egg coloration of European cuckoos (Cuculus canorus) and that of their 25 main hosts, as well as parasitism and rejection rate of hosts. Cuckoo hosts parasitized with more blue, green, and ultraviolet cuckoo eggs, or those laying more blue‐green eggs, were more prone to accept experimental parasitism with artificial cuckoo eggs. In addition, coloration of cuckoo eggs is more variable when parasitizing hosts laying bluer‐greener eggs, even after controlling for the effect of host egg coloration (i.e. degree of egg matching). Globally, our results are consistent with the proposed hypothesis that host egg traits that are related to phenotypic quality of hosts, such as egg coloration, may have important implications for the coevolutionary interaction between hosts and brood parasites. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 154–168.     

Multiple responses to increasing spring temperatures in the breeding cycle of blue and great tits (Cyanistes caeruleus,Parus major)

Many bird species start laying their eggs earlier in response to increasing spring temperatures, but the causes of variation between and within species have not been fully explained. Moreover, synchronization of the nestling period with the food supply not only depends on first‐egg dates but also on additional reproductive parameters including laying interruptions, incubation time and nestling growth rate. We studied the breeding cycle of two sympatric and closely related species, the blue tit Cyanistes caeruleus and the great tit Parus major in a rich oak‐beech forest, and found that both advanced their mean first‐egg dates by 11–12 days over the last three decades. In addition, the time from first egg to fledging has shortened by 2–3 days, through a decrease in laying interruptions, incubation time (not statistically significant) and nestling development time. This decrease is correlated with a gradual increase of temperatures during laying, suggesting a major effect of the reduction in laying interruptions. In both species, the occurrence of second clutches has strongly decreased over time. As a consequence, the average time of fledging (all broods combined) has advanced by 15.4 and 18.6 days for blue and great tits, respectively, and variance in fledging dates has decreased by 70–75%. Indirect estimates of the food peak suggest that both species have maintained synchronization with the food supply. We found consistent selection for large clutch size, early laying and short nest time (laying to fledging), but no consistent changes in selection over time. Analyses of within‐individual variation show that most of the change can be explained by individual plasticity in laying date, fledging date and nest time. This study highlights the importance of studying all components of the reproductive cycle, including second clutches, in order to assess how natural populations respond to climate change.     

Genetic analysis of ryanodine receptor function in<Emphasis Type="Italic"> Caenorhabditis elegans</Emphasis> based on<Emphasis Type="Italic"> unc-68</Emphasis> revertants

The Caenorhabditis elegans ryanodine receptor is encoded by the unc-68 gene, and functions as a Ca²⁺-induced Ca²⁺ release channel during muscle contraction. To investigate the factors that suppress calcium release and identify molecules that interact with the ryanodine receptor, we isolated revertants from two unc-68 mutants. Three of the revertants obtained from the null allele unc-68(e540), which displayed normal motility, had intragenic mutations that resulted in failure to splice out intron 21. The other two, kh53 and kh55, had amino acid insertions in the third of the four RyR domains. The brood size and the egg laying rate remain abnormal in these revertants. This suggests the third RyR domain may be required for egg laying and embryogenesis, although we can not determine a molecular mechanism. Five ketamine sensitive revertants recovered from the missense mutant unc-68(kh30) showed altered responses to caffeine, ryanodine, levamisole and ouabain relative to those of the unc-68(kh30) animals. These may carry second-site suppressor mutations, which may define genes for proteins that regulate the Ca²⁺ concentration in body-wall muscle. One of these mutants, kh52 , shows lower motility and higher sensitivity to drugs, and this mutation was mapped to chromosome X. These observations provide a basis for the study of ryanodine receptor functions in embryogenesis and in calcium-mediated regulation of muscle contraction in C. elegans. This is the first study to show that the conserved RyR domain of the receptor acts in egg laying and embryogenesis.Communicated by C. P. Hollenberg     

Effects of Enterococcus faecalis on egg production,egg quality and caecal microbiota of hens during the late laying period

