Dong B & Gardner A (2025) Kin competition drives the evolution of earlier metamorphosis. Ecology and Evolution 15, e70806.

Metamorphosis, the discrete morphological change between postembryonic life stages, is widespread across the animal kingdom. The suggested advantages of metamorphosis have usually been framed in terms of population benefits, i.e., ecological explanations. In contrast, evolutionary explanations concern whether and how metamorphosis spreads through a population owing to individual-fitness benefits. However, how kin selection modulates evolution of metamorphosis remains to be investigated formally. Here we develop a mathematical model to investigate how kin selection shapes the optimal timing of metamorphosis from foraging, non-reproductive larva to reproductive adult, when larvae tend to cluster with their genetic relatives. We consider the full range of larval competition intensities—from no competition to full competition—and the full range of relatedness coefficients—from unrelated to clonality. We provide testable predictions as to how kin selection modulates the timing of metamorphosis, as well as a conceptual framework within which empirical observations may be understood.

 

Chokechaipaisarn C & Gardner A (2024) Density-dependent dispersal reduces conflict over the sex ratio. Biology Letters 20, 20240378.

Haplodiploids—in particular, wasps—are the workhorses of sex-allocation research. This is owing to their unusual system of sex determination, which provides a ready means of sex ratio adjustment. Notably, their sexually asymmetrical mode of genetic inheritance leads mothers and fathers to come into conflict over the sex ratio of their offspring. In the simplest outbreeding scenario, a mother is favoured to employ an even sex ratio while a father prefers that all his mate’s offspring are female. An important modulator of evolutionary conflict between mating partners is genetic relatedness, raising the possibility that this sex ratio conflict is reduced in low-dispersal settings with mating occurring between relatives. However, the impact of population viscosity on sex ratio conflict in haplodiploids remains unknown. Here, we develop and analyse a kin-selection model to investigate how the rate of dispersal modulates sex ratio conflict in a haplodiploid, viscous population setting. We find that population viscosity is associated with a reduction in the extent of sex ratio conflict—the effect being very weak under density-independent dispersal and much stronger under density-dependent dispersal.

Dong B, Paracchini S & Gardner A (in press) Kin selection as a modulator of human handedness: sex-specific, parental and parent-of-origin effects. Evolutionary Human Sciences.

Image: Wikimedia Commons

The frequency of left-handedness in humans is ∼10% worldwide and slightly higher in males than females. Twin and family studies estimate the heritability of human handedness at around 25%. The low but substantial frequency of left-handedness has been suggested to imply negative frequency-dependent selection, e.g. owing to a “surprise” advantage of left-handers in combat against opponents more used to fighting right-handers. Because such game-theoretic hypotheses involve social interaction, here, we perform an analysis of the evolution of handedness based on kin-selection, which is understood to play a major role in the evolution of social behaviour generally. We show that: (1) relatedness modulates the balance of right-handedness versus left-handedness, according to whether left-handedness is marginally selfish versus marginally altruistic; (2) sex differences in relatedness to social partners may drive sex differences in handedness; (3) differential relatedness of parents and offspring may generate parent-offspring conflict and sexual conflict leading to the evolution of maternal and paternal genetic effects in relation to handedness; and (4) differential relatedness of maternal-origin versus paternal-origin genes may generate intragenomic conflict leading to the evolution of parent-of-origin-specific gene effects—such as “genomic imprinting”—and associated maladaptation.

Hitchcock TJ & Gardner A (2023) Sexual antagonism in sequential hermaphrodites. Proceedings of the Royal Society of London Series B – Biological Sciences 290, 20232222.

Image: Wikimedia Commons

Females and males may have distinct phenotypic optima, but share essentially the same complement of genes, potentially leading to trade-offs between attaining high fitness through female versus male reproductive success. Such sexual antagonism may be particularly acute in hermaphrodites, where both reproductive strategies are housed within a single individual. While previous models have focused on simultaneous hermaphroditism, we lack theory for how sexual antagonism may play out under sequential hermaphroditism, which has the additional complexities of age-structure. Here, we develop a formal theory of sexual antagonism in sequential hermaphrodites. First, we construct a general theoretical overview of the problem, then consider different types of sexually antagonistic and life-history trade-offs, under different modes of genetic inheritance (autosomal or cytoplasmic), and different forms of sequential hermaphroditism (protogynous, protoandrous or bidirectional). Finally, we provide a concrete illustration of these general patterns by developing a two-stage two-sex model, which yields conditions for both invasion of sexually antagonistic alleles and maintenance of sexually antagonistic polymorphisms.

Gardner A (in press) A geometric approach to the evolution of altruism. Journal of Theoretical Biology https://doi.org/10.1016/j.jtbi.2023.111653

Fisher’s geometric model provides a powerful tool for making predictions about key properties of Darwinian adaptation. Here, I apply the geometric model to predict differences between the evolution of altruistic versus nonsocial phenotypes. I recover Kimura’s prediction that probability of fixation is greater for mutations of intermediate size, but I find that the effect size that maximises probability of fixation is relatively small in the context of altruism and relatively large in the context of nonsocial phenotypes, and that the overall probability of fixation is lower for altruism and is higher for nonsocial phenotypes. Accordingly, the first selective substitution is expected to be smaller, and to take longer, in the context of the evolution of altruism. These results strengthen the justification for employing streamlined social evolutionary methodologies that assume adaptations are underpinned by many genes of small effect.

