New approaches to the pre- and post-contact history of Arctic peoples.

The last few years have witnessed the addition of new techniques and research strategies to the study of the population history of Arctic peoples. Osteon-photon analysis of bone cores provides an improved method of assigning age at death to skeletons. Consequently, it is possible to improve calculations of life expectancy and relate them to pathological correlates such as osteoporosis, separate neural arches, spina bifida and arthritis along with regular growth changes. This capability enables much better utilization of pre-contact skeletons and therefore of the numbers, density and composition of populations before European contact. The general picture emerging from skeletal studies, census records and living populations is, in Arctic Eskimos, one of high fertility, high mortality and short length of life, with a slow population growth rate. Aleuts show lower fertility, lower mortality and longer length of life, also with a low population growth rate.     

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.     

On generating birth rates from skeletal populations

Sattenspiel and Harpending (1983, American Antiquity 48(3): 489-498) have stated that the life expectancy at birth (e0(0] which paleodemographers calculate from skeletal population data is actually the mean age at death (ad) of the population. Yet, only when a population is neither growing or declining (i.e., is stationary) are these two statistics equivalent. They further assert, that the mean age at the death (ad) is more accurately interpreted as a measure of the fertility of the population. While we support their statement that since paleodemographic calculations use skeletal evidence of death, these do not a priori produce life expectancy values, we disagree that the inverse of the birth rate is a substitute for the average age at death (ad). The following pages demonstrate that: 1) An exact expression for the relationship between ad and 1/b can be derived using standard stable population theory, wherein ad = 1/b is shown to be a special case. 2) There are only two cases when ad = 1/b is an identity. 3) Whereas empirically ad and 1/b appear to correspond closely, this is an artifact of heavy mortality at early ages, which is a characteristic of the populations being considered. 4) Without insights into the behavioral dynamics of the situation any assessment of the demographics of the population is questionable.     

Population and prehistory I: Food-dependent population growth in constant environments

We present a demographic model that describes the feedbacks between food supply, human mortality and fertility rates, and labor availability in expanding populations, where arable land area is not limiting. This model provides a quantitative framework to describe how environment, technology, and culture interact to influence the fates of preindustrial agricultural populations. We present equilibrium conditions and derive approximations for the equilibrium population growth rate, food availability, and other food-dependent measures of population well-being. We examine how the approximations respond to environmental changes and to human choices, and find that the impact of environmental quality depends upon whether it manifests through agricultural yield or maximum (food-independent) survival rates. Human choices can complement or offset environmental effects: greater labor investments increase both population growth and well-being, and therefore can counteract lower agricultural yield, while fertility control decreases the growth rate but can increase or decrease well-being. Finally we establish equilibrium stability criteria, and argue that the potential for loss of local stability at low population growth rates could have important consequences for populations that suffer significant environmental or demographic shocks.     

Life history implications of allocation to growth versus reproduction in dynamic energy budgets

We compare the implications of determinate vs. indeterminate growth of a parthenogenetic iteroparous ectotherm at constant food density in the context of the dynamic energy budget theory, which specifies the tight links between life history traits, such as feeding, aging, growth and reproduction. We do a comparative analysis using, as measure of fitness, the life span reproduction, the population growth rate, and the conversion efficiency of food to biomass. When extrinsic mortality is constant, indeterminate growth cannot maximize fitness if measured by the population growth rate or the conversion efficiency, except when mortality is low, in which case both types of animals are similar. If the fitness measure is life span reproduction, indeterminate growth maximizes fitness even with constant mortality, provided it is not very high. When mortality decreases with size, indeterminate growth maximizes fitness for almost all measures of fitness. Finally, we suggest an evolutionary link between allocation strategies and expected life span. In populations of long living species, each type of animal can establish in the population of the other. In populations of short living species, determinate growers can invade, and displace, a population of indeterminate ones. However, when the mortality risk of organisms with small size is much higher than those of large size, indeterminate growers can be superior.     

