Impact of developmental variance on behavior of models for insect populations I. Models for populations with unrestricted growth

The concept of developmental variance is discussed with reference to its use in models for insect populations. When included in a model, developmental variance is typically used to describe the variation of developmental periods among individuals. However, its presence in a model can also have indirect impact on survival and fertility schedules. This impact can lead to significant changes in population growth rates and generation times. These relationships between developmental variance and population growth in models are quantified and discussed.     

High temperatures reveal cryptic genetic variation in a polymorphic female sperm storage organ

Variation in female reproductive morphology may play a decisive role in reproductive isolation by affecting the relative fertilization success of alternative male phenotypes. Yet, knowledge of how environmental variation may influence the development of the female reproductive tract and thus alter the arena of postcopulatory sexual selection is limited. Yellow dung fly females possess either three or four sperm storage compartments, a polymorphism with documented influence on sperm precedence. We performed a quantitative genetics study including 12 populations reared at three developmental temperatures complemented by extensive field data to show that warm developmental temperatures increase the frequency of females with four compartments, revealing striking hidden genetic variation for the polymorphism. Systematic genetic differentiation in growth rate and spermathecal number along latitude, and phenotypic covariance between the traits across temperature treatments suggest that the genetic architecture underlying the polymorphism is shaped by selection on metabolic rate. Our findings illustrate how temperature can modulate the preconditions for sexual selection by differentially exposing novel variation in reproductive morphology. This implies that environmental change may substantially alter the dynamics of sexual selection. We further discuss how temperature-dependent developmental plasticity may have contributed to observed rapid evolutionary transitions in spermathecal morphology.     

The influence of variation and of developmental constraints on the rate of multivariate phenotypic evolution

A model of multivariate phenotypic evolution is analysed under the assumption that all characters have the same variance or at least constant ratios of variance. The rate of evolution is examined as a function of the amount of phenotypic variance in a variety of adaptive landscapes (fitness functions). It is demonstrated that the effect of variation depends on the type of adaptive landscape. In “well behaved” adaptive landscapes the rate of evolution can theoretically increase without limits, depending on the amount of heritable phenotypic variation. However, in other adaptive landscapes there are upper limits to the rate of evolution which cannot be exceeded if phenotypic variation is developmentally unconstrained, i. e. if it is the same for all characters. Further it is shown that the maximal rate of evolution becomes small if the number of characters becomes large. Fitness functions of this type are called malignant. It is argued that malignant fitness functions are more adequate models for the evolution of typical organismic systems, because they are models of functionally interdependent characters. It is concluded that there are upper limits to the rate of phenotypic evolution if the variation of functionally interdependent characters is developmentally unconstrained. The possible role of developmental constraints in adaptive phenotypic evolution is discussed.     

Individual variability and mortality required for constant final yield in simulated plant populations

When plant monocultures are sown over a wide range of densities for a given period of time, the total biomass yield increases with density at low densities and then levels off at high densities, a phenomenon called constant final yield (CFY). There are several reported cases, however, where the total yield decreases at very high densities, but the reasons for such exceptions are not known. We used a spatially explicit, individual-based “field of neighborhood” simulation model to investigate the potential roles of spatial pattern, individual variation, and competitive stress tolerance for CFY. In the model, individual plants compete asymmetrically for light when their fields overlap, and this competition decreases growth and increases mortality. We varied (1) the initial size variation, (2) the spatial pattern, and (3) ability to survive intense competition and examined the effects on the density-biomass relationship. CFY was always observed when there was high variability among individuals, but not always when variability was low. This high size variation could be the result of high initial size variability or variation in the degree of local crowding. For very different reasons, very high and very low tolerance for competition resulted in decreasing total biomass at very high densities. Our results emphasize the importance of individual variation for population processes and suggest that we should look for exceptions to CFY in homogeneous, even-aged, regularly spaced populations such as plantations.     

