Life tables for worker honeybees

Life tables for worker honeybees covering all life span, and those for adults, were prepared for three seasonal cohorts, June bees, July bees and wintering bees. Survivorship curves for June and July bees show a convex type being exceptional for insects, with relatively high mortality at egg and feeding larval stages and at later adult stage after most bees became potential foragers. Adult longevity greatly lengthens in Winteriing bees and survivorship curve drops approximately with the same rate. A remarkable similarity of survivorship curves for men and honeybees was demonstrated, apparently due to highly developed social care in both. Some comments were given on mortality factors. The importance of life tables for population researches was shown by applying our result to the population growth curve made byBodenheimer , based upon the data byNolan . At the asymptote of the uncorrected curve, the ratio of total population estimated by uncorrected curve to that by corrected curve reaches about 3∶2.     

哺乳动物存活曲线类型的分析方法探讨

动物的存活曲线通常可分为3种类型：凸型（Ⅰ型或A型）、直线型（Ⅱ型或B型）和凹型（Ⅲ型或C型）。存活率曲线与存活曲线在概念上有一定的区别。存活率要经过对数转换（或者直接采用对数标尺）后才能得到存活曲线类型,从动物的存活率曲线上直接判读存活曲线类型可能会导致错误的结论。鉴于存活曲线分析研究中出现的问题,建议在进行物种存活曲线类型的分析时,务必先进行对数转换。     

山西七里峪茶条槭的种群结构与空间分布格局

从生命表特征、径级结构、存活曲线及不同海拔种群和不同龄级个体空间分布格局等方面研究了山西七里峪茶条槭的种群结构和空间分布规律.结果显示:(1)七里峪茶条槭种群径级结构呈现幼苗比例大,幼树、小树比例次之的规律,为增长型种群;(2)不同海拔种群中,1 300～1 600m为增长型种群,随着海拔的升高,种群结构逐渐趋于稳定成熟;(3)茶条槭种群的存活曲线接近于Deevey-Ⅲ,除1 400m和1 700m种群外,其它海拔种群存活曲线为Deevey-Ⅱ型;茶条槭种群不同径级的个体呈现出集群分布或随机分布,其空间分布格局随龄级增加而发生改变;(4)不同海拔高度种群的分布格局表现出从集群分布趋向随机分布的规律.研究表明,七里峪茶条槭种群的生活史特征与空间分布具有较为明显的特性,其幼苗储备丰富、自身的生物学特性及种群依存的微环境因素是导致这种现象的主要原因.     

Size-dependent survivorship in the web-building spiderAgelena limbata

Summary Stage-specific mortality rates and mortality factors for the web-building spiderAgelena limbata, which is suggested to be food-limited, were studied, and the relationship between body size of spiders and survivorship for instar 3 to adults was examined. The mortality rate of the egg sac stage including eggs, deutova (prenymphal stage), and overwintering instar 1 nymphs was low. The low mortality of this stage was partly due to maternal care that reduced the mortality caused by predation and/or abiotic factors. From emergence of instar 1 nymphs from egg sacs to reproduction, the stagespecific mortality rates were almost constant, 32–47%, and the time-specific mortality rates were also constant. These results suggest a Deevey (1947) type II survivorship curve inA. limbata, in contrast to other reports on the wandering or burrowing spiders which suggested type III curves. Important mortality factors for nymphs and adults were parasitism by an ichneumonid wasp and predation by spiders. There were great variations in body size (carapace width) ofA. limbata in the field. Smaller individuals survived at a lower rate to the next stage than larger individuals. This tendency was clearer for the population living under poorer prey availability.A. limbata was unlikely to starve to death in the field because every stage ofA. limbata could survive starvation for a long time in the laboratory, 22–65 days on average. I suggest that the size-dependent survivorship of this spider is associated with vulnerability of smaller individuals to parasitism and predation.     