Freshwater goby life history in a Mediterranean stream

The Italian goby Gobius nigricans is a running-water-dwelling species inhabiting the Tuscano-Latium district and threatened by several human activities. Many aspects of its biology are currently unknown, such as the dynamic potentialities. Since age determination and dynamic assessment are considered an important tool for a correct management strategy, we aimed to investigate some aspects of the Italian goby life history by applying the fish stock assessment principles, collecting data on fertility, population structure (age-class number), growth (i.e., curvature parameter and asymptotic length), mortality (natural and due to the fishing), and longevity of a central Italian goby population to know the status and dynamic properties of the Italian goby in Mediterranean river systems and to provide a baseline for comparison with other populations. Females are subject to a strong selective pressure to quickly reach a small size to optimize reproductive output, while acquisition of a large size favored males for a more easy access to breeder females (the asymptotic length was higher in males than in females). Moreover, appreciable between-sex differences were found in both growth pattern (the growth rate was lower in females than in males) and dynamic potentialities. In particular, the mortality rate was lower in females, whereas the longevity was slightly lower in males, although the two sexes showed similar values for the two latter properties. There is considerable scope for further work on the G. nigricans growth and other freshwater gobies because information on population and dynamic properties, and assessment of factors affecting growth are still very insufficient. Handling editor: P. Viaroli     

Evaluating the consequences of habitat fragmentation: a case study in the common forest herb Trillium camschatcense

The effects of habitat fragmentation on remnant plant populations have rarely been studied extensively using a single species. We have attempted to quantify the effects of forest fragmentation (primarily that of population size) on populations of Trillium camschatcense, a representative spring herb in the Tokachi plain of Hokkaido, Japan. In this region, intensive agricultural development over the past 100 years has divided once-large, continuous populations of this species into small, isolated fragments. Small populations generally produced fewer seeds than large populations, although this result differed between years. The level of seed production is unlikely to explain demographic structures based on life-history stages. Instead, the stage structure was better explained by population size, seedling recruitment being limited in smaller populations. This could be associated with edge effects because the stage structure in small populations corresponded well to that observed in forest edges, where altered microclimatic conditions strongly limit seedling recruitment. Small populations also experienced stochastic loss of rare alleles at allozyme loci as well as biparental inbreeding. Although one consequence of these changes is reduced fertility, the long-term effects on population growth can be controversial in long-lived forest herbs, since the negative effect on fertility may vary across years, and population growth rate may not be sensitive to changes in fertility. Further studies of long-term demography will reveal whether and how habitat fragmentation could limit population growth of remnant populations more than a century after fragmentation.     

A General Method for Modeling Population Dynamics and Its Applications

Studying populations, be it a microbe colony or mankind, is important for understanding how complex systems evolve and exist. Such knowledge also often provides insights into evolution, history and different aspects of human life. By and large, populations’ prosperity and decline is about transformation of certain resources into quantity and other characteristics of populations through growth, replication, expansion and acquisition of resources. We introduce a general model of population change, applicable to different types of populations, which interconnects numerous factors influencing population dynamics, such as nutrient influx and nutrient consumption, reproduction period, reproduction rate, etc. It is also possible to take into account specific growth features of individual organisms. We considered two recently discovered distinct growth scenarios: first, when organisms do not change their grown mass regardless of nutrients availability, and the second when organisms can reduce their grown mass by several times in a nutritionally poor environment. We found that nutrient supply and reproduction period are two major factors influencing the shape of population growth curves. There is also a difference in population dynamics between these two groups. Organisms belonging to the second group are significantly more adaptive to reduction of nutrients and far more resistant to extinction. Also, such organisms have substantially more frequent and lesser in amplitude fluctuations of population quantity for the same periodic nutrient supply (compared to the first group). Proposed model allows adequately describing virtually any possible growth scenario, including complex ones with periodic and irregular nutrient supply and other changing parameters, which present approaches cannot do.     

Population Variation in the Life History of a Land Fish,Alticus arnoldorum,and the Effects of Predation and Density

Life history variation can often reflect differences in age-specific mortality within populations, with the general expectation that reproduction should be shifted away from ages experiencing increased mortality. Investigators of life history in vertebrates frequently focus on the impact of predation, but there is increasing evidence that predation may have unexpected impacts on population density that in turn prompt unexpected changes in life history. There are also other reasons why density might impact life history independently of predation or mortality more generally. We investigated the consequences of predation and density on life history variation among populations of the Pacific leaping blenny, Alticus arnoldorum. This fish from the island of Guam spends its adult life out of the water on rocks in the splash zone, where it is vulnerable to predation and can be expected to be sensitive to changes in population density that impact resource availability. We found populations invested more in reproduction as predation decreased, while growth rate varied primarily in response to population density. These differences in life history among populations are likely plastic given the extensive gene flow among populations revealed by a previous study. The influence of predation and density on life history was unlikely to have operated independently of each other, with predation rate tending to be associated with reduced population densities. Taken together, our results suggest predation and density can have complex influences on life history, and that plastic life history traits could allow populations to persist in new or rapidly changing environments.     