High variation in developmental instability under non-normal developmental error: a Bayesian perspective

The developmental mechanisms behind developmental instability (DI) are only poorly understood. Nevertheless, fluctuating asymmetry (FA) is often used a surrogate for DI. Based on statistical arguments it is often assumed that individual levels of FA are only weakly associated with the underlying DI. Patterns in FA therefore need to be interpreted with caution, and should ideally be transformed into patterns in DI. In order to be able to achieve that, assumptions about the distribution of developmental errors must be made. Current models assume that errors during development are additive and independent such that they yield a normal distribution. The observation that the distribution of FA is often leptokurtic has been interpreted as evidence for between-individual variation in DI. This approach has led to unrealistically high estimates of between-individual variation in DI, and potentially incorrect interpretations of patterns in FA, especially at the individual level. Recently, it has been suggested that the high estimates of variation in DI may be biased upward because either developmental errors are log-normal or gamma distributed and/or low measurement resolution of FA. A proper estimation of the amount (and shape) of heterogeneity in DI is crucial for the interpretation of patterns in FA and their transformation into patterns in DI. Yet, incorrect model assumptions may render misleading inferences. We therefore develop a statistical model to evaluate the sensitivity of results under the normal error model against the two alternative distributions as well as to investigate the importance of low measurement resolution. An analysis of simulated and empirical data sets indicated that bias due to misspecification of the developmental error distribution can be substantial, yet, did not appear to reduce estimates of variation in DI in empirical data sets to a large extent. Effects of low measurement resolution were neglectable. The importance of these results are discussed in the context of the interpretation of patterns in FA.     

Estimating developmental and mortality rates and stage recruitment from insect stage-frequency data

A model for the analysis of insect stage-frequency data is developed which includes stage-specific variable developmental periods and stage-specific daily survival rates. The model can predict the development of an insect population through its developmental stages and consequently may form the basis for a simulation model of the population.     

The demography of the annual halophyte Spergularia marina on a baltic seashore meadow

The survivorship and reproduction of the annual halophyte Spergularia marina was studied during four years on three different sites on a Baltic seashore meadow. The survivorship showed variation both between years and between sites. The mortality rate was not density-dependent in spite of very high densities in some years. At two of the sites the seed output was related to density, but individuals differed in their reproductive capacity. Limitation of the flower production was the main reason for this difference. At the third site the length of the flowering season determined the reproductive output.     

An evolutionary correlate of genome size change in plethodontid salamanders

Variation in the amount of nuclear DNA, the C-value, does not correlate with differences in morphological complexity. There are two classes of explanations for this observation, which is known as the ''C-value paradox''. The quantity of DNA may serve a ''nucleotypic'' function that is positively selected. Alternatively, large genomes may consist of junk DNA, which increases until it negatively affects fitness. Attempts to resolve the C-value paradox focus on the link between genome size and fitness. This link is usually sought in life history traits, particularly developmental rates. I examined the relationship among two life history traits, egg size and embryonic developmental time and genome size, in 15 species of plethodontid salamanders. Surprisingly, there is no correlation between egg size and developmental time, a relationship included in models of life history evolution. However, genome size is positively correlated with embryonic developmental time, a result that is robust with respect to many sources of variation in the data. Without information on the targets of natural selection it is not possible with these data to distinguish between nucleotypic and junk DNA explanations for the C-value paradox.     

Effect of aggregation on the reproductive rate of populations

The effect of spatial distribution on the reproductive rate of a population was examined by applying an approximate equation to several data based on the entomological literature. This study suggests that aggregation generally reduces the equilibrium level of the population density if the population density is not very high.     

Reproductive allocation patterns in different density populations of spring wheat

The effects of increased intraspecific competition on size hierarchies （size inequality） and reproductive allocation were investigated in populations of the annual plant, spring wheat （Triticurn aestivurn）. A series of densities （100, 300, 1 000, 3 000 and 10 000 plants/m^2） along a gradient of competition intensity were designed in this experiment. The results showed that average shoot biomass decreased with increased density. Reproductive allocation was negatively correlated to Gini coefficient （R^2 = 0.927）, which suggested that reproductive allocation is inclined to decrease as size inequality increases. These results suggest that both vegetative and reproductive structures were significantly affected by intensive competition. However, results also indicated that there were different relationships between plant size and reproductive allocation pattern in different densities. In the lowest density population, lacking competition （100 plants/m^2）, individual reproductive allocation was size independent but, in high density populations （300, 1 000, 3 000 and 10 000 plants/m^2）, where competition occurred, individual reproductive allocation was size dependent： the small proportion of larger individuals were winners in competition and got higher reproductive allocation （lower marginal reproductive allocation; MRA）, and the larger proportion of smaller individuals were suppressed and got lower reproductive allocation （higher MRA）. In conclusion, our results support the prediction that elevated intraspecific competition would result in higher levels of size inequality and decreased reproductive allocation （with a negative relationship between them）. However, deeper analysis indicated that these frequency- and size-dependent reproductive strategies were not evolutionarily stable strategies.     