This study was conducted to determine the effects of diet supplementation of laying hens with Enterococcus faecalis (EF) on egg production, egg quality and caecal microbiota. A total of 360 Hy-Line Brown laying hens (72 weeks old) were divided into three groups with four replicates of 30 birds each. The laying hens were fed with the basal diet (Control), the basal diet + 3.75 · 10⁸ cfu EF/kg (Group I) or the basal diet + 7.5 · 10⁸ cfu EF/kg (Group II). The experiment lasted for 45 d. Eggs and caecal samples were collected at the end of the experiment. Results showed that dietary supplementation with EF did not affect the average daily egg weight, cracked egg rate, mortality and egg quality. However, EF supplementation caused a significantly increased laying rate and decreased feed/egg ratio (p < 0.05). The differences in caecal microbiota between Group II and the Control were significant. The relative abundance of Verrucomicrobia and Cyanobacteria at the phylum level, Rikenellaceae, Christensenellaceae and Veillonellaceae at the family level, and the Faecalibacterium, Christensenellaceae R-7 group and Eubacterium coprostanoligenes group at the genus level changed significantly in Group II compared with the Control (p < 0.05). In conclusion, the tested dietary supplementations with EF improved product performance and affected the caecal microbial community structure of laying hens during the late laying period.     

Geographical variation in reproductive traits and trade‐offs between size and number of eggs in the king ratsnake,Elaphe carinata

We collected gravid king ratsnakes (Elaphe carinata) from three geographically separated populations in Chenzhou (CZ), Lishui (LS) and Dinghai (DH) of China to study the geographical variation in female reproductive traits and trade‐offs between the size and number of eggs. Not all reproductive traits varied among the three populations. Of the traits examined, five (egg‐laying date, post‐oviposition body mass, clutch size, egg mass and egg width) differed among the three populations. The egg‐laying date, ranging from late June to early August, varied among populations in a geographically continuous trend, with females at the most northern latitude (DH) laying eggs latest, and females at the most southern latitude (CZ) laying eggs earliest. Such a trend was less evident or even absent in the other traits that differed among the three populations. CZ and DH females, although separated by a distance of approximately 1100 km as the crow flies, were similar to each other in most traits examined. LS females were distinguished from CZ and DH females by the fact that they laid a greater number of eggs, but these were smaller. The egg size–number trade‐off was evident in each of the three populations and, at a given level of relative fecundity, egg mass was significantly greater in the DH population than in the LS population. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 701–709.     

广聚萤叶甲成虫产卵行为及其产卵部位的选择性

【目的】明确广聚萤叶甲成虫产卵行为及其产卵部位的选择性。【方法】在室内条件下,对广聚萤叶甲成虫交配及产卵的系列行为、产卵场所选择、不同部位豚草植株叶片叶绿素b的含量进行了观察和测定:(1)将1对成虫放到养虫笼内的一株豚草上,观察交配时间,记录产卵数量、前后2粒卵之间的产卵时间间隔;(2)在均匀分为5部分(0～10、11～20、21～30、31～40和41～50 cm)的豚草植株上,随机放置10对成虫,观察雌虫对于产卵场所的选择。(3)将上述5个部位的豚草叶片通过丙酮匀浆法处理,用紫外分光光度计测定其在645和663 nm的吸光值,计算叶绿素b含量。【结果】广聚萤叶甲成虫完成一次成功交配平均需96.09 min。雌虫一般需45 min的时间来寻找其适应的产卵场所,在产卵过程中,成虫习惯将卵产于叶片背面,雌虫喜欢用口器来清理刚产下的卵粒。在一株50 cm高的豚草植株上,雌虫喜欢将卵产在植株中部21～30 cm和中上部31～40cm的叶片上(从下往上划分)。卵块数量和豚草不同部位叶片叶绿素b含量呈显著的正相关性。【结论】广聚萤叶甲成虫喜欢产卵在叶绿素b含量较高的叶片背面,可能以视觉识别叶片颜色来选择和定位产卵场所。     

Egg mass in an asynchronously hatching parrot: does variation offset constraints imposed by laying order?

Life history theory predicts phenotypic trade-offs between the number and quality of offspring produced. Intraspecific variation in egg mass is common in birds and increased egg size can have positive effects on offspring fitness. However, evidence of a trade-off with clutch size is limited. We analyzed variation in mass of 5,743 Green-rumped parrotlet (Forpus passerinus) eggs laid over 15 years to evaluate the potential for facultative adjustment of egg mass and factors governing variation. Heavier eggs had an increased probability of both hatching and fledging but egg mass did not affect postfledging recruitment. Offspring egg mass differed between populations and the potential for seasonal adjustment to egg mass may be related to environmental factors such as seed density. Egg mass was moderately heritable (h ²=0.42) which accounts for some of the individual variation detected, and these results are likely attributable to strong maternal effects. We found an effect of female age on egg mass, but no effects of previous reproductive experience. Finally, egg mass was strongly governed by position within the laying sequence, independent of clutch size, and such adjustment may facilitate brood reduction under some conditions in this highly asynchronous species.