Adult population density,fecundity and productivity in Drosophila melanogaster and Drosophila simulans

Summary Single-species cultures of D. melanogaster Oregon-R-C and D. simulans v were set up with 5, 50, 100, 200, 300 or 400 pairs of parents. These parents were discarded after 48 hours, and the numbers and wet weights of emerging progeny recorded twice daily. For each culture, the fitness components estimated were total number of progeny, total progeny biomass, average male and average female wet weights, mean developmental period, and sex ratio. D. melanogaster had higher progeny productivity and longer mean developmental period. For both species, as adult density increased, progeny number per culture increased to a maximum and then decreased, but the average number of progeny per female decreased rapidly from the lowest density. The cause of this decreased progeny number per female differed in the two species. For simulans, it was due to decreased fecundity per female, possibly a behavioural response to crowding. For melanogaster, the decreased progeny number per female was mainly due to reduced immature stage viability as a result of increased larval crowding. Reduction in fecundity per female was relatively small, as compared with simulans.     

Interactions within and between species at different densities of the bark beetleIps typographus and its predatorThanasimus formicarius

Interactions between the bark beetleIps typographus and one of its predators,Thanasimus formicarius, were investigated in caged spruce logs containing both species in eight different density combinations. Productively (offspring per female) ofIps was adversely affected by highIps gallery density as well as highThanasimus density.Thanasimus productivity was enhanced by highIps gallery density but negatively affected by highThanasimus density.Ips productivity andThanasimus developmental rate differed between tree individuals, probably owing to tree-related differences in phloem thickness. Relative predator-causedIps mortality was ca 20% higher at high gallery density (ca 300 egg galeries per m²) than at low gallery density (ca 100 egg galleries per m²), indicating that mortality was density dependent. This difference was due to the fact thatThanasimus larval density was positively related toIps gallery density. Mortality increased by ca 0.4% with each additionalThanasimus larva per m², independently of gallery density. Relative population levels ofIps andThanasimus were monitored with pheromone traps in two regions differing in theirIps typographus outbreak history. Absolute catch as well as theThanasimus/Ips catch ratio were ten times greater in the outbreak region than in the non-outbreak region. Coupled with the results in the caging experiment, this indicates thatT. formicarius responds numerically to changes inI. typographus numbers per unit bark area as well as toIps population changes at the regional level. The findings suggest that predation under bark may be a significant factor in supressingI. typographus outbreaks.     

Acceptability and suitability of Spodoptera exigua (Hübner) for Cotesia icipe Fernandez‐Triana & Fiaboe on amaranth

The beet armyworm Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae) is a polyphagous insect that is distributed worldwide and was recently reported as an important pest on African indigenous vegetables. Cotesia icipe Fernandez‐Triana & Fiaboe (Hymenoptera: Braconidae) is a recently described parasitoid, reported from various Afrotropical countries. This work investigated the performance of C. icipe on S. exigua infesting Amaranthus dubius Mart. ex Thell. under laboratory conditions. Cotesia icipe was aggressive on the host and successfully oviposited on S. exigua with 70% of parasitoid females ovipositing after 2 hr of exposure. Parasitoid densities significantly affected the parasitism rate and the nonreproductive larval mortality. Parasitism rate was 9.7 ± 0.8% and 59.5 ± 3.1% for a single and cohort of five females released, respectively, when offered 50 host larvae. The cohort female release resulted in significantly higher larval nonreproductive mortality than the single release. However, there was no significant difference between parasitoid release densities in regard to pupal nonreproductive mortality. The larval and pupal mortalities in the presence of C. icipe were significantly higher than the natural mortalities at both parasitoid release densities. The parasitoid sex ratio was female‐biased for the cohort females but balanced when a single female was released. The hind tibia and forewing lengths were not affected by the density of female parasitoids but there were variations according to sex. The implication of these findings on the potential use of C. icipe for biological control of S. exigua in amaranth production systems is discussed.     

