Evolution drives, and is driven by, demography. A genotype moulds its phenotype’s age patterns of mortality and fertility in an environment; these two patterns in turn determine the genotype’s fitness in that environment. Hence, to understand the evolution of ageing, age patterns of mortality and reproduction need to be compared for species across the tree of life. However, few studies have done so and only for a limited range of taxa. Here we contrast standardized patterns over age for 11 mammals, 12 other vertebrates, 10 invertebrates, 12 vascular plants and a green alga. Although it has been predicted that evolution should inevitably lead to increasing mortality and declining fertility with age after maturity, there is great variation among these species, including increasing, constant, decreasing, humped and bowed trajectories for both long- and short-lived species. This diversity challenges theoreticians to develop broader perspectives on the evolution of ageing and empiricists to study the demography of more species.
Summary1. Schedules of survival, growth and reproduction are key life-history traits. Data on how these traits vary among species and populations are fundamental to our understanding of the ecological conditions that have shaped plant evolution. Because these demographic schedules determine population *Correspondence author. E-mails: salguero@demogr.mpg.de; compadre-contact@demogr.mpg.de † Joint senior author. This is an open access article under the terms of the Creative Commons Attribution-NonCommercial License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited and is not used for commercial purposes. 2015, 103, 202-218 doi: 10.1111/1365-2745.12334 growth or decline, such data help us understand how different biomes shape plant ecology, how plant populations and communities respond to global change and how to develop successful management tools for endangered or invasive species. Journal of Ecology2. Matrix population models summarize the life cycle components of survival, growth and reproduction, while explicitly acknowledging heterogeneity among classes of individuals in the population. Matrix models have comparable structures, and their emergent measures of population dynamics, such as population growth rate or mean life expectancy, have direct biological interpretations, facilitating comparisons among populations and species. 3. Thousands of plant matrix population models have been parameterized from empirical data, but they are largely dispersed through peer-reviewed and grey literature, and thus remain inaccessible for synthetic analysis. Here, we introduce the COMPADRE Plant Matrix Database version 3.0, an opensource online repository containing 468 studies from 598 species world-wide (672 species hits, when accounting for species studied in more than one source), with a total of 5621 matrices. COMPADRE also contains relevant ancillary information (e.g. ecoregion, growth form, taxonomy, phylogeny) that facilitates interpretation of the numerous demographic metrics that can be derived from the matrices. 4. Synthesis. Large collections of data allow broad questions to be addressed at the global scale, for example, in genetics (GENBANK), functional plant ecology (TRY, BIEN, D3) and grassland community ecology (NUTNET). Here, we present COMPADRE, a similarly data-rich and ecologically relevant resource for plant demography. Open access to this information, its frequent updates and its integration with other online resources will allow researchers to address timely and important ecological and evolutionary questions.
The human lifespan has traversed a long evolutionary and historical path, from short-lived primate ancestors to contemporary Japan, Sweden, and other longevity frontrunners. Analyzing this trajectory is crucial for understanding biological and sociocultural processes that determine the span of life. Here we reveal a fundamental regularity. Two straight lines describe the joint rise of life expectancy and lifespan equality: one for primates and the second one over the full range of human experience from average lifespans as low as 2 y during mortality crises to more than 87 y for Japanese women today. Across the primate order and across human populations, the lives of females tend to be longer and less variable than the lives of males, suggesting deep evolutionary roots to the male disadvantage. Our findings cast fresh light on primate evolution and human history, opening directions for research on inequality, sociality, and aging.biodemography | equality | lifespan | pace and shape | senescence L ongevous populations have two characteristics: The average length of life is long and relative variation in lifespans is low. For example, life tables for contemporary Sweden and Japan indicate that most deaths occur at ages between the late 70s and early 90s. Our primate relatives, in contrast, have lifespans that are highly variable in length but short on average and rarely longer than 30 y (Fig. 1). An association between the average length of life and its variability has been found for industrialized societies (1, 2). However, detailed knowledge is lacking about whether and how this association varies across species separated by millions of years of primate evolution or whether it has changed over the past several centuries of unprecedented social progress in human populations. Fuller comprehension of the relationship between rising lifespans and reduced lifespan variability across evolution and history holds potential insights that might illuminate past, current, and future longevity.We pose three related questions aimed at filling this knowledge gap: How long and variable are lifespans for humans compared with nonhuman primates, for humans today compared with the past, and for males compared with females? We provide answers to these questions by applying a powerful framework that simultaneously examines changes in both the average length of life in a population or species-the "pace" of life-and relative variation in the length of life, i.e., the "shape" of the distribution of ages at death (3-5). Studying variation in lifespan links to increasing interest in social, economic, and health inequalities and to key sociological findings that relate social factors-including high social status and social integration-to longer, healthier lifespans in human and animal societies (6-10).Estimating the average length of life (here measured by life expectancy, the mean age at death) and variation in lifespans relative to the average (measured here as "lifespan equality"; Box 1) requires data on the ages at death of individuals...
