Omar Tonsi Eldakar

The role of multilevel selection in the evolution of sexual conflict in the water strider aquarius remigis.

In evolution, exploitative strategies often create a paradox in which the most successful individual strategy “within” the group is also the most detrimental strategy “for” the group, potentially resulting in extinction. With regard to sexual conflict, the overexploitation of females by harmful males can yield similar consequences. Despite these evolutionary implications, little research has addressed why sexual conflict does not ultimately drive populations to extinction. One possibility is that groups experiencing less sexual conflict are more productive than groups with greater conflict. However, most studies of sexual conflict are conducted in a single isolated group, disregarding the potential for selection among groups. We observed Aquarius remigis water striders in a naturalistic multigroup pool in which individuals could freely disperse among groups. The free movement of individuals generated variation in aggression and sex‐ratio among groups, thereby increasing the importance of between‐group selection compared to within‐group selection. Females dispersed away from local aggression, creating more favorable mating environments for less‐aggressive males. Furthermore, the use of contextual analysis revealed that individual male aggression positively predicted fitness whereas aggression at the group level negatively predicted fitness, empirically demonstrating the conflict between levels of selection acting on mating aggression.

Aggressive Mating as a Tragedy of the Commons in the Water Strider Aquarius remigis

The tragedy of the commons usually refers to the overexploitation of resources such as food or water. Here, we show in a laboratory study that competition among males for females can also result in a tragedy of the commons’ situation. Male water striders (Aquarius remigis) vary widely in their aggressiveness toward pursuing females. The most aggressive males prevent females from feeding and cause them to leave the water surface, where they are unavailable to all males. Groups of nonaggressive males are collectively three times more fit than groups of hyperaggressive males, but hyperaggressive males are more successful than nonaggressive males within mixed groups. This is the classic tragedy of the commons’ situation, and it is likely to occur in many species that exhibit sexual conflict. We have also shown that individual differences in male aggression are stable across time and are not influenced by food or light regime, although all individuals become nonaggressive in the presence of fish predators.

The thermoregulatory theory of yawning: what we know from over 5 years of research

Over the past 5 years numerous reports have confirmed and replicated the specific brain cooling and thermal window predictions derived from the thermoregulatory theory of yawning, and no study has found evidence contrary to these findings. Here we review the comparative research supporting this model of yawning among homeotherms, while highlighting a recent report showing how the expression of contagious yawning in humans is altered by seasonal climate variation. The fact that yawning is constrained to a thermal window of ambient temperature provides unique and compelling support in favor of this theory. Heretofore, no existing alternative hypothesis of yawning can explain these results, which have important implications for understanding the potential functional role of this behavior, both physiologically and socially, in humans and other animals. In discussion we stress the broader applications of this work in clinical settings, and counter the various criticisms of this theory.

EIGHT CRITICISMS NOT TO MAKE ABOUT GROUP SELECTION

Group selection, which was once widely rejected as a significant evolutionary force, is now accepted by all who seriously study the subject. There is still widespread confusion about group selection, however, not only among students and the general public, but among professional evolutionists who do not directly study the subject. We list eight criticisms that are frequently invoked against group selection, which can be permanently laid to rest based upon current knowledge. Experts will always find something to critique about group selection, as for any important subject, but these eight criticisms are not among them. Laying them to rest will enable authors to openly use the term group selection without being handicapped during the review process.

Contagious Yawning and Seasonal Climate Variation

Recent evidence suggests that yawning is a thermoregulatory behavior. To explore this possibility further, the frequency of contagious yawning in humans was measured while outdoors in a desert climate in the United States during two distinct temperature ranges and seasons (winter: 22°C; early summer: 37°C). As predicted, the proportion of pedestrians who yawned in response to seeing pictures of people yawning differed significantly between the two conditions (winter: 45%; summer: 24%). Across conditions yawning occurred at lower ambient temperatures, and the tendency to yawn during each season was associated with the length of time spent outside prior to being tested. Participants were more likely to yawn in the milder climate after spending long periods of time outside, while prolonged exposure to ambient temperatures at or above body temperature was associated with reduced yawning. This is the first report to show that the incidence of yawning in humans is associated with seasonal climate variation, further demonstrating that yawn-induced contagion effects can be mediated by factors unrelated to individual social characteristics or cognitive development.

A thermal window for yawning in humans: Yawning as a brain cooling mechanism

The thermoregulatory theory of yawning posits that yawns function to cool the brain in part due to counter-current heat exchange with the deep inhalation of ambient air. Consequently, yawning should be constrained to an optimal thermal zone or range of temperature, i.e., a thermal window, in which we should expect a lower frequency at extreme temperatures. Previous research shows that yawn frequency diminishes as ambient temperatures rise and approach body temperature, but a lower bound to the thermal window has not been demonstrated. To test this, a total of 120 pedestrians were sampled for susceptibly to self-reported yawn contagion during distinct temperature ranges and seasons (winter: 1.4°C, n=60; summer: 19.4°C, n=60). As predicted, the proportion of pedestrians reporting yawning was significantly lower during winter than in summer (18.3% vs. 41.7%), with temperature being the only significant predictor of these differences across seasons. The underlying mechanism for yawning in humans, both spontaneous and contagious, appears to be involved in brain thermoregulation.

Allelopathy as an emergent, exploitable public good in the bloom-forming microalga Prymnesium parvum.