Age,growth, and sex ratio among populations of least brook lamprey,Lampetra aepyptera,larvae: an argument for environmental sex determination

Synopsis Sex ratios of least brook lamprey,Lampetra aepyptera, larvae varied widely among 12 geographically-diverse streams of the eastern United States. The extremes were 29 and 71% male, and the proportion of males increased significantly with relative population density, which was estimated among the streams from the number of larvae collected per m2 of substrate. The skewed sex ratios were not likely due to differential mortality between the sexes or differential recruitment to the adult stock, since they were established at the time of gonadal differentiation (at ca. 2 years of age) and remained relatively constant over the subsequent 2–3 years of larval life. Furthermore, although females seemed to predominate in the oldest larval age class, thus appearing to metamorphose later than males, their numbers were small and were omitted from the overall sex ratio. Sex ratio did not vary significantly with water hardness, pH, annual thermal units, or latitude. The possible adaptive significance of density-dependent sex determination in lampreys, however, remains elusive. It has been proposed that growth-promoting conditions might yield female-biased sex ratios as a tactic for ensuring that relatively large individuals become females, thereby increasing their fecundity. As predicted, larval size at a given age was generally greater in low-density populations, but there was no relationship between sex ratio and larval size, and female larvae were not consistently larger than the males.     

Effects of larval density on a natural population of Culex restuans (Diptera: Culicidae): no evidence of compensatory mortality

1. This study investigated the effects of strong density dependence on larval growth, development, and survival of the mosquito Culex restuans (Theobald). It also tested the hypothesis that density reduction early in larval development could result in as many or more individuals surviving to adulthood (compensation or over‐compensation, respectively), or increased reproductive performance via rapid development and greater adult size. 2. In a field study of a natural population of C. restuans, the effects of a 75% lower density on percentage survivorship to adulthood, number of adults, development time, adult size, adult longevity, and size dependent fecundity were tested. 3. No evidence was found of compensation or over‐compensation in adult production, or of effects of lower density on percentage survivorship. Low density yielded significant increases in adult size, adult longevity, and size‐dependent fecundity, and a decrease in development time. 4. Estimated per‐capita population growth rate was significantly greater in the low‐density treatment than in the high‐density treatment. It is inferred that this difference was due to greater per‐capita resources, which increased female size and fecundity, and reduced development time. Greater per‐capita population growth could therefore result from early mortality of larvae, meaning that the hydra effect, which predicts greater equilibrium population with, as opposed to without, extrinsic mortality, may be possible for these mosquitoes.     

Interactions between ecological factors in the developmental environment modulate pupal and adult traits in a polyphagous fly

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缙云山细枝柃种群性比及空间分布

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Sex ratio bias in the dung beetle <Emphasis Type="Italic">Onthophagus taurus</Emphasis>: adaptive allocation or sex-specific offspring mortality?

Sex allocation theory predicts that females should adjust the sex of their offspring when the fitness returns of one sex are higher than the other. However, biased sex ratios may also arise if mortality differs between the sexes. Here, we examine whether offspring sex ratio bias in the dung beetle, Onthophagus taurus, represents adaptive sex allocation by females or is due to sex-specific mortality. First, we re-analyze an existing data set to show that females produce an excess of daughters when mating to smaller, less attractive males and near equal sex ratio with large, more attractive males. We show, that this results from females adjusting larval provisions after mating to males of variable attractiveness which in turn influences the likelihood that sons die during development. Second, we conduct a manipulative experiment varying the quantity and quality of larval provisions and show that the mortality of sons increased when larval provisions were reduced. Collectively, our work demonstrates that offspring mortality is contingent on the amount of resources provisioned by females and that sons have greater nutritional demands than daughters during development, leading to higher mortality. Our results therefore demonstrate the importance of considering sex-specific offspring mortality in studies of sex ratio evolution.     

Parthenogenesis is rare in the reproduction of a sexual field population of the isogamous brown alga Scytosiphon (Scytosiphonaceae,Ectocarpales)

Parthenogenetic development of unfused gametes is commonly observed in laboratory cultures among various brown algal taxa. There is, however, little information on the contribution of parthenogenesis to the reproduction of field populations. In this study, we investigated whether parthenogenesis is present in a sexual population of the isogamous brown alga Scytosiphon with a 1:1 sex ratio. In culture, both female and male gametes showed higher mortality and slower development compared to zygotes. More than 90% of surviving partheno‐germlings formed parthenosporophytes irrespective of the culture conditions tested. Therefore, if parthenogenesis occurs in the field, most unfused gametes are expected to form parthenosporophytes. Contrary to this expectation, parthenosporophytes were rare in the field population. We collected 126 sporophytic thalli and isolated and cultured a unilocular sporangium from each of them. We confirmed that cultures of 120 unilocular sporangia produced both female and male gametophytes by the observation of zygotes or amplification of PCR‐based sex markers indicating that these sporangia originated from zygotic sporophytes. Only females were detected in cultures from two sporangia and only males from four sporangia suggesting that these sporangia originated from parthenosporophytes. In the Scytosiphon population, although parthenogenesis is observable in culture, our results demonstrate that the contribution of parthenogenesis to reproduction is small (≤4.8%) compared to sexual reproduction. Unfused gametes may not survive to form mature parthenosporophytes in significant numbers in the field partly due to their higher mortality and slower development compared from zygotes.     

