Can road mortality limit populations of pool-breeding amphibians?

We integrated road maps, traffic volume data, and pool locations in a modeling study to estimate the potential effects of road mortality on populations of pool-breeding spotted salamanders (Ambystoma maculatum Shaw). Population projections based on spotted salamander life tables imply that an annual risk of road mortality for adults of >10% can lead to local population extirpation; mitigation efforts (tunnels, road closures, and other measures) should seek to reduce road mortality rates to below this threshold. For central and western Massachusetts, we estimated that salamanders would be exposed to at least this threshold level of risk at 22–73% of populations (assuming a 100 vs. 500 m migration distance, respectively). We conclude that road mortality can be a significant source of additive mortality for individual spotted salamanders in many parts of the species’ range. Efforts to prevent such mortality by transportation planners are likely warranted strictly on a biological basis in areas with road densities >2.5 km per km² of landscape and traffic volumes >250 vehicles/lane/day within the migration range of a breeding population of spotted salamanders.     

An improved life table method.

A life table estimates probabilities of surviving and of dying as well as death rates, as these would apply in a stationary population with the same underlying continuous mortality curve as the observed population. We have derived approximations to the probability of surviving that require no iteration, do not depend on graduation or interpolation, and appear to give as precise results as interpolated or iterated tables. On the side of theroy we show that methods due to T.N.E. Greville and to Reed and Merrell are special cases of our formula (3). The new approach is extended to cause-deleted tables and to multiple decrement.     

The effects of prorating risk in the development of life‐tables

This study used computer models to investigate two different strategies for assessing risk in the development of age‐based life‐tables from studbook data sets. One methodology is similar to that currently employed in American Zoo and Aquarium Association population management, which prorates animals at risk within age‐classes. The other follows the method used in human life‐tables that assumes animals are at risk for the entire age‐class. This study concludes that prorating risk may invalidate population growth projections by significantly and unequally over‐estimating fecundity and mortality rates. This effect is most pronounced in species that have distinct breeding seasons (birth pulse populations), seasonal mortality, and small data sets. Recommendations include using a non‐prorated methodology, tabulating life‐tables using only completely known age‐class data, and combining population parameters for emigrations, releases, and deaths for population growth projections. Zoo Biol 20:279–291, 2001. © 2001 Wiley‐Liss, Inc.     

An agent-based modelling approach to estimate error in gyrodactylid population growth

Comparative studies of gyrodactylid monogeneans on different host species or strains rely upon the observation of growth on individual fish maintained within a common environment, summarised using maximum likelihood statistical approaches. Here we describe an agent-based model of gyrodactylid population growth, which we use to evaluate errors due to stochastic reproductive variation in such experimental studies. Parameters for the model use available fecundity and mortality data derived from previously published life tables of Gyrodactylus salaris, and use a new data set of fecundity and mortality statistics for this species on the Neva stock of Atlantic salmon, Salmo salar. Mortality data were analysed using a mark-recapture analysis software package, allowing maximum-likelihood estimation of daily survivorship and mortality. We consistently found that a constant age-specific mortality schedule was most appropriate for G. salaris in experimental datasets, with a daily survivorship of 0.84 at 13°C. This, however, gave unrealistically low population growth rates when used as parameters in the model, and a schedule of constantly increasing mortality was chosen as the best compromise for the model. The model also predicted a realistic age structure for the simulated populations, with 0.32 of the population not yet having given birth for the first time (pre-first birth). The model demonstrated that the population growth rate can be a useful parameter for comparing gyrodactylid populations when these are larger than 20-30 individuals, but that stochastic error rendered the parameter unusable in smaller populations. It also showed that the declining parasite population growth rate typically observed during the course of G. salaris infections cannot be explained through stochastic error and must therefore have a biological basis. Finally, the study showed that most gyrodactylid-host studies of this type are too small to detect subtle differences in local adaptation of gyrodactylid monogeneans between fish stocks.     

Periodic dynamics in a two-stage Allee effect model are driven by tension between stage equilibria

Single species difference population models can show complex dynamics such as periodicity and chaos under certain circumstances, but usually only when rates of intrinsic population growth or other life history parameter are unrealistically high. Single species models with Allee effects (positive density dependence at low density) have also been shown to exhibit complex dynamics when combined with over-compensatory density dependence or a narrow fertility window. Here we present a simple two-stage model with Allee effects which shows large amplitude periodic fluctuations for some initial conditions, without these requirements. Periodicity arises out of a tension between the critical equilibrium of each stage, i.e. when the initial population vector is such that the adult stage is above the critical value, while the juvenile stage is below the critical value. Within this area of parameter space, the range of initial conditions giving rise to periodic dynamics is driven mainly by adult mortality rates. Periodic dynamics become more important as adult mortality increases up to a certain point, after which periodic dynamics are replaced by extinction. This model has more realistic life history parameter values than most 'chaotic' models. Conditions for periodic dynamics might arise in some marine species which are exploited (high adult mortality) leading to recruitment limitation (low juvenile density) and might be an additional source of extinction risk.     