Growth and survival rate of the larvae ofHynobius nebulosus Tokyoensis Tago (Amphibia,Hynobiidae)

Growth and population density of the larvae, Hynobius nebulosus tokyoensisTago , were estimated in a small pond within the study site settled in Habu village of Hinodemachi, a suburb of Tokyo City, during the period from 1975 to 1980. The mortality factors which influenced the survival rate of larvae were also evaluated from the ecological point of view. Laboratory experiments on the growth of larvae and predation by newts were conducted in pararell with the field survey. The results showed that growth rate of larvae under the natural condition was very slow, as compared with that under the laboratory condition with sufficient food supply, and mean body size at metamorphosis was negatively correlated with the density at that time. This suggested that food resources were in short supply in the pond, and there occurred a severe intraspecific competition for food among larvae. The mortality rate of larvae was so high, 80–99% in each year, and the density of larvae survived until metamorphosis varied so greatly from year to year that the larval stage was the most important stage throughout the life cycle to the maintenance of a population for this salamander. The most important factors which contributed to this high mortality were the predation by the newt, Triturus pyrrhogaster pyrrhogasterBoie , and cannibalism. From the laboratory experiment, it was found that predators could attack only small larvae successfully, and successful attack rate decreased sharply as larvae grew larger. This relationship resulted in the characteristic L-shaped pattern of survivorship curve of larvae; that is, heavy mortality just after hatching period.     

湖北神农架地区巴山冷杉（Abies fargesii）种群结构特征与空间分布格局

巴山冷杉群落是湖北神农架地区主要的群落类型之一,从生命表特征、年龄结构、存活曲线以及种群和不同龄级个体空间分布格局等方面研究了湖北神农架地区巴山冷杉的种群结构特征与空间分布规律.研究结果表明:巴山冷杉的种群年龄结构图呈现的是幼苗比例最大,中龄树、幼树和成龄树比例次之的规律;巴山冷杉幼苗储备丰富,巴山冷杉自身的生物学特性及环境因素是形成这种现象的主要原因;湖北神农架地区巴山冷杉种群的存活曲线接近于Deevey C型,I径级巴山冷杉死亡率最高,II径级开始巴山冷杉幼树死亡率大幅度下降,直至老龄期死亡率逐渐升高;巴山冷杉种群不同年龄级的个体呈现出集群分布或随机分布,处于不同发育阶段的个体群,其空间分布格局是随时间变化的.     

Darwinian fitness and the intensity of natural selection: studies in sensitivity analysis

Darwinian fitness, the capacity of a variant type to establish itself in competition with the resident population, is determined by evolutionary entropy, a measure of the uncertainty in age of the mother of a randomly chosen newborn. This article shows that the intensity of natural selection, as measured by the sensitivity of entropy with respect to changes in the age-specific fecundity and mortality variables, is a convex function of age, decreasing at early and increasing at later ages. We exploit this result to provide quantitative evolutionary explanations of the large variation in survivorship curves observed in natural populations. Previous studies to explain variation in survivorship curves have been based on the proposition that Darwinian fitness is determined by the Malthusian parameter. Hence the intensity of natural selection will be determined by the sensitivity of the Malthusian parameter with respect to changes in the age-specific fecundity and mortality variables. This measure of the selection gradient is known to be a decreasing function of age, with implications which are inconsistent with empirical observations of survivorship curves in human and animal populations. The analysis described in this paper point to the mitigated import of sensitivity studies based on the Malthusian parameter. Our analysis provides theoretical and empirical support for the ecological and evolutionary significance of sensitivity analysis based on entropy, which is the appropriate measure of Darwinian fitness.     

Contest and scramble competitions inCallosobruchus maculatus (Coleoptera: Bruchidae)