Observations on the reproductive and larval biology ofBlennius pavo (Pisces: Teleostei)

Social behaviour and spawning of adultBlennius pavo kept in the laboratory are described. Eggs are deposited in batches on the walls of artificial spawning places (PVC pipes). One male guards and tends the eggs of different females in one spawning place. Larval hatching occurs in groups according to oviposition. Minimum incubation temperature is around 14–15°C. Larval survival in 1-1 rearing jars is not related to larval total length but to density of larval stock. An experimental population of laboratory reared juvenile and adolescentB. pavo displays a male to female ratio of 1:1.4. Factors possibly influencing the sex ratio of this littoral fish are discussed in view of the situation in its natural environment.     

Effective size of density‐dependent two‐sex populations: the effect of mating systems

Density dependence in vital rates is a key feature affecting temporal fluctuations of natural populations. This has important implications for the rate of random genetic drift. Mating systems also greatly affect effective population sizes, but knowledge of how mating system and density regulation interact to affect random genetic drift is poor. Using theoretical models and simulations, we compare N_e in short‐lived, density‐dependent animal populations with different mating systems. We study the impact of a fluctuating, density‐dependent sex ratio and consider both a stable and a fluctuating environment. We find a negative relationship between annual N_e/N and adult population size N due to density dependence, suggesting that loss of genetic variation is reduced at small densities. The magnitude of this decrease was affected by mating system and life history. A male‐biased, density‐dependent sex ratio reduces the rate of genetic drift compared to an equal, density‐independent sex ratio, but a stochastic change towards male bias reduces the N_e/N ratio. Environmental stochasticity amplifies temporal fluctuations in population size and is thus vital to consider in estimation of effective population sizes over longer time periods. Our results on the reduced loss of genetic variation at small densities, particularly in polygamous populations, indicate that density regulation may facilitate adaptive evolution at small population sizes.     

Differences in environmental response between the sexes of the dioecious shrub Baccharis halimifolia (Compositae)

Summary Baccharis halimifolia (Compositae) is a dioecious shrub which grows on the upland fringe of tidal marshes along the Atlantic and Gulf Coasts of North America. We examined the responses of the two sexes to variation in nutrient and moisture availability plant density, and defoliation. By growing plants from seedlings to flowering adults under various combinations of soil type, fertilization rate and plant density, we were able to establish different rates of plant growth and mortality. Plants grown at high density and low nutrient and water supply grew the least, incurrent the most mortality and showed a male-biased sex ratio (73% male). At low density with abundant nutrients and water, plants grew more, survived well, flowered frequently, and were female-biased (75% female). Changes in sex ratio were probably the result of sex-related mortality rather than sexual lability of the seedlings. While changes in sex ratio occurred under experimental conditions in the green-house, no evidence for differences in habitat utilization between the sexes were found in the field and the sex ratio (59% female) did not vary across habitats. In the marsh habitats we sampled where water and nutrients were apparently available, there was no evidence for differential mortality between the sexes. When defoliated (75% of leaf tissue), both sexes showed similar reductions in reproductive effort (number of flower heads/shoot). Our results indicate that differences between the sexes of Baccharis in their response to environmental growing conditions is an important ecological factor associated with the separation of male and female function into separate individuals.     

Influence of gizzard shad on phytoplankton size and primary productivity in mesocosms and earthen ponds in the southeastern U.S.

Gizzard shad (Dorosoma cepedianum), a filter feeding omnivore, can consume phytoplankton, zooplankton and detritus and is a common prey fish in U.S. water bodies. Because of their feeding habits and abundance, shad have the potential to affect primary productivity (and hence water quality) directly through phytoplankton grazing and indirectly through zooplankton grazing and nutrient recycling. To test the ability of shad to influence primary productivity, we conducted a 16-day enclosure study (in 2.36-m³ mesocosms) and a 3-year whole-pond manipulation in 2–5 ha earthen ponds. In the mesocosm experiment, shad reduced zooplankton density and indirectly enhanced chlorophyll a concentration, primary productivity, and photosynthetic efficiency (assimilation number). While shad did not affect total phytoplankton density in the mesocosms, the density of large phytoplankton was directly reduced with shad. Results from the pond study were not consistent as predicted. There were few changes in the zooplankton and phytoplankton communities in ponds with versus ponds without gizzard shad. One apparent difference from systems in which previous work had been conducted was the presence of high densities of a potential competitor (i.e., larval bluegill) in our ponds. We suggest that the presence of these extremely high larval bluegill densities (20–350 larval bluegill m^–3; 3–700 times higher density than that of larval gizzard shad) led to the lack of differences between ponds with versus ponds without gizzard shad. That is, the influence of gizzard shad on zooplankton or phytoplankton was less than the influence of abundant bluegill larvae. Differences in systems across regions must be incorporated into our understanding of factors affecting trophic interactions in aquatic systems if we are to be able to manage these systems for both water quality and fisheries.     