A censored quantile regression approach for relative survival analysis: Relative survival quantile regression

We propose a censored quantile regression model for the analysis of relative survival data. We create a hybrid data set consisting of the study observations and counterpart randomly sampled pseudopopulation observations imputed from population life tables that adjust for expected mortality. We then fit a censored quantile regression model to the hybrid data incorporating demographic variables (e.g., age, biologic sex, calendar time) corresponding to the population life tables of demographically-similar individuals, a population versus study covariate, and its interactions with the variables of interest. These latter variables can be interpreted as relative survival parameters that depict the differences in failure quantiles between the study participants and their population counterparts.     

Ecotypic variation in the asymmetric Hawk-Dove game: When is Bourgeois an evolutionarily stable strategy?

Summary This paper develops a mathematical model of an iterated, asymmetric Hawk-Dove game with the novel feature that not only are successive pairs of interactants — in the roles of owner and intruder contesting a site — drawn randomly from the population, but also the behaviour adopted at one interaction affects the role of a contestant in the next. Under the assumption that a site is essential for reproduction, the evolutionarily stable strategy (ESS) of the population is found to depend on the probability, w, that the game will continue for at least a further period (which is inversely related to predation risk), and five other parameters; two of them are measures of site scarcity, two are measures of fighting costs, and the last is a measure of resource holding potential (RHP). Among the four strategies — Hawk (H), Dove (D), Bourgeois (B) and anti-Bourgeois (X) — only D is incapable of being an ESS; and regions of parameter space are found in which the ESS can be only H, or only X, or only B; or either H or X; or either X or B; or either H or B; or any of the three. The scarcer the sites or the lower the costs of fighting, or the lower the value of w, the more likely it is that H is an ESS; the more abundant the sites or the higher the costs of fighting, or the higher the value of w, the more likely it is that X or B is an ESS. The different ESSs are interpreted as different ecotypes. The analysis suggests how a non-fighting population could evolve from a fighting population under decreasing risk of predation. If there were no RHP, or if RHP were low, then the ESS in the non-fighting population would be X; only if RHP were sufficiently high would the ESS be B, and the scarcer the sites, the higher the RHP would have to be. These conclusions support the thesis that if long-term territories are essential for reproduction and sites are scarce, then ownership is ruled out not only as an uncorrelated asymmetry for settling disputes in favour of owner, but also as a correlated asymmetry.     

Population dynamics of Marginopora kudakajimensis Gudmundsson (Foraminifera: Soritidae) in the Ryukyu Islands, the subtropical northwest Pacific

The population dynamics of Marginopora kudakajimensis Gudmundsson, a dinoflagellate endosymbiont-bearing soritid foraminifer, was studied in the Ryukyu Islands, the subtropical northwest Pacific. Macroalgal samples were collected monthly between November 1995 and November 1996 at a 1-m-deep lagoonal site colonized by this species. Monthly variations in the size–frequency distributions and population density indicate that this foraminiferal population replaces itself in one year. Asexual reproduction occurs twice a year, in late spring and winter; in late spring, some of the adult individuals reproduce by multiple fission simultaneously. As a result, two cohorts (a late spring and a winter cohort) are found in the population during a year. Life span of the former cohort is up to one year, while that of the latter cohort is up to six months. Megalospheric specimens comprise up to 99% of the population with a few microspheric individuals throughout the year. The life tables and survivorship curves revealed that size-specific mortality rates were very low during the first several size classes and increased thereafter, indicating low juvenile mortality with high mortality later in life. The carbonate production rate by this Marginopora population is approximately 5 kg CaCO₃ m⁻² yr⁻¹, which is extremely higher than those reported for other larger foraminiferal species.     