Variation due to maternal age in the onion root maggot,Hylemya antiqua (Meigen)

A variety of characteristics were studied in populations of the onion root maggot, Hylemya antiqua, to determine the extent to which the age of the female producing a batch of eggs might affect heterogeneity in her offspring. Early-born offspring were found to be reproductively most successful. They had the longest mean expectation of life, a relatively low mortality rate into mid-life, the highest average fecundity, the highest mean rate of egg production, and they produced the highest percentage of female offspring. On the other hand, mid-born offspring were hardiest. They also had a long mean expectation of life, comparable to that for early-born offspring but, in addition, they had the lowest sustained mortality rate and the greatest ability to survive food stress. Their robustness may have been acquired at the expense of certain reproductive capabilities however, in that their average fecundity was lower than their early-born siblings', and their rate of egg production was more variable. Late-born offspring were the most mortality-prone and the least fecund, but they had the shortest mean generation time. These differences must be evaluated in terms of their effects on the rate of natural increase and on population growth. Maternal age was found to influence pupal size, although the provenance groups differed significantly from each other and, in fact, showed opposite trends within their respective cohort groups. Size itself, and the ability of a species to alter the size at which it pupates, may be characteristics which have evolved to fit a specific set of environmental conditions.     

不同生境条件下羊草种群无性系和有性生殖特征的比较研究

比较研究了东北草原不同生境条件下4个羊草种群无性系和有性生殖数量特征的差异,结果表明,由于各种群所处立地的资源差异(特别是土壤水分),导致各种群在无性系(包括营养枝和根茎芽密度)和有性生殖(生殖枝密度、生殖枝分化率、生殖贡献)等数量特征方面差异显著,尤其在低平原种群与沙丘种群之间最为显著.与沙丘种群相比,低平原种群营养枝平均密度高50%;根茎芽平均密度高30%以上;生殖枝密度平均值高200%以上.     

Selection on floral characters in natural Spanish populations of Silene latifolia

In insect-pollinated plants, floral characters are expected to play an important role in paternal and maternal reproductive success. Bateman's principle states that male reproductive success increases with more mating opportunities, while female reproductive success is limited by the amount of resources available to produce progeny, thus there should be greater selection on male floral characters than on female. In the case of the dioecious plant Silene latifolia, floral characters are likely to be influenced by its association within its native European range with moths of the genus Hadena, which serve as both pollinators and seed predators. The present study addresses relationships between male and female reproductive success, spatial location and floral characters (corolla, calyx and claw) over a 2-year period in two Spanish populations of S. latifolia in the presence of Hadena moths. A maximum likelihood paternity analysis using genetic variation at six allozyme markers showed heterogeneity in male reproductive success. There was much less variation in female reproductive success. When this analysis was extended to include interplant distance as a causal factor underlying variation in male success, we found that successful pollination tended to be limited to nearby females, in accordance with exponential decay of pollen dispersal with increasing distance. When the paternity analysis included floral characters as a causal factor underlying variation in male success, our data showed little evidence for directional selection, but there was stabilizing selection in one of the two years for calyx diameter. Selection on female characters varied widely between sites and years, in most of the site/year combinations there was little selection on female floral characters; however, in one site/year there was evidence for selection on all three floral characters. We conclude that pollinators visit flowers that are close together, and that while floral characters are important for the attraction of pollinators, larger flowers do not necessarily attract more pollinators at all sites and that variation among sites and years makes difficult any conclusions about the long-term importance of the predictions suggested by Bateman's principle.     