Harvesting in a resource dependent age structured Leslie type population model

We analyse the effect of harvesting in a resource dependent age structured population model, deriving the conditions for the existence of a stable steady state as a function of fertility coefficients, harvesting mortality and carrying capacity of the resources. Under the effect of proportional harvest, we give a sufficient condition for a population to extinguish, and we show that the magnitude of proportional harvest depends on the resources available to the population. We show that the harvesting yield can be periodic, quasi-periodic or chaotic, depending on the dynamics of the harvested population. For populations with large fertility numbers, small harvesting mortality leads to abrupt extinction, but larger harvesting mortality leads to controlled population numbers by avoiding over consumption of resources. Harvesting can be a strategy in order to stabilise periodic or quasi-periodic oscillations in the number of individuals of a population.     

DOES INCREASED MORTALITY FAVOR THE EVOLUTION OF MORE RAPID SENESCENCE?

It is widely believed (following the 1957 hypothesis of G. C. Williams) that greater rates of “extrinsic” (age- and condition-independent) mortality favor more rapid senescence. However, a recent analysis of mammalian life tables failed to find a significant correlation between minimum adult mortality rate and the rate of senescence. This article presents a simple theoretical analysis of how extrinsic mortality should affect the rate of senescence (i.e., the rate at which probability of mortality increases with age) under different evolutionary and population dynamical assumptions. If population dynamics are density independent, extrinsic mortality should not alter the senescence rate favored by natural selection. If population growth is density dependent and populations are stable, the effect of extrinsic mortality depends on the age specificity of the density dependence and on whether survival or reproduction (or both) are functions of density. It is possible that higher extrinsic mortality will increase the rate of senescence at all ages, decrease the rate at all ages, or increase it at some ages while decreasing it at others. Williams's hypothesis is most likely to be supported when density dependence acts primarily on fertility and does not differentially decrease the fertilities of older individuals. Patterns contrary to Williams's prediction are possible when density dependence acts primarily on the survival or fertility of later ages or when most variation in mortality rates is due to variation in nonextrinsic mortality.     

Causes of early human population growth

The archaeological record indicates large increases in human population coincident with the emergence of food production about 10,000 years ago. The cause of the growth is unclear. Extreme views attribute the change to increases in the birth rate or to decreases in the death rate. Many argue that sedentism led to improved ovarian function and higher fertility through higher caloric intakes or reduced activity levels. Similarly, shortened lactation periods may have reduced birth spacing and increased fertility. Others attribute the rise in population to decreases in mortality, arguing that the evidence from skeletal populations indicates improvements in health and the expectation of life at birth, though others use the same evidence to argue that mortality increased. An analysis presented here draws on findings that indicate substantial increases in the survival of young children as populations switch from nomadic to sedentary lives. Projections indicate that this improvement in child survival is so critical that it may be followed by substantially larger decreases in survival at later ages, yet result in higher population growth rates and reduced expectation of life at birth. Increases in the birth rate are not necessary for population growth, even when overall mortality increases. Large increases in overall mortality can be associated with large increases in population. Because positive population growth can occur while the expectation of life at birth declines, this analysis shows that this summary statistic is not an appropriate indicator of population fitness. © 1996 Wiley-Liss, Inc.     

Closing the larval loop: linking larval ecology to the population dynamics of marine benthic invertebrates

The majority of marine benthic invertebrates exhibit a complex life cycle that includes separate planktonic larval, and bottom-dwelling juvenile and adult phases. To understand and predict changes in the spatial and temporal distributions, abundances, population growth rate, and population structure of a species with such a complex life cycle, it is necessary to understand the relative importance of the physical, chemical and biological properties and processes that affect individuals within both the planktonic and benthic phases. To accomplish this goal, it is necessary to study both phases within a common, quantitative framework defined in terms of some common currency. This can be done efficiently through construction and evaluation of a population dynamics model that describes the complete life cycle.