Summary 1. Humans age, but how much more or less do we age compared with other species? Do humans age more than chimps, birds more than fish or sheep more than buffalos? In this article, I argue that current methods to compare patterns of ageing across species are limited because they confound two dimensions of age‐specific change – the pace and the shape of ageing. 2. Based on the two axes of pace and shape, I introduce a new conceptual framework to classify how species age. 3. With this method, I rank species according to how strongly they age (shape) and how fast they age (pace). Depending on whether they are ranked by pace or by shape, species are ordered differently. 4. Alternative pace measures turn out to be highly correlated. Alternative shape measures are also highly correlated. The correlation between pace and shape ranking is negative but weak. Among the examples here, no species is long lived yet exhibits negligible ageing – contrary to the commonly held view that long‐lived species are good candidates for negligible ageing. 5. Analysis of species in pace–shape space provides a tool to identify key determinants of the evolution of ageing for species across the tree of life.
To quantify the force of selection, Hamilton [Hamilton, W. D. (1966) J. Theor. Biol. 12, 12-45] derived expressions for the change in fitness with respect to age-specific mutations. Hamilton's indicators are decreasing functions of age. He concluded that senescence is inevitable: survival and fertility decline with age. I show that alternative parameterizations of mutational effects lead to indicators that can increase with age. I then consider the case of deleterious mutations with age-specific effects. In this case, it is the balance between mutation and selection pressure that determines the equilibrium number of mutations in a population. In this balance, the effects of different parameterizations cancel out, but only to a linear approximation. I show that mutation accumulation has little impact at ages when this linear approximation holds. When mutation accumulation matters, nonlinear effects become important, and the parameterizations of mutational effects make a difference. The results also suggest that mutation accumulation may be relatively unimportant over most of the reproductive lifespan of any species.
International audienceIt is commonly thought that temporal fluctuations in demographic parameters should be selected against because of the deleterious impacts variation can have on fitness. A critical underpinning of this prediction is the assumption that changes in environmental conditions map linearly into changes in demographic parameters over time. We detail why this assumption may often break down and why selection should not always favor buffering of demographic parameters against environmental stochasticity. To the contrary, nonlinear relationships between the environment and demographic performance can produce asymmetric temporal variation in demographic parameters that actually enhances fitness. We extend this result to structured populations using simulation and show that ‘demographic lability' rather than ‘buffering' may be adaptive, particularly in organisms with low juvenile or adult survival. Finally, we review previous ecological work, and indicate cases where ‘demographic lability' may be adaptive, then conclude by identifying research that is needed to develop a theory of life-history evolution that encompasses both demographic buffering and lability
Summary1. Demographic senescence, the decay in fertility and increase in the risk of mortality with age, is one of the most striking phenomena in ecology and evolution. Comparative studies of senescence patterns of plants are scarce, and consequently, little is known about senescence and its determinants in the plant kingdom. 2. Senescence patterns of mortality can be classified by distinguishing between two metrics: pace and shape. The pace of mortality captures the speed at which life proceeds and can be measured by life expectancy, while the shape of mortality captures whether mortality increases ('senescence'), decreases ('negative senescence') or remains constant over age ('negligible senescence'). 3. We extract mortality trajectories from ComPADRe III, a data base that contains demographic information for several hundred plant species. We apply age-from-stage matrix decomposition methods to obtain age-specific trajectories from 290 angiosperm species of various growth forms distributed globally. From these trajectories, we survey pace and shape values and investigate how growth form and ecoregion influence these two aspects of mortality using a Bayesian regression analysis that accounts for phylogenetic relationships using a resolved supertree. 4. In contrast to the animal kingdom, most angiosperms (93%) show no senescence. Senescence is observed among phanerophytes (i.e. trees), but not among any other growth form (e.g. epiphytes, chamaephytes or cryptopyhtes). Yet, most phanerophytes (81%) do not senesce. We find that growth form relates to differences in pace, that is, life span, as woody plants are typically longer lived than nonwoody plants, while differences in shape, that is, whether or not angiosperms senesce, are related to ancestral history. 5. Synthesis: The age trajectory of mortality captures a fundamental life-history pattern for a species that is crucial to ecological understanding. We contribute to ecological knowledge by surveying these patterns across angiosperms. The novelty and strength of our study lies in the comprehensiveness of the data set, the use of a novel Bayesian analysis that accounts for phylogenetic history and in the distinction between metrics of pace and shape as two separate aspects of mortality. We believe that our approach could prove useful in future comparative studies of mortality patterns.
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