Demographics in an alpine reindeer herd: effects of density and winter weather

We examined how population density, winter weather, snow conditions, and 2 large-scale climatic indices (North Atlantic Oscillation, NAO, and Arctic Oscillation, AO) influenced demography (reproduction and mortality) in an alpine herd of semi-domesticated reindeer Rangifer tarandus between 1959 and 2000 in Finnish Lapland. The herd lived on heavily grazed lichen pastures, with winter densities between 0.8 and 3.9 individuals km⁻². Icing conditions occurred every 7th yr, on an average, and decreased reproductive rate (calves/females) by 49%. In general linear models icing remarkably increased the fit of snow models to reproductive rate. Incorporation of an interaction term between icing and the snow depth index provided better fit than a model without interaction. Delayed snowmelt decreased reproductive rate. For the day of snowmelt, however, the model without interaction was better than the interaction model. These 3 models provided the best fit to the data and accounted for 51–54% of the variation in reproductive rate. Winter mortality was related to density and large-scale climatic indices, but not to local winter weather except a slight increase in mortality during an icing winter. The best model for winter mortality, including reindeer density and NAO, accounted for 26% of variation in mortality. Three factors may be involved explaining weak density dependence or the lack of such dependence; climate change scenarios that predict higher winter temperature, more frequent thawing-freezing periods, and deeper snow would be expected to decrease reproductive rate and increase winter mortality of reindeer and thus to reduce profitability of reindeer husbandry. In contrast, early springs would be advantageous for reindeer in the short term.     

Population density, phenotype and reproductive output in the grasshopper Chorthippus brunneus

ABSTRACT. 1. The effects of population density on the reproductive output of individual female grasshoppers ( Chorthippus brunneus Thunberg) and the relationships between reproductive output and component elements of each female's phenotype, were investigated.
2. Reproductive output was primarily determined by the rate of egg-pod production. Increased density led to significant reductions in the rate of egg-pod production and reproductive output.
3. Exoskeleton size (reflecting nymphal experience) was positively correlated with clutch size at both high and low densities, but condition (reflecting adult experience) showed no such correlation. At high density, exoskeleton size was more strongly correlated with the rate of egg-pod production and reproductive output than condition. It is concluded that the nymphal aspects of size are more important than the adult aspects.
4. At low density, females with small exoskeletons compensated for their smaller clutches by producing egg-pods at a faster rate. Thus, there was no overall relationship between reproductive output and any of the phenotypic characteristics.
5. The effects of competition were weakly asymmetric in high density populations. The weakness of the asymmetry suggests scramble-like interactions for resources. It is concluded that not only are the effects of competition influenced by individual differences, but also that competition may reveal differences that would not otherwise be apparent.     

The effect of developmental variation on hybridization and rarity in Stipoid grasses

Developmental variation in some Achnatherum species was evaluated for two kinds of groups, (1) species pairs that do or do not hybridize and (2) rare and common species. Variation was assessed in two different ways, one that captures developmental events expressed in an individual and one reflecting developmental events that are part of the information systems of a species. The former captures the effect of the environment on development; the latter expresses developmental variation without the information controlling ontogenetic events being filtered through the environment. Development variation is lower for species pair that hybridizes when the effect of development in an individual is expressed. When that variation is of the species information system, the non-hybridizing species pair shows a lower level of developmental variation, likely the effect of greater similarity between those species. It is lower for rare species when variation in development is that of the information system of a species. The lower level of developmental variation seen in species pairs that hybridize likely reflects the necessity of compatible developmental programs in order for a hybrid to appear. Lower variation in development in rare species is expected. Here, though, the lower variation is a property of the species and not of the environment.     

Effect of diazinon on life stages and resting egg hatchability of rotifer Brachionus plicatilis