不同干扰条件下红豆树种群数量特征的比较

红豆树（Ormosia hosiei）为中国特有种,兼具材用、药用、园艺观赏及科学研究等价值,但是日益严重的干扰对红豆树种群造成了极大影响。本文基于红豆树胸径与年龄之间的关系,初步分析了受干扰程度不同的3个红豆树种群的年龄结构、种群静态生命表、存活曲线及死亡率和消失率曲线。结果显示：受严重干扰的种群幼苗严重不足,种群表现出明显的衰退趋势;近年来受中度干扰的种群幼苗个体数占绝对优势,为各级数量的7877%;长期受轻微干扰的种群年龄级结构表现出为稳定的特征。这3个种群生命表最大的差异在于受干扰最严重种群的第Ⅰ龄级和第Ⅱ龄级的死亡率为负,这说明其幼苗库不足。存活曲线都存在波动,长期受轻微干扰的种群死亡率变幅最小。研究认为,红豆树种质资源和分布面积都有所减少,应加强科学研究和保护,进行适度干扰,使种群朝着进展演替的方向发展。     

Mortality and Life Expectancy in Homeless Men and Women in Rotterdam: 2001–2010

Background

Data on mortality among homeless people are limited. Therefore, this study aimed to describe mortality patterns within a cohort of homeless adults in Rotterdam (the Netherlands) and to assess excess mortality as compared to the general population in that city.

Methods

Based on 10-year follow-up of homeless adults aged ≥ 20 years who visited services for homeless people in Rotterdam in 2001, and on vital statistics, we assessed the association of mortality with age, sex and type of service used (e.g. only day care, convalescence care, other) within the homeless cohort, and also compared mortality between the homeless and general population using Poisson regression. Life tables and decomposition methods were used to examine differences in life expectancy.

Results

During follow-up, of the 2096 adult homeless 265 died. Among the homeless, at age 30 years no significant sex differences were found in overall mortality rates and life expectancy. Compared with the general Rotterdam population, mortality rates were 3.5 times higher in the homeless cohort. Excess mortality was larger in women (rate ratio [RR] RR 5.56, 95% CI 3.95–7.82) as compared to men (RR 3.31, 95% CI 2.91–3.77), and decreased with age (RR 7.67, 95% CI 6.87–8.56 for the age group 20–44 and RR 1.63, 95% CI 1.41–1.88 for the age group 60+ years). Life expectancy at age 30 years was 11.0 (95% CI 9.1–12.9) and 15.9 (95% CI 10.3–21.5) years lower for homeless men and women compared to men and women in the general population respectively.

Conclusion

Homeless adults face excessive losses in life expectancy, with greatest disadvantages among homeless women and the younger age groups.     

Differential mortality in Turkana agriculturalists and pastoralists

Nomadic pastoral populations appear to have much lower rates of growth than the otherwise very high growth rates now characteristic of populations in developing nations. Because dramatic declines in infant mortality have been a primary contributor to increased population growth rates in these countries, it has been assumed that nomadic pastoral populations are still characterized by high levels of mortality in the first few years of life. Few studies, however, have been undertaken to estimate demographic parameters for nomadic pastoral populations, and even fewer of a comparative nature have been undertaken to document the impact of subsistence strategy on demographic processes. This study compares indirect childhood mortality estimates for Turkana nomadic pastoralists with childhood mortality in a settled agricultural group within the same population and finds that pastoralists have substantially higher levels of mortality. Based on the childhood mortality estimates, model life tables are selected for pastoral and agricultural groups from which values for mean life expectancy and infant mortality are estimated and compared. Recent improvements in primary health care for the settled agricultural group are ruled out as being an important cause of their lower mortality levels, and some aspects of life-style associated with subsistence strategy are discussed as likely determinants of the mortality differences.     

Abridged life tables for Pakistan based on the 1971 Population Growth Survey

A new set of abridged life tables for Pakistan is presented. Data from the 1971 Population Growth Survey were 1st analyzed to estimate the degree of completeness of the reporting of male and female deaths; female deaths were substantially more underreported than male deaths. Age-sex specific mortality schedules were adjusted accordingly. Life expectancy was around 50 years at birth, but increased by 8-9 years for those surviving the substantial risks of dying in the 1st year of life. No significant sex differential in mortality could be discerned from the life tables. The present life tables exhibit mortality levels which are very similar to those observed in the life tables based on the 1962-64 Population Growth Estimation data. The only exception in the present life tables is the lack of a significant improvement in female mortality beyond the reproductive ages possibly because the extent of underreporting of femal deaths found in the 1971 Population Growth Survey was substantially higher than that for males.     

Demographic analysis of delayed mating in mating disruption: a case study with Cryptophelbia illepida (Lepidoptera: Tortricidae)

Laboratory-derived life tables were used to determine the effect of delaying mating of adult female koa seedworm, Cryptophlebia illepida (Butler) (Lepidoptera: Tortricidae), 4 and 6 d on population growth rates. Leslie matrices were developed from the life tables and used to project the effects for approximately four generations. Delay of mating caused a decrease in population growth rate and also resulted in asynchronous population cycling between control (1-d delay) and the delayed treatments. By the fourth generation, the control population began to increase 10 and 14 d before the 4- and 6-d delay treatments, respectively. Increasing the mortality of females during the first 7 d of adult life resulted in a greater reduction of the populations where mating was delayed than in the control populations. This result suggests that even at relatively low levels of natural enemy mortality, there is a synergistic effect when mating is delayed. The implications of these effects on mating disruption management programs are discussed.     