Two forms that such a model might assume are reviewed: a stage-based, population matrix model, and a model that specifies discrete stages of the population, on the bottom and in the water column, in terms of simultaneous differential equations that may be solved in both space and time. Terms to be incorporated in each type of model can be formulated to describe the critical properties and processes that can affect populations within each stage of the life cycle. For both types of model it is shown how this might be accomplished using an idealized balanomorph barnacle as an example species. The critical properties and processes that affect the planktonic and benthic phases are reviewed. For larvae, these include benthic adult fecundity and fertilization success, growth and larval stage duration, mortality, larval behavior, dispersal by currents and turbulence, and larval settlement. It is possible to predict or estimate empirically all of the key terms that should be built into the larval and benthic components of the model. Thus, the challenge of formulating and evaluating a full life cycle model is achievable. Development and evaluation of such a model will be challenging because of the diverse processes which must be considered, and because of the disparities in the spatial and temporal scales appropriate to the benthic and planktonic larval phases. In evaluating model predictions it is critical that sampling schemes be matched to the spatial and temporal scales of model resolution.     

Kin-structured migration and the rate of advance of an advantageous gene

Hemoglobin E, an allele generally considered to confer malarial resistance in heterozygotes, is found in high frequencies across a wide area of Southeast Asia. Apparently it originated as a single-point mutation which was spread by gene flow. The rate of diffusion of this adaptive allele is studied using four computer simulation models. It is shown that in small populations deterministic equations for gene flow may overestimate rates of diffusion. Other aspects of population structure, however, such as kin-structuring of migrant groups, increase the rate of advance. Finally, population growth coupled with the spread of the allele leads to much more rapid diffusion. These results suggest that population structure can be an important factor affecting the diffusion of advantageous genes.     

Statistical inference on associated fertility life table parameters using jackknife technique: computational aspects

Knowledge of population growth potential is crucial for studying population dynamics and for establishing management tactics for pest control. Estimation of population growth can be achieved with fertility life tables because they synthesize data on reproduction and mortality of a population. The five main parameters associated with a fertility life table are as follows: (1) the net reproductive rate (Ro), (2) the intrinsic rate of increase (rm), 3) the mean generation time (T), (4) the doubling time (Dt), and (5) the finite rate of increase (lambda). Jackknife and bootstrap techniques are used to calculate the variance of the rm estimate, which can be extended to the other parameters of life tables. Those methods are computer-intensive, their application requires the development of efficient algorithms, and their implementation is based on a programming language that encompasses quickness and reliability. The objectives of this article are to discuss statistical and computational aspects related to estimation of life table parameters and to present a SAS program that uses jackknife to estimate parameters for fertility life tables. The SAS program presented here allows the calculation of confidence intervals for all estimated parameters, as well as provides one-sided and two-sided t-tests to perform pairwise or multiple comparison between groups, with their respective P values.     

Demographic variance in heterogeneous populations: matrix models and sensitivity analysis

The demographic consequences of stochasticity in processes such as survival and reproduction are modulated by the heterogeneity within the population. Therefore, to study effects of stochasticity on population growth and extinction risk, it is critical to use structured population models in which the most important sources of heterogeneity (e.g. age, size, developmental stage) are incorporated as i‐state variables. Demographic stochasticity in heterogeneous populations has often been studied using one of two approaches: multitype branching processes and diffusion approximations. Here, we link these approaches, through the demographic stochasticity in age‐ or stage‐structured matrix population models. We derive the demographic variance, σ²_d, which measures the per capita contribution to the variance in population growth increment, and we show how it can be decomposed into contributions from transition probabilities and fertility across ages or stages. Furthermore, using matrix calculus we derive the sensitivity of σ²_d to age‐ or stage‐specific mortality and fertility. We apply the methods to an extensive set of data from age‐classified human populations (long‐term time‐series for Sweden, Japan and the Netherlands; two hunter–gatherer populations, and the high‐fertility Hutterites), and to a size‐classified population of the herbaceous plant Calathea ovandensis. For the human populations our analysis reveals substantial temporal changes in the demographic variance as well as its main components across age. These new methods provide a powerful approach for calculating the demographic variance for any structured model, and for analyzing its main components and sensitivities. This will make possible new analyses of demographic variance across different kinds of heterogeneity in different life cycles, which will in turn improve our understanding of mechanisms underpinning extinction risk and other important biological outcomes.     