Population dynamics ofTelenomus fariai (Hymenoptera: Scelionidae), a parasite of Chagas' disease vectors IX. Larval competition and population size regulation under laboratory conditions

A series of experiments were carried out with the endophagous egg parasite Telenomus fariai on its host Triatoma phyllosoma pallidipennis to determine the possible role of intraspecific competition by the parasite progeny in population regulation of the parasite. Eight parasite densities (1, 3, 5, 10, 20, 30, 40, and 50 individuals per vial) were used, and the design of sequentially sacrificed replicates applied. Survivorship curves for each density indicated smaller number of progeny per host at higher densities, and the shapes of the curves suggested a relatively early mortality process.Morris' linear regression technique for determining within-generation density-dependence was used, and the results showed that only larval mortality could be identified as density dependent. The same technique applied within the larval stage proved that only mortality of larvae in their second, third, and fourth day of development were responsible for population regulation. The applicability of the technique, as well as the relevance of the results for natural population, is discussed.     

Developmental mortality increases sex‐ratio bias of a size‐dimorphic bark beetle

1. Given sexual size dimorphism, differential mortality owing to body size can lead to sex‐biased mortality, proximately biasing sex ratios. This mechanism may apply to mountain pine beetles, Dendroctonus ponderosae Hopkins, which typically have female‐biased adult populations (2 : 1) with females larger than males. Smaller males could be more susceptible to stresses than larger females as developing beetles overwinter and populations experience high mortality. 2. Survival of naturally‐established mountain pine beetles during the juvenile stage and the resulting adult sex ratios and body sizes (volume) were studied. Three treatments were applied to vary survival in logs cut from trees containing broods of mountain pine beetles. Logs were removed from the forest either in early winter, or in spring after overwintering below snow or after overwintering above snow. Upon removal, logs were placed at room temperature to allow beetles to complete development under similar conditions. 3. Compared with beetles from logs removed in early winter, mortality was higher and the sex ratio was more female‐biased in overwintering logs. The bias increased with overwinter mortality. However, sex ratios were female‐biased even in early winter, so additional mechanisms, other than overwintering mortality, contributed to the sex‐ratio bias. Body volume varied little relative to sex‐biased mortality, suggesting other size‐independent causes of male‐biased mortality. 4. Overwintering mortality is considered a major determinant of mountain pine beetle population dynamics. The disproportionate survival of females, who initiate colonisation of live pine trees, may affect population dynamics in ways that have not been previously considered.     

Biology ofhyphantria cunea drury (lepidoptera : Arctiidae) in Japan. IX. population dynamics

Fluctuations in population density of Hyphantria cunea in Japan are characterized by a gradation-like pattern. Analysis of the life table data taken from two stations during eight successive generations showed that (1) mortality during egg and early larval stages was density-independent, (2) mortality during later larval stages was inversely density-dependent, and (3) mortality during prepupal and pupal stages was density-independent. Thus the overall mortality process from egg to adults eclosion was inversely density-dependent. The inverse density-dependence in mortality process during later larval instars was mainly attributed to the ‘escape’ (VOÛTE, 1946) of H. cunea populations from the predation pressure of polyphagous predators such as birds and Politses wasps. This inverse density-dependence was considered to be a cause of gradation-like fluctuation. Field collection of egg-masses showed that the mean number of eggs per egg-mass, which was believed to be a good representation of mean fecundity, varied from 425 to 1050 during 4 years. Density-dependent reduction in the mean number of eggs per egg-mass was demonstrated, and this reduction was a factor regulating the population density. Assuming fixed sex ratio and survival rate of adults, a preliminary population model was constructed. The number of eggs laid in the survey station could be predicted well by the model based on the number of eggs laid in the previous generation, in 9 out of 13 cases. An attempt to apply a model of the same type to mimic the fluctuation of abundance (peak number of larval colonies per tree) on road-side trees suggested another density-dependent process, that is, insecticide application by man. Discussion was also presented on the causes responsible for the turn of population trend from decreasing to increasing in the 1st generation of 1968.     

