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Clonal organisms persist at a range of population sex ratios, from equal numbers of males and females to single-sex systems. When intersexual competition is strong enough to drive one sex locally extinct, the maintenance of the sexes is facilitated by the semi-independent dynamics of populations within a metapopulation. These semi-independent dynamics are influenced by dispersal and recolonization rates, which are affected by the spatial arrangement of populations. To establish the quantitative relationship between spatially complex metapopulations and the maintenance of the sexes, we used a mathematical model of the liverwort Marchantia inflexa. This clonal organism is found in discrete patches on rocks and along the banks of streams, which form single-sex and two-sex metapopulations. In this system, asexual propagules mainly disperse short distances. Long-distance between-patch dispersal and recolonization mainly occurs via sexual propagules, which require both sexes to be present. Dispersal of these two types of propagules could interact with the spatial arrangement of populations to affect the maintenance of the sexes. With our mathematical model, we found that at intermediate distances between populations, metapopulations maintained both sexes, and the spatial arrangement of populations changed the threshold at which one sex was lost. On the other hand, when populations were close to one another, one sex was lost and the single-sex metapopulation persisted through dispersal of asexual propagules. When populations were far apart, one sex was lost, and the metapopulation either went extinct due to lack of recolonization by asexual propagules or persisted because clumped populations facilitated recolonization. These idealized spatial arrangements help clarify the effects of the spatial arrangement on the maintenance of the sexes and the persistence of metapopulations of clonal organisms, which can help explain geographic parthenogenesis and the distribution of asexual populations, the persistence of asexual species, and inform the conservation of clonal organisms.

In principle, the intensity of sexual conflict is best measured as a loss of fitness associated with the expression of conflict-related traits. But because the relevant traits may be difficult to manipulate and fitness difficult to assess, proxy variables linked to conflict intensity may provide important tools for empirical measurement. Here we identify two common types of sexual conflict—one within mating pairs over the less expensive male role, and one between mating pairs and intruders seeking to obtain fertilizations—and consider how they vary in intensity along gradients of population density and predation risk. To do this, we develop and analyze a model of mating dynamics in the chalk bass, an egg-trading simultaneous hermaphrodite that lives on Caribbean coral reefs. In this species, within-pair sexual conflict leads each female-role partner to provide in each mating episode only a subset (parcel) of its egg clutch to its mate for fertilization. Pair-intruder sexual conflict (i.e., sperm competition) increases the proportion of the gonad allocated to male function. In the model, more parceling and greater male allocation both resulted in lower fitness at the ESS, our measure of conflict intensity. Male allocation increased along the density gradient but decreased along the predation-risk gradient, reflecting shifts in intrusion frequency. Parcel number sharply increased and then decreased more gradually along a gradient of increasing local density, initially responding to increased availability of alternative mates across low densities and then to diminishing clutch size toward higher densities. Parcel number decreased with predation risk as each mating episode became more dangerous. Conflict intensities were usually greatest at intermediate positions along the two environmental gradients, and each conflict ameliorated the intensity of the other. Overall, parceling and sex allocation may be good though imperfect proxies for intensities of within-pair and pair-intruder sexual conflicts among chalk bass.

In theoretical and empirical studies of the evolution of cooperation, the tit-for-tat strategy (i.e. cooperate unless your partner did not cooperate in the previous interaction) is widely considered to be of central importance. Nevertheless, surprisingly little is known about the conditions in which tit-for-tat appears and disappears across generations in a population of interacting individuals. Here, we apply a newly developed classifier-system model (EvA) in addressing this issue when the key features of interactions are caricatured using the iterated prisoner's dilemma game. Our simple representation of behavioural strategies as algorithms composed of two interacting rules allowed us to determine conditions in which tit-for-tat can replace noncooperative strategies and vice versa. Using direct game-theoretic analysis and simulations with the EvA model, we determined that no strategy is evolutionarily stable, but larger population sizes and longer sequences of interactions between individuals can yield transient dominance by tit-for-tat. Genetic drift among behaviourally equivalent strategies is the key mechanism underlying this dominance. Our analysis suggests that tit-for-tat could be important in nature for cognitively simple organisms of limited memory capacity, in strongly kin-selected or group-selected populations, when interaction sequences between individuals are relatively short, in moderate-sized populations of widely interacting individuals and when defectors appear in the population with moderate frequency.

We generated a computer model to analyse the effects of ‘shadow’ competition for sit-and-wait predators, particularly antlion larvae. The model used a simple foraging assessment rule to determine the quality of an antlion's location, and antlions relocated randomly in their habitat when a location proved to be of low quality. Shadow competition, or competition for food caused when one sit-and-wait predator intercepts moving prey before a second sit-and-wait predator is encountered, was incorporated into the model by restricting antlions to a bounded arena, and having prey for the antlions enter from the arena periphery. Antlions responded to shadow competition by relocating their pits to peripheral areas of their habitat. This peripheral accumulation of pits was most pronounced when antlion densities were high, and when prey availabilities were intermediate. An experimental test with the antlionMyrmeleon immaculatus supported the importance of shadow competition as a cause of observed pit distributions. Only the treatment which incorporated shadowing had pit distributions near the periphery, while the pit distributions in the control treatments did not differ from randomly generated distributions. We conclude that shadowing can influence sit-and-wait predator distributions when the prey distributions and movement patterns generate the conditions necessary for shadowing. But when prey availability is unpredictable, making assessment of patches difficult, or when prey do not originate in the periphery of the habitat, other factors, such as temperature or moisture, could be more important.