Larval competition curves and resource sharing patterns of 5 strains of Callosobruchus maculatus (iQ, yQ, aaQ, wQ, and tQ) were examined. Offspring emergences as a function of the initial larval density were recorded to construct competition curves. Elytron length of emerged adults was used as the indicator of resource sharing patterns among competing larvae inside a bean. In the large beans, strain iQ showed a saturated competition curve and tQ strain showed a humped curve. Competition curves of the other 3 strains (yQ, aaQ, and wQ) were between those two extremes. In the small beans, strains iQ and tQ also showed a saturated and a humped competition curves, respectively, whereas the competition curves of the 3 intermediate scramble strains could not be distinguished from that of the iQ strain. Thus, the classification based on competition curves was sensitive to the resource condition (bean size). In both the large and the small beans, the elytron lengths of iQ strain remained constant irrespective of initial larval density. On the contrary, the elytron lengths of the 4 other strains decreased monotonically with higher initial larval density. Thus, the judgment based on the resource sharing pattern was shown to be robust. Only iQ strain should be designated as a contest type, and the remaining strains as scaramble types. Contest and scramble types in C. maculatus were also compared with those observed in C. analis and C. phaseoli using competition curves, resource sharing patterns, and other physiological characters.     

The age-structured lottery model

The lottery model of competition between species in a variable environmental has been influential in understanding how coexistence may result from interactions between fluctuating environmental and competitive factors. Of most importance, it has led to the concept of the storage effect as a mechanism of species coexistence. Interactions between environment and competition in the lottery model stem from the life-history assumption that environmental variation and competition affect recruitment to the adult population, but not adult survival. The strong role of life-history attributes in this coexistence mechanism implies that its robustness should be checked for a variety of life-history scenarios. Here, age structure is added to the adult population, and the results are compared with the original lottery model. This investigation uses recently developed shape characteristics for mortality and fecundity schedules to quantify the effects of age structure on the long-term low-density growth rate of a species in competition with its competitor when applying the standard invasibility coexistence criterion. Coexistence conditions are found to be affected to a small degree by the presence of age structure in the adult population: Type III mortality broadens coexistence conditions, and type I mortality makes them narrower. The rates of recovery from low density for coexisting species, and the rates of competitive exclusion in other cases, are modified to a greater degree by age structure. The absolute rates of recovery or decline of a species from low density are increased by type I mortality or early peak reproduction, but reduced by type III mortality or late peak reproduction. Analytical approximations show how the most important effects can be considered as simple modifications of the long-term low-density growth rates for the original lottery model.     

Pheromone trapping models for pest control: Effects of mating patterns and immigration

Several models are presented which examine pest population behaviour with the release of female sex pheromones for the attraction and annihilation of males. These models include male polygamy and female monogamy, various mating frequencies, delayed mating of females, immigration of one or all individual types, and differential survivorship of males and females. In all the models there are two steady states, a stable s.s. at the origin and an unstable s.s. in the positive domain for a given value of pheromone release rate. In all the models, control relies on the reduced ability of males to fertilize virgin females following trapping and male annihilation. As such, control is very sensitive to mating frequency, being very difficult when males mate frequently. Control is also very difficult with the immigration of even a moderate number of fertilized females. Control is much easier when mating is delayed, especially if survivorship is low, or with density dependent population regulation.     

The 'ghost of settlement past' determines mortality and fecundity in the barnacle, Semibalanus balanoides

Intraspecific competition is common in many organisms. For many sessile marine invertebrates high settlement densities can lead to competition with conspecifics that can affect the probability of mortality, morphology, and reproductive output. This work aims to determine the affect of recruit density on shell morphology and fecundity and the temporal nature of intraspecific competition for the acorn barnacle, Semibalanus balanoides . Two tile arrays were used; experiment 1 was deployed at the start of the settlement season (43,264 recruits, 4.1% survivorship to adult), experiment 2 was deployed in the middle of the season (5,454 recruits, 42.8% survivorship). Experiment 2 displayed density dependent mortality between recruits and juveniles and adults, whereas no relationship was found for exp. 1. Experiment 2 had higher egg production per individual, but also higher egg production per area of tile compared to exp. 1. A negative relationship was found between recruit density and egg production, however during the summer period, this relationship switched to a positive relationship between juvenile density and egg production, due to high mortality on tiles with high recruit density. Cumulative population density (CPD) was calculated as the cumulative mean density of barnacles per cm^-2 of tile and was used as an index of intraspecific competition. There was a linear negative relationship between mass of eggs per individual and CPD. However, CPD over the summer period more fully explained egg production per individual. A time window was identified when cumulative intraspecific competitive effects were shown to have a strong negative effect on egg production and an increase in mortality, causing a flip from density independent fecundity to density dependent fecundity. These results suggest that recruit density is an important driver of subsequent population processes.     