Sexual dimorphism in Odonata: age, size, and sex ratio at emergence

Males and females of many organisms differ in important life-history and behavioral characters. Following a recent optimization analysis of sexually dimorphic life histories, we employed an odonate-like parameter set to identify patterns of life history and behavior to be expected in an odonate population. The default parameter magnitudes generated a smaller body size and shorter development time for males than for females, which resulted in a male-biased sex ratio. Whether population growth was density dependent or density independent, and whether development time was fixed or flexible had major impacts on life-history features. The model generated five general predictions for odonate systems. (1) For species with fixed development times, males and females should differ more in activity level, growth and mortality rates than for species with flexible life cycles. (2) In species with fixed development times, populations at high latitude or high altitude should be more active, emerge and reproduce at smaller size and have a more male-biased sex ratio than low latitude and low altitude populations. (3) In density-dependent populations, with density dependence mediated by activity-dependent mortality, higher predation rates should increase activity levels and reduce development time in species with flexible development times. (4) For species with flexible development times, in strongly density-dependent populations with density dependence mediated by mortality, activity levels should decrease and development times should increase at high prey abundance. (5) Males should be larger at emergence relative to females, and the sex ratio at emergence should be more female-biased in territorial than in non-territorial species. Existing empirical evidence concerning these predictions is generally sparse and equivocal; focused tests are clearly needed.     

Unexpected Demography in the Recovery of an Endangered Primate Population

Assessments of the status of endangered species have focused on population sizes, often without knowledge of demographic and behavioral processes underlying population recovery. We analyzed demographic data from a 28-year study of a critically endangered primate, the northern muriqui, to investigate possible changes in demographic rates as this population recovered from near extirpation. As the population increased from 60 to nearly 300 individuals, its growth rate declined due to increased mortality and male-biased birth sex ratios; the increased mortality was not uniform across ages and sexes, and there has been a recent increase in mortality of prime-aged males. If not for a concurrent increase in fertility rates, the population would have stabilized at 200 individuals instead of continuing to grow. The unexpected increase in fertility rates and in adult male mortality can be attributed to the muriquis’ expansion of their habitat by spending more time on the ground. The demographic consequences of this behavioral shift must be incorporated into management tactics for this population and emphasize the importance of understanding demographic rates in the recovery of endangered species.     

Conflicting processes in the evolution of body size and development time

Body size and development time of Manduca sexta are both determined by the same set of three developmental–physiological factors. These define a parameter space within which it is possible to analyse and explain how phenotypic change is associated with changes in the underlying factors. Body size and development time are determined by the identical set of underlying factors, so they are not independent, but because the mechanisms by which these factors produce each phenotype are different, the two phenotypes are only weakly correlated, and the correlation is context dependent. We use a mathematical model of this mechanism to explore the association between body size and development time and show that the correlation between these two life-history traits can be positive, zero or negative, depending entirely on where in parameter space a population is located, and on which of the underlying factors has a greater variation. The gradient within this parameter space predicts the unconstrained evolutionary trajectory under directional selection on each trait. Calculations of the gradients for body size and development time revealed that these are nearly orthogonal through much of the parameter space. Therefore, simultaneous directional selection on body size and development time can be neither synergistic nor antagonistic but leads to conflicting selection on the underlying developmental parameters.     

The population ecology of a natural population of the pierid butterfly Colias alexandra

Summary Key factor analysis techniques were used to examine factors determining the abundance of a population of non-pest Colias. The number of individuals entering each successive life stage in the sample population are summarized in life tables for 1975 to 1979. Survivorship to the adult is a relatively consistent proportionality (-x=1.2%, S.D.=1.14; 1975–1979). Factors resulting in reduced natality and, less importantly, mortality during larval diapause determine the population trends for C. alexandra. Egg mortality, pre-diapause larval mortality and postdiapause mortality contribute little to these trends. Possible key sources contributing to reduced natality are examined. Mortality of adults (including removal by collectors), poor weather conditions during the oviposition period, unseasonal snow or drought which affect nectar sources or oviposition sites are among the factors which cause reduced natality and result in population depression.     