Mortality,fertility, and the OY ratio in a model hunter–gatherer system

An agent‐based model (ABM) is used to explore how the ratio of old to young adults (the OY ratio) in a sample of dead individuals is related to aspects of mortality, fertility, and longevity experienced by the living population from which the sample was drawn. The ABM features representations of rules, behaviors, and constraints that affect person‐ and household‐level decisions about marriage, reproduction, and infant mortality in hunter–gatherer systems. The demographic characteristics of the larger model system emerge through human‐level interactions playing out in the context of “global” parameters that can be adjusted to produce a range of mortality and fertility conditions. Model data show a relationship between the OY ratios of living populations (the living OY ratio) and assemblages of dead individuals drawn from those populations (the dead OY ratio) that is consistent with that from empirically known ethnographic hunter–gatherer cases. The dead OY ratio is clearly related to the mean ages, mean adult mortality rates, and mean total fertility rates experienced by living populations in the model. Sample size exerts a strong effect on the accuracy with which the calculated dead OY ratio reflects the actual dead OY ratio of the complete assemblage. These results demonstrate that the dead OY ratio is a potentially useful metric for paleodemographic analysis of changes in mortality and mean age, and suggest that, in general, hunter–gatherer populations with higher mortality, higher fertility, and lower mean ages are characterized by lower dead OY ratios. Am J Phys Anthropol 154:222–231, 2014. © 2014 Wiley Periodicals, Inc.     

Modeling the prehistoric Maori population

Skeletal and comparative evidence of mortality is combined with fertility estimates for the precontact Maori population of New Zealand to determine the implied rate of precontact population growth. This rate is found to be too low to populate New Zealand within the time constraints of its prehistoric sequence, the probable founding population size, and the probable population size at contact. Rates of growth necessary to populate New Zealand within the accepted time span are calculated. The differences between this minimum necessary rate and the skeletally derived rate are too large to result solely from inadequacies in the primary data. Four alternative explanations of this conundrum are proposed: 1) skeletal evidence of precontact mortality is highly inaccurate; 2) skeletal evidence of fertility is severely underestimating actual levels; 3) there was very rapid population growth in the earliest part of the sequence up to 1150 A.D., from which no skeletal evidence currently is available; or 4) the prehistoric sequence of New Zealand may have been longer than the generally accepted 1,000-1,200 years. These alternatives are examined, and a combination of the last two is found to be the most probable. The implications of this model for New Zealand prehistory and Oceanic paleodemography are discussed.     

Offspring fitness in relation to population size and genetic variation in the rare perennial plant species Gentiana pneumonanthe (Gentianaceae)

Seeds were sampled from 19 populations of the rare Gentiana pneumonanthe, ranging in size from 5 to more than 50,000 flowering plants. An analysis was made of variation in a number of life-history characters in relation to population size and offspring heterozygosity (based on seven polymorphic isozyme loci). Life-his-tory characters included seed weight, germination rate, proportion of seeds germinating, seedling mortality, seedling weight, adult weight, flower production per plant and proportion of plants flowering per family. Principal component analysis (PCA) reduced the dataset to three main fitness components. The first component was highly correlated with adult weight and flowering performance, the second with germination performance and the third component with seed and seedling weight and seedling mortality. The latter two components were considered as being maternally influenced, since these comprised life-history traits that were significantly correlated with seed weight. Multiple regression analysis showed that variation in the first fitness component was mainly associated with heterozygosity and not with population size, while the third fitness component was only correlated with population size and not with heterozygosity. The latter relationship appeared to be non-linear, which suggests a stronger loss of fitness in the smallest populations. The second (germination) component was neither correlated with population size nor with genetic variation. There was only a weak association between population size, heterozygosity and the population coefficients of variation for each life history character. Most correlation coefficients were negative, however, which suggests that there is more variation among progeny from smaller populations. We conclude that progeny from small populations of Gentiana pneumonanthe show reduced fitness and may be phenotypically more variable. One of the possible causes of the loss of fitness is a combination of unfavourable environmental circumstances for maternal plants in small populations and increased inbreeding. The higher phenotypic variation in small populations may also be a result of inbreeding, which can lead to deviation of individuals from the average phenotype through a loss of developmental stability.