Behavioral signature of intraspecific competition and density dependence in colony‐breeding marine predators

In populations of colony‐breeding marine animals, foraging around colonies can lead to intraspecific competition. This competition affects individual foraging behavior and can cause density‐dependent population growth. Where behavioral data are available, it may be possible to infer the mechanism of intraspecific competition. If these mechanics are understood, they can be used to predict the population‐level functional response resulting from the competition. Using satellite relocation and dive data, we studied the use of space and foraging behavior of juvenile and adult gray seals (Halichoerus grypus) from a large (over 200,000) and growing population breeding at Sable Island, Nova Scotia (44.0 ^oN 60.0 ^oW). These data were first analyzed using a behaviorally switching state‐space model to infer foraging areas followed by randomization analysis of foraging region overlap of competing age classes. Patterns of habitat use and behavioral time budgets indicate that young‐of‐year juveniles (YOY) were likely displaced from foraging areas near (<10 km) the breeding colony by adult females. This displacement was most pronounced in the summer. Additionally, our data suggest that YOY are less capable divers than adults and this limits the habitat available to them. However, other segregating mechanisms cannot be ruled out, and we discuss several alternate hypotheses. Mark–resight data indicate juveniles born between 1998 and 2002 have much reduced survivorship compared with cohorts born in the late 1980s, while adult survivorship has remained steady. Combined with behavioral observations, our data suggest YOY are losing an intraspecific competition between adults and juveniles, resulting in the currently observed decelerating logistic population growth. Competition theory predicts that intraspecific competition resulting in a clear losing competitor should cause compensatory population regulation. This functional response produces a smooth logistic growth curve as carrying capacity is approached, and is consistent with census data collected from this population over the past 50 years. The competitive mechanism causing compensatory regulation likely stems from the capital‐breeding life‐history strategy employed by gray seals. This strategy decouples reproductive success from resources available around breeding colonies and prevents females from competing with each other while young are dependent.     

Demographic mechanisms in the coexistence of two closely related perennials in a fluctuating environment

The demographic variability and life history differentiation of two closely related shrubs (Atriplex canescens and A. acanthocarpa) were investigated in the Chihuahuan Desert, and the results were interpreted in the context of theories of coexistence in fluctuating environments. Demographic information was recorded during three annual intervals and analyzed employing matrix projection models. A. canescens had lower λ (finite rate of population increase), higher longevity and generation time and slower convergence to a stable population structure than A. acanthocarpa. In favorable years for recruitment (those when, for both species, λ > 1), the λ of A. acanthocarpa was higher than that of A. canescens; in unfavorable years (when λ < 1), the reverse was true. Regardless of conditions (year), A. acanthocarpa had a type 2 survivorship curve (constant rate of mortality with age), while A. canescens had a type 3 survivorship curve (declining mortality with age). Elasticity analyses highlighted the larger influence that fecundity and growth would have in modifying the λ of A. acanthocarpa in comparison to that of A. canescens. In contrast, survival would have a larger influence on the λ of A. canescens. Atriplex acanthocarpa behaved as an opportunistic species that benefitted from sporadic favorable conditions and declined rapidly when conditions deteriorated. In contrast, A. canescens behaved as a tolerant species able to withstand years when conditions were poor, but which could not gain any advantage over A. acanthocarpa when conditions improved. By each having a relative advantage over the other on opposite ends of the contrasting climatic conditions experienced in the Chihuahuan Desert, they are able to coexist. Their contrasting life histories agreed with the theoretical predictions for the operation of the two mechanisms of species coexistence in fluctuating environments: the storage effect and the relative non-linearity of competition. Based on these results, we conclude by speculating on the nature of succession in arid communities.     