Gardner A (2023) R. A. Fisher on J. A. Cobb’s The problem of the sex-ratio. Notes and Records http://doi.org/10.1098/rsnr.2023.0067.

The logic of the rarer-sex effect, concerning how natural selection acts to balance the sex ratio among newborns, was long supposed to have originated with Ronald Aylmer Fisher in his 1930 book The genetical theory of natural selection. However, the principle is now understood to have originated with John Austin Cobb in his 1914 paper ‘The problem of the sex-ratio’. Fisher did not provide a citation of Cobb’s sex-ratio paper, and it has been unclear whether he was aware of its existence. Here, I show that Fisher was indeed aware of Cobb’s paper in 1930, as revealed by him citing it elsewhere that same year. Fisher’s willingness to highlight Cobb’s sex-ratio work lends support to the view that his failure to mention it in his book reflects the less stringent citation standards of the time rather than an attempt to deceive readers as to the provenance of the rarer-sex effect.

Twyman KZ & Gardner A (2023) Kin selection of time travel: the social evolutionary causes and consequences of dormancy. Proceedings of the Royal Society of London Series B – Biological Sciences 290, 20231247.

 

A basic mechanism of kin selection is limited dispersal, whereby individuals remain close to their place of origin such that even indiscriminate social interaction tends to modify the fitness of genealogical kin. Accordingly, the causes and consequences of dispersal have received an enormous amount of attention in the social evolution literature. This work has focused on dispersal of individuals in space, yet similar logic should apply to dispersal of individuals in time (e.g. dormancy). We investigate how kin selection drives the evolution of dormancy and how dormancy modulates the evolution of altruism. We recover dormancy analogues of key results that have previously been given for dispersal, showing that: (1) kin selection favours dormancy as a means of relaxing competition between relatives; (2) when individuals may adjust their dormancy behaviour to local density, they are favoured to do so, resulting in greater dormancy in high-density neighbourhoods and a concomitant ‘constant non-dormant principle’; (3) when dormancy is constrained to be independent of density, there is no relationship between the rate of dormancy and the evolutionary potential for altruism; and (4) when dormancy is able to evolve in a density-dependent manner, a greater potential for altruism is expected in populations with lower dormancy.

Image: https://hotpot.ai/art-generator

Gardner A (in press) W. D. Hamilton and the golden sex ratio. Journal of Theoretical Biology. doi: 10.1016/j.jtbi.2023.111599

In his famous two-part paper, published in Journal of Theoretical Biology in 1964, W. D. Hamilton predicted that natural selection acting in male-haploid populations favours a ratio of males to females that is in accordance with the golden ratio. This prediction has found its way into the pages of one of the best-selling books of all time, Dan Brown’s 2003 novel The da Vinci Code, and is therefore in the running for the most widely known quantitative result in the history of evolutionary biology. Unfortunately, this golden-ratio result is wrong, and was later corrected by Hamilton, who showed that natural selection actually favours an unbiased sex ratio in this setting. But it has been unclear exactly how Hamilton arrived at the golden-ratio result in the first place. Here I show that the solution to this puzzle is found in unpublished work held in the British Library’s W. D. Hamilton collection. Specifically, in addition to employing a faulty method for calculating relatedness, Hamilton had also employed a faulty method for calculating reproductive value, considering only genetic contributions to the next generation rather than to the distant future. Repeating both mistakes recovers his erroneous golden-ratio result.

Image: https://hotpot.ai/art-generator

Chokechaipaisarn C & Gardner A (2023) A simulation test of the prediction that density-dependent dispersal promotes female-biased sex allocation in viscous populations. microPublication Biology doi: 10.17912/micropub.biology.000821

Stucky K & Gardner A (in press) Kin selection favours religious traditions: ancestor worship as a cultural descendant-leaving strategy Religion, Brain & Behavior doi: 10.1080/2153599X.2023.2215854

Recent years have seen renewed interest in the role of religious systems as drivers of the evolution of cooperation in human societies. One suggestion is that a cultural tradition of ancestor worship might have evolved as a “descendant-leaving strategy” of ancestors by encouraging increased altruism particularly between distant kin. Specifically, Coe and others have suggested a mechanism of cultural transmission exploiting social learning biases, whereby ancestors have been able to establish parental manipulation of kin recognition and perceived relatedness as a traditional behavior, leading to increased altruism among co-descendants and thereby maximizing the ancestor’s long-term inclusive fitness. Here, we develop a demographically explicit model in order to quantify the resulting increase in altruism and concomitant “ancestor-descendant conflict”, and to determine the evolutionary feasibility of religiously motivated cultural norms that promote altruism among co-descendants. Our analysis reveals that such norms could indeed drive an overall increase in altruism with potential for ancestor-descendant conflict, particularly in low-dispersal settings. Moreover, we find that natural selection can favor traditions encouraging increased altruism towards co-descendants under a range of conditions, with the inclusive-fitness costs of enacting an inappropriately high level of altruism being offset by inclusive-fitness benefits derived from the cultural tradition facilitating kin recognition.