Distribution, density and productivity of great bustards Otis tarda in northwestern Spain: a regional approach

This paper describes at a regional scale the distribution pattern, density, productivity and sex ratio of great bustards in northwestern Spain and explores the role played by habitat type, terrain characteristics and human disturbance on variation in its demographic parameters. Data from 136 plots covering an area of 7314 km² were obtained in two censuses carried out in the spring and summer of 1998. The density of the great bustards was 1.39 individuals/km² in the pre-breeding period, decreasing by 22% in the post-breeding period. Density was significantly higher in central plots than in peripheral plots. Mean productivity was 0.24 chicks per female and showed a high variability among plots, being significantly lower in the densest plots. The overall sex ratio was 1.35 females per male during the pre-breeding period. Productivity related positively to areas holding small fields and a high interspersion of land uses. Density and productivity were negatively affected by river density and altitude, and seasonal density variation was positively correlated with human population density.     

Male‐killer dynamics in Danaus chrysippus (L.) (Lepidoptera: Nymphalidae) in East Africa

Within certain regions in East Africa, the butterfly Danaus chrysippus (L.) shows female‐biased population sex ratio, because of the production by some females of all‐female broods, as a result of infection by maternally inherited, male‐killing bacterium of the genus Spiroplasma. In this study, we describe a 3‐year field survey for the population dynamics of the male‐killing Spiroplasma in D. chrysippus in four independent localities, namely Uganda, Ghana, Sudan and Madagascar. The prevalence of the bacterium was found to show extensive variations at multiple scales among different sites, in various countries, seasons and years. A novel, selection‐based hypothesis was suggested to explain the high variability of male‐killer prevalence over space and time, based on the existence of an adaptive link between larval food‐plant density and the magnitude of resource reallocation fitness advantage for the male‐killer.     

Food limitation of a Delaware salt marsh population of the mummichog,Fundulus heteroclitus (L.)

Summary The relationship of the mummichog, Fundulus heteroclitus, to its food supply was examined using 8 m x 20 m enclosures under field conditions. Density effects were examined by stocking these enclosures with fish at densities ranging from 1/8 to four times the natural population density. Mean growth rate in each pen was estimated from measurements on individually marked fish. Fish enclosed at normal density grew at the same rate as fish from the natural population, indicating that caging had minimal effects on growth rate. When enclosed below natural density, mummichogs had a growth rate that was 2–3 times higher than that of the natural population. Fish enclosed at four times normal density had a negative mean growth rate, a higher mortality rate than fish enclosed at natural density, and no egg production.In further experiments, food was added to the same type of enclosure at fish densities ranging from one-half to four times natural density. Food additions had a positive effect on growth rate at all densities. Food additions also caused egg production of fish enclosed at the highest density to increase to a rate equivalent with the natural population. We conclude that the total food supply may regulate the maximum size of the Canary Creek mummichog population by affecting the individual growth rate, mortality rate, and fecundity.     

Biotic and abiotic effects on density,body size,sex ratio,and survival in immature stages of the European woodwasp,Sirex noctilio

Resource quality can have direct or indirect effects on female oviposition choice, offspring growth and survival, and ultimately on body size and sex ratio. We examined these patterns in Sirex noctilio Fabricus, the globally invasive European pine woodwasp, in South African Pinus patula plantations. We studied how tree position as well as natural variation in biotic and abiotic factors influenced sex‐specific density, larval size, tunnel length, male proportion, and survival across development. Twenty infested trees divided into top, middle, and bottom sections were sampled at three time points during larval development. We measured moisture content, bluestain fungal colonization, and co‐occurring insect density and counted, measured, and sexed all immature wasps. A subset of larval tunnels was measured to assess tunnel length and resource use efficiency (tunnel length as a function of immature wasp size). Wasp density increased from the bottoms to the tops of trees for both males and females. However, the largest individuals and the longest tunnels were found in bottom sections. Male bias was strong (~10:1) and likewise differed among sections, with the highest proportion in the middle and top sections. Sex ratios became more strongly male biased due to high female mortality, especially in top and middle sections. Biotic and abiotic factors such as colonization by Diplodia sapinea, weevil (Pissodes sp.) density, and wood moisture explained modest residual variation in our primary mixed effects models (0%–22%). These findings contribute to a more comprehensive understanding of sex‐specific resource quality for S. noctilio and of how variation in key biotic and abiotic factors can influence body size, sex ratio, and survival in this economically important woodwasp.