Supply-side control over the survivorship of Chthamalus spp. recruits in northern Baja California

The constant supply of larvae to coastal habitats is important for the persistence of populations and can vary depending on oceanic conditions that may affect physical transport processes. We evaluated the survivorship of Chthamalus recruits in Playa San Miguel, Baja California, for over a year to evaluate if the supply of new recruits was of greater importance than the post-recruitment factors of competition and predation in determining adult population size. We hypothesized that the number of Chthamalus recruits would predict the number of adults present and that predation and competition would not play a significant role in determining adult densities. Recruit density was a robust predictor of adult density despite the presence of weak density-dependent mortality. Neither predation nor competition significantly affected the survivorship of recruits and did not decouple the relationship between recruit and adult densities, suggesting that these post-recruitment controls did not play an important role in determining the population size of Chthamalus barnacles at this site. The data support the hypothesis that supply-side control is the primary factor structuring the population of Chthamalus in the intertidal community at Playa San Miguel.     

Resource competition induces heterogeneity and can increase cohort survivorship: selection-event duration matters

Determining when resource competition increases survivorship can reveal processes underlying population dynamics and reinforce the importance of heterogeneity among individuals in conservation. We ran an experiment mimicking the effects of competition in a growing season on survivorship during a selection event (e.g., overwinter starvation, drought). Using a model fish species (Poecilia reticulata), we studied how food availability and competition affect mass in a treatment stage, and subsequently survivorship in a challenge stage of increased temperature and starvation. The post-treatment mean mass was strongly related to the mean time to mortality and mass at mortality at all levels of competition. However, competition increased variance in mass and extended the right tail of the survivorship curve, resulting in a greater number of individuals alive beyond a critical temporal threshold ( $T^{*}$ ) than without competition. To realize the benefits from previously experienced competition, the duration of the challenge ( $T_{c}$ ) following the competition must exceed the critical threshold $T^{*}$ (i.e., competition increases survivorship when $T_{c} > T^{*}$ ). Furthermore, this benefit was equivalent to increasing food availability by 20 % in a group without competition in our experiment. The relationship of $T^{*}$ to treatment and challenge conditions was modeled by characterizing mortality through mass loss in terms of the stochastic rate of loss of vitality (individual’s survival capacity). In essence, when the duration of a selection event exceeds $T^{*}$ , competition-induced heterogeneity buffers against mortality through overcompensation processes among individuals of a cohort. Overall, our study demonstrates an approach to quantify how early life stage heterogeneity affects survivorship.     

Post-settlement emigration affects mortality estimates for two Bahamian wrasses

Understanding the dynamics of open marine populations is inherently complex, and this complexity has led to decades of debate regarding the relative importance of pre- versus post-settlement processes in structuring these populations. Movement between patches may be an important modifier of patterns established at settlement, yet local immigration and emigration have received less attention than other demographic rates. I examined loss rates from tagged populations of juvenile wrasses (yellowhead wrasse Halichoeres garnoti and bluehead wrasse Thalassoma bifasciatum) at two sites in the Bahamas. Assuming that all losses were due solely to mortality would have significantly underestimated survivorship of yellowhead wrasse by 29% and bluehead wrasse by 14%. On average, per capita mortality and emigration rates were higher for yellowhead than bluehead wrasse, but neither demographic rate differed between sites for either species. With respect to within-species density, bluehead wrasse mortality was density-dependent at the patch reef site, but mortality rates of yellowhead wrasse were consistently density-independent. Evaluating the effects of between-species density, yellowhead wrasse mortality increased with a decrease in bluehead wrasse density, but this effect was limited to the patch reef site. Emigration rates were not a function of either within-species or between-species density, but instead varied inversely with isolation distance. Numerous previous studies of coral-reef fish, conducted on patch reefs separated by only a few meters of sand and often using untagged fish, may have confounded losses due to emigration with those due to mortality. A better understanding of the factors affecting emigration in marine fishes is important to their effective management using spatial tools such as marine protected areas.     