Many microbes cooperatively secrete extracellular products that favorably modify their environment. Consistent with social evolution theory, structured habitats play a role in maintaining these traits in microbial model systems, by localizing the benefits and separating strains that invest in these products from ‘cheater’ strains that benefit without paying the cost. It is thus surprising that many unicellular, well‐mixed microalgal populations invest in extracellular toxins that confer ecological benefits upon the entire population, for example, by eliminating nutrient competitors (allelopathy). Here we test the hypotheses that microalgal exotoxins are (1) exploitable public goods that benefit all cells, regardless of investment, or (2) nonexploitable private goods involved in cell‐level functions. We test these hypotheses with high‐toxicity (TOX+) and low‐toxicity (TOX−) strains of the damaging, mixotrophic microalga Prymnesium parvum and two common competitors: green algae and diatoms. TOX+ actually benefits from dense populations of competing green algae, which can also be prey for P. parvum, yielding a relative fitness advantage over coexisting TOX−. However, with nonprey competitors (diatoms), TOX− increases in frequency over TOX+, despite benefiting from the exclusion of diatoms by TOX+. An evolutionary unstable, ecologically devastating public good may emerge from traits selected at lower levels expressed in novel environments.

The group-level consequences of sexual conflict in multigroup populations.

In typical sexual conflict scenarios, males best equipped to exploit females are favored locally over more prudent males, despite reducing female fitness. However, local advantage is not the only relevant form of selection. In multigroup populations, groups with less sexual conflict will contribute more offspring to the next generation than higher conflict groups, countering the local advantage of harmful males. Here, we varied male aggression within-and between-groups in a laboratory population of water striders and measured resulting differences in local population growth over a period of three weeks. The overall pool fitness (i.e., adults produced) of less aggressive pools exceeded that of high aggression pools by a factor of three, with the high aggression pools essentially experiencing no population growth over the course of the study. When comparing the fitness of individuals across groups, aggression appeared to be under stabilizing selection in the multigroup population. The use of contextual analysis revealed that overall stabilizing selection was a product of selection favoring aggression within groups, but selected against it at the group-level. Therefore, this report provides further evidence to show that what evolves in the total population is not merely an extension of within-group dynamics.

Population Structure Influences Sexual Conflict in Wild Populations of Water Striders.

In sexual conflict, aggressive males frequently diminish the long-term reproductive success of females in efforts to gain a short-term advantage over rival males. This short-term advantage can selectively favour high-exploitation males. However, just as the over-exploitation of resources can lead to local extinction, the over-exploitation of females in the form of harassment by aggressive males can yield similar consequences resulting in reduced female fecundity, increased female mortality and overall decline in mating activity. This outcome may often be prevented by selection acting at multiple levels of biological organization. Directional selection favouring aggressive exploitation within groups can be balanced by directional selection amongst groups opposing exploitation. Such between-group selection has recently been demonstrated in laboratory studies of water striders, where the conditional dispersal of individuals increased variation amongst groups and influenced the balance of selection toward reduced male aggression. This multilevel selection (MLS) framework also provides predictive value when investigating natural populations differing in their relative strength of selection within versus among groups. For water striders, the consequences of local exploitation cause fitness differences between groups, favouring less aggressive males. Inconsistently flowing ephemeral streams consist of isolated pools that prevent aggressive male water striders from escaping the consequences of local exploitation. We, therefore, predicted that inconsistently flowing ephemeral streams would favour the evolution of less aggressive males than would perennial streams, which allow aggressive males to move more freely and to escape the group-level costs of their aggression. Comparing two neighbouring streams during the mating season, we found that males dispersed naturally between pools at much higher rates in the perennial stream than in the ephemeral stream. As predicted, we found that males from the perennial stream were significantly more aggressive than those from the ephemeral stream. We also found that dispersers were significantly more aggressive than non-dispersers within each stream. These field results illustrate the relevance of the MLS framework in our understanding of the evolution of sexual conflict.

High mating frequency and variation with lineage ratio in dependent-lineage harvester ants

Explaining the evolution of multiple mating is a challenge because of the associated costs. For social insects, mating frequency may have fitness consequences due to effects on social interactions or genetic diversity within colonies. Here, we investigated the evolution of mating frequency in a social insect species with a unique genetic system that requires multiple mating. In certain populations of Pogonomyrmex harvester ants, there are two interbreeding yet genetically distinct mitochondrial lineages. Queens must mate with males of the opposite lineage to produce workers and with males of the same lineage to produce reproductive females. We expected queens of the dependent-lineage system to exhibit high mating frequencies relative to other social insects. Furthermore, we expected queens from populations of highly asymmetric lineage ratios to exhibit even higher mating frequencies, to adequately sample the population and successfully mate with males of the less common lineage. To test these predictions, we estimated the mating frequency of 16 P. barbatus queens, and compared these mating frequencies between two populations, one with relatively equal lineage ratio (60:40) and a second with a highly asymmetrical lineage ratio (96:4). Overall, observed mating frequency exceeded 10, which is high in comparison to other social insects, and our estimates of effective mating frequency were among the highest of Pogonomyrmex species. Mating frequency at the site with the asymmetrical lineage ratio was also significantly higher than the site with the more even ratio. Our results suggest that obligate multiple mating as well as lineage ratio contribute to the evolution of high mating frequency in dependent-lineage populations.