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Moorad et al [1] reinforces and elaborates on warnings made previously [2, 3] that Williams’ “hypothesis” should be treated with scepticism. A proper hypothesis should be based upon the best information available at the time, and for the case of the evolutionary theory of senescence, Hamilton’s model of selection is superior to Williams’. Hamilton’s insights improved over Williams’ by his appreciation for how fertility and juvenile survival schedules play a critical role in defining selection. While Williams’ model is not articulated mathematically, it is clear that his hypothesis is motivated by his belief that the strength of selection against mortality specific to some age follows from the frequency of individuals that survived to that age [4]. However, Hamilton [5] was quite clear: the strength of selection is equal to the proportion of newborns that came from parents that survived to that age (see [1] or Equation 1 in the Appendix). Furthermore, Williams’ explicitly states that juvenile mortality cannot affect the evolution of senescence while Hamilton’s formulae show us that it can. It should be emphasized that Williams himself came to appreciate that Hamilton’s model describes the best way to think about how selection works [e.g., 6].

The evolutionary theory of senescence underpins research in life history evolution and the biology of aging. In 1957 G.C. Williams predicted that higher adult death rates select for earlier senescence and shorter length of life, but preadult mortality does not matter to the evolution of senescence. This was subsequently interpreted as predicting that senescence should be caused by 'extrinsic' sources of mortality. This idea still motivates empirical studies, although formal, mathematical theory shows it is wrong. It has nonetheless prospered because it offers an intuitive explanation for patterns observed in nature. We review the flaws in Williams' model, explore alternative explanations for comparative patterns that are consistent with the evolutionary theory of senescence, and discuss how hypotheses based on it can be tested. We argue that focusing on how sources of mortality affect ages differently offers greater insight into evolutionary processes.

Long-term ecological observation affords a picture of the past that uniquely informs our understanding of present and future ecological communities and processes. Without a long-term perspective, our vision is prone to environmental myopia. Long-term experiments (LTEs) in particular can reveal the mechanisms that underlie change in communities and ecosystem functioning in a way that cannot be understood by long-term monitoring alone. Despite the urgent need to know more about how climate change will affect ecosystems and their functioning, the continued existence of LTEs is extremely precarious and we believe that dedicated funds are needed to support them. A new non-profit organization called the Ecological Continuity Trust seeks to provide a solution to this problem by establishing an endowment that will be specifically earmarked to sustain LTEs as a scientific tool for the benefit of future generations.

How large numbers of competing plant species manage to coexist is a major unresolved question in community ecology. The classical explanation, that each species occupies its own niche, seems at first unlikely because most plants require the same set of resources and acquire these in a limited number of ways. However, recent studies, although few in number and incomplete in many ways, do suggest that plants segregate along various environmental niche axes, including gradients of light, soil moisture and root depth, and that partitioning of soil nutrients occurs, possibly through the mediation of microbial symbionts, some of which are more species specific than was previously thought. Although it is unlikely that niche separation along environmental axes is the only mechanism of coexistence in any large community, the evidence now suggests that it plays a more significant role than has been previously appreciated. More research into the consequences of various known tradeoffs is likely to uncover further cases of niche separation facilitating coexistence.

Sustainable exploitation is widely advocated as a strategy for reconciling economic pressures upon natural habitats with nature conservation. Two recent papers examine different aspects of the sustainability of the nut harvest on wild populations of Brazil nut trees Bertholletia excelsa in Amazonia. Peres et al. find that many populations of the Brazil nut tree lack juvenile trees and are not regenerating. In a socioeconomic study, Escobal and Aldana find that nut-gathering provides insufficient income on its own to support nut-gatherers and that their other income-raising activities damage the forest. The existence of a market for rainforest products is, therefore, not sufficient on its own to prevent habitat destruction or the overexploitation of the resource and a more sophisticated approach to sustainability is required. Development of a market in ethically traded Brazil nuts might be one solution.

Species-rich plant communities appear to defy the competitive exclusion principle, showing relatively few obvious niche differences between coexisting species. Here we explore alternatives to the potentially endless search for new niche axes. Spatial aggregation in populations, non-transitive competition, episodes of density-independent mortality and various non-equilibrium theories allow trophically similar species to coexist for extended periods. In perennial plants or annuals with a seed pool, asynchrony between species in recruitment permits coexistence by the 'storage effect'. There is increasing evidence that species-specific herbivores and pathogens regulate populations of tropical trees to low levels at which competitive exclusion does not occur. The wide variety of alternatives to niche differentiation lead us to question whether plants need occupy different niches to coexist.

Correlations between habitat and seed size suggest that this character is adaptive. Mean seed size is a relatively invariant species characteristic and seed size has marked effects upon fitness. These observations have previously led to the conclusion that seed size is under stabilizing selection. This conclusion, originally based mainly on evidence from crop plants grown in controlled environments, is questioned here on the grounds that recent studies of wild plants show marked phenotypic plasticity and low heritability of seed size. If seed size is not readily altered by natural selection in the wild, then its effects on fitness are evidence that this character is a constraint on habitat distribution. Constancy of mean seed size may be due to developmental canalization to a size set by previous selection, rather than a continuing process of stabilizing selection.