Studies on theAbies population of Mt. Shimagare

The survivorship curve is estimated for theAbies population of the subalpine wave-regenerated forest on Mt. Shimagare in the northern Yatsugatake Mountains, Central Japan. The repetition of decay and regeneration of nearly even-aged stands is the nature of this forest in a steady-state, so that the survivorship curve is given as the decrease of density with stand age. Four distinct stages are recognized on the curve. The mortality is high in the first 10 years (stage i). Though germination is observed from the floor under dense canopy to the forest edge, only a portion of the seedlings can survive and form a nearly even-aged young regeneration. The established regeneration enjoys a low mortality before it attains a full density state (stage ii, until 32 years). When full density is reached, as indicated by the ?3/2 power law, the mortality increases again due to an intensive self-thinning (stage iii). However, mortality rate declines with age. This decline is thought to occur because canopy trees become uniform in size and spatial distribution. In the last stage the aggregative dying-off of trees takes place (stage iv, about 90 years). The uniformity among trees is believed to promote the collapse of the stand.     

Spatial pattern of central African rainforests can be predicted from average tree size

When considering all trees irrespective of their species, natural tropical rain forests typically exhibit spatial patterns that range from random to regular. The regularity is often interpreted as a footprint of tree competition. Using 23 permanent sample plots totalling 61 ha in the rain forests of central Africa, we characterized their spatial patterns and modelled those that exhibited regularity by a Strauss point process. This Strauss process is obtained as a Markov point process whose interaction function is an exponential function of a competition index commonly used in forestry. The parameter of this Strauss process characterizes the strength of competition. The 23 plots in central Africa differed in tree density and basal area, and could be discriminated depending on the type of spatial patterns: plots having a large basal area with respect to their density had a non regular pattern, whereas those having a small basal area with respect to their density had a regular pattern. For those plots that exhibited regularity, average tree size could be used to predict the strength of competition. The parameter of the Strauss process was significantly related to the average size by a linear relationship, such that competition decreases as average tree size increases. This relationship extrapolated to a null value of the Strauss parameter when average tree size reaches 32 cm in diameter. This relationship between average tree size and spatial pattern is a testable feature for future studies on the relationship between competition and spatial pattern in natural forests.     

灵空山辽东栎种群年龄结构与动态

利用生存分析理论,分析了山西灵空山自然保护区辽东栎种群的年龄结构,以及树高与胸径的关系,绘制了静态生命表和存活曲线.结果表明： 辽东栎种群年龄结构金字塔呈两头小中间大的纺锤型,处于稳定状态.辽东栎种群的树高和胸径的相关关系可用一元线性方程拟合.辽东栎种群的最高生命期望值出现在死亡率最高的龄级之后,且极大值与死亡密度最小值相对应,说明在经过自然筛选和种内竞争后,存活下来的幼苗幼树具有较强的生命期望寿命.辽东栎种群的存活曲线呈Deevey-Ⅱ型,为稳定种群.辽东栎种群动态表现为前期衰退、中期增长、后期稳定.
     

Intraspecific competition and density dependence in an Ephestia kuehniella–Venturia canescens laboratory system

A model host-parasitoid system of Ephestia kuehniella and Venturia canescens was used to examine the influence of host and parasitoid density on host and parasitoid life-history parameters via a two-way factorial experimental design (5 initial host densities×3 parasitoid densities). In the absence of parasitoids, E. kuehniella experienced scramble-type competition with reduced growth, diminished adult size and a subsequent fecundity trade-off for mortality. The mortality that did occur was confined to the late larval and pupal stages. In the presence of parasitoids attacking the late larval stage, competition changed from scramble for food to contest for enemy-free space, with hosts escaping parasitism being small with low fecundity and reduced egg size, and with parasitoid adult size inversely dependent on host density. Total insect emergence (host+parasitoid), a measure of the influence of host resource competition on survivorship, exhibited a threshold effect as a function of initial host density; the threshold value was increased to a higher initial host density in the presence of parasitoids. Models of host self-limitation were fitted to the data, with the generalized Beverton-Holt model that incorporates a threshold effect providing the best fit, and the Ricker model with no threshold providing a very poor fit to the data.