The effects of organophosphate pesticide, diazinon, on life history parameters and hatchability of resting eggs of rotifer Brachionus plicalitis were assessed. Newly hatched (<1 h-old) neonates were individually cultured in six varying concentrations (0/control, 0.1, 1.0, 2.5, 5.0 and 10.0 mg/l) of diazinon. The life history parameters such as time (h) the rotifers bear first egg and release first neonate, reproductive period, net reproductive rate, mixis, intrinsic rate of population increase, and life span were evaluated. Results showed that among the life history parameters, the time the rotifers took to release neonates is the most sensitive, giving the lowest EC₅₀ value of 1.24 mg/l. The fecundity of maternal females, amictic and mictic daughters was also investigated. Rotifers exposed to 10.0 mg/l produced significantly fewer amictic daughters, and at this concentration, rotifers did not produce any mictic daughter. At 5.0 mg/l, the number of male offspring was significantly lower than the control. Furthermore, the hatchability of resting eggs produced by the rotifers was evaluated when exposed to diazinon: from birth until they produced resting eggs (early development); during late developmental stage of resting eggs (before diapause); and during diapausing stage. The hatchability of the resting eggs was not affected when exposure was timed at late developmental and diapausing stages. Overall results showed that even though amictic females reproduced normally in the presence of low-concentration of diazinon, sexual reproduction is severely affected, especially the hatchability of resting eggs when the exposure was timed on its early developmental stages. Guest editors: S. S. S. Sarma, R. D. Gulati, R. L. Wallace, S. Nandini, H. J. Dumont & R. Rico-Martínez Advances in Rotifer Research     

Life on the edge: the demography of short-season populations of deer mice

We documented populations of deer mice in the Kananaskis Valley, southwestern Alberta, Canada from 1979 to 1997 to determine whether these short-season populations were more, or less, variable than populations in more temperate environments. We then examined patterns of reproduction, age-specific survival, and immigration to explain variation in summer population growth. Population densities showed no multi-annual periodicity and were generally low. At maximum, numbers doubled over the breeding season, but declined over the breeding season in 4 of 16 yr. Variability in population density was low, and similar to that of Peromyscus populations in more temperate environments. No demographic parameters were related to spring population densities, and immigration rates were low when conditions for survival of nestlings and adults were favorable. Variation in summer population growth was attributed primarily to variation in nestling survival among years.     

Factors influencing development and survival ofCulex pipiens pallens larvae (Diptera: Culicidae) in polluted urban creeks

Factors influencing development and survival of Culex pipiens pallens immatures in polluted urban creeks were studied in Saga, Japan. Addition of food shortened developmental durations and increased pupal size in floating cages at only the least polluted site out of 4 study sites. There was no evidence for developmental delay due to overcrowding in natural populations. Survival was not increased by artificial feeding at any site. Using the developmental parameters obtained for caged cohorts, survival in natural populations was estimated from differences between the actual number and the expectation if there were no mortality. Pupation rates ranged from 1> to >10% by site and season. Mortality due to predation, evaluated from the difference in pupation rates between caged and natural populations, exceeded 30% in 4 out of 6 cases (twice at 2 sites and once at the other 2 sites). Lethal factors in the creek water caused the predominant mortality in 2 other cases. Regulation of C. pipiens pallens population breeding in eutrophic, open creeks was discussed in the context of larval dispersal through behavioral interference.     

Reproductive assurance and the evolutionary ecology of self-pollination in Clarkia xantiana (Onagraceae)

The reproductive assurance hypothesis posits that selection favors self-pollination in flowering plants where mates and/or pollinators are scarce. A corollary is that self-pollinating populations are expected to be superior colonizers of mate- and pollinator-scarce environments. The California annual Clarkia xantiana includes outcrossing populations (ssp. xantiana) and autogamously self-pollinating populations (ssp. parviflora). Outcrossing is ancestral, and the subspecies have parapatric distributions with a narrow contact zone. We tested aspects of the reproductive assurance hypothesis by examining geographic and subspecies variation in the densities of mates and pollinators (native bees) and the density dependence of pollinator visitation and pollen receipt. Plant and flower densities, pollinator density, and pollinator visitation rates were lowest in the region of exclusively self-pollinating populations. Pollinator assemblages there lacked Clarkia-associated pollinator taxa that were common elsewhere. Self-pollinating populations in the contact zone generally had densities and visitation rates intermediate between allopatric self-pollinating populations and outcrossing populations. Visitation rate and pollen receipt increased significantly with plant density. These findings suggest that selection for reproductive assurance influenced the origin of self-pollination and/or that reproductive assurance influenced the geographic distribution of self-pollination. Geographic variation in pollinator assemblages may have generated variation in the value of reproductive assurance.