Annual league tables of mortality in neonatal intensive care units: longitudinal study

Objective: To assess whether crude league tables of mortality and league tables of risk adjusted mortality accurately reflect the performance of hospitals. Design: Longitudinal study of mortality occurring in hospital. Setting: 9 neonatal intensive care units in the United Kingdom. Subjects: 2671 very low birth weight or preterm infants admitted to neonatal intensive care units between 1988 and 1994. Main outcome measures: Crude hospital mortality and hospital mortality adjusted using the clinical risk index for babies (CRIB) score. Results: Hospitals had wide and overlapping confidence intervals when ranked by mortality in annual league tables; this made it impossible to discriminate between hospitals reliably. In most years there was no significant difference between hospitals, only random variation. The apparent performance of individual hospitals fluctuated substantially from year to year. Conclusions: Annual league tables are not reliable indicators of performance or best practice; they do not reflect consistent differences between hospitals. Any action prompted by the annual league tables would have been equally likely to have been beneficial, detrimental, or irrelevant. Mortality should be compared between groups of hospitals using specific criteria—such as differences in the volume of patients, staffing policy, training of staff, or aspects of clinical practice—after adjusting for risk. This will produce more reliable estimates with narrower confidence intervals, and more reliable and rapid conclusions.

Key messages

• League tables are being used increasingly to evaluate hospital performance in the United Kingdom
• In annual league tables the rankings of nine neonatal intensive care units in different hospitals had wide and overlapping confidence intervals and their rankings fluctuated substantially over six years
• Annual league tables of hospital mortality were inherently unreliable for comparing hospital performance or for indicating best practices
• The UK government’s commitment to using annual league tables of outcomes such as mortality to monitor services and the spread of best practices should be reconsidered
• Prospective studies of risk adjusted outcome in hospitals grouped according to specific characteristics would provide better information and be a better use of resources

     

Impact of Pregnancy-Related Deaths on Female Life Expectancy in Zambia: Application of Life Table Techniques to Census Data

Introduction

Since 2000, the world has been coalesced around efforts to reduce maternal mortality. However, few studies have estimated the significance of eliminating maternal deaths on female life expectancy. We estimated, based on census data, the potential gains in female life expectancy assuming complete elimination of pregnancy-related mortality in Zambia.

Methods

We used data on all-cause and pregnancy-related deaths of females aged 15–49 reported in the Zambia 2010 census, and evaluated, adjusted and smoothed them using existing and verified techniques. We used associated single decrement life tables, assuming complete elimination of pregnancy-related deaths to estimate the potential gains in female life expectancy at birth, at age 15, and over the ages 15–49. We compared these gains with the gains from eliminating deaths from accidents, injury, violence and suicide.

Results

Complete elimination of pregnancy-related deaths would extend life expectancy at birth among Zambian women by 1.35 years and life expectancy at age 15 by 1.65 years. In rural areas, this would be 1.69 years and 2.19 years, respectively, and in urban areas, 0.78 years and 0.85 years. An additional 0.72 years would be spent in the reproductive age group 15–49; 1.00 years in rural areas and 0.35 years in urban areas. Eliminating deaths from accidents, injury, suicide and violence among women aged 15–49 would cumulatively contribute 0.55 years to female life expectancy at birth.

Conclusion

Eliminating pregnancy-related mortality would extend female life expectancy in Zambia substantially, with more gains among adolescents and females in rural areas. The application of life table techniques to census data proved very valuable, although rigorous evaluation and adjustment of reported deaths and age was necessary to attain plausible estimates. The collection of detailed high quality cause-specific mortality data in future censuses is indispensable.     

Evolution of reproductive effort in a metapopulation with local extinctions and ecological succession

Abstract Using a metapopulation model, we study how local extinctions, limited population life span, and local demographic disequilibrium affect the evolution of the reproductive effort in a species with overlapping generations but no senescence. We show that in a metapopulation with saturation of all sites and an infinite deme maximal life span (no succession), local extinctions simply constitute an additional source of extrinsic mortality. When either the hypothesis of an infinite deme maximal life span or the saturation hypothesis is relaxed, nontrivial predictions arise. in particular, we find interactions between the evolutionarily stable reproductive effort strategy and the demographic dynamics in the metapopulation. We predict that larger reproductive effort may be selected for in habitats of poorer productivity, contrary to what would be predicted in a single population. Also, we predict that higher dispersal rates should favor selection for lower reproductive efforts. However, metapopulation parameters that favor high dispersal rates also favor larger reproductive efforts. Conflicting selection pressures in the metapopulation also allow maintaining evolutionarily stable polymorphism between a low and high reproductive effort for particular trade-offs between survival and fecundity.