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Two of John et al.'s examples of proxy failures in ecological situations are not failures: Runaway sexual selection and marsupial neonate competition. Instead, more appropriate ecological examples may be paternal genetic kin recognition and warning coloration. These differ in proxy effectiveness and failure in ways that illustrate the importance of "costs" in the evolution of ecological proxy traits.

Eusocial Hymenoptera are often characterized by having facultatively or obligately sterile worker castes. However, findings across an increasing number of species are that some workers are non-natal-they have 'drifted' away from where they were born and raised. Moreover, drifters are often indistinguishable from natal workers in the work and benefits provided to joined groups. This seems an evolutionary paradox of providing benefits to potentially unrelated individuals over close kin. Rather than being mistakes, drifting is proposed to be adaptive if joiners either gain inclusive fitness by preferentially moving to other kin groups or through generalized reciprocity in which exchanging workers across groups raises group-level genetic diversity and creates social heterosis. It is unclear, however, if reciprocity is unlikely because of a susceptibility to cheating. In resolving this question, a series of evolutionary simulations show: (1) Reciprocity can persist under a range of genetic assumptions and scenarios of cheating, (2) cheating almost always evolves, but can be expressed in a variety of ways that are not always predictable, (3) the inclusive fitness hypothesis is equally or more susceptible to cheating. Moreover, existing data in Hymenoptera (although not extensive) are more consistent with generalized reciprocity. This supports a hypothesis that drifting, as a phenomenon, may more often reflect maximization of group and parental fitness rather than fitness gains for the individual drifters.

In viral infections and cancer tumours, negative health outcomes often correlate with increasing genetic diversity. Possible evolutionary processes for such relationships include mutant lineages escaping host control or diversity, per se, creating too many immune system targets. Another possibility is social heterosis where mutations and replicative errors create clonal lineages varying in intrinsic capability for successful dispersal; improved environmental buffering; resource extraction or effective defence against immune systems. Rather than these capabilities existing in one genome, social heterosis proposes complementary synergies occur across lineages in close proximity. Diverse groups overcome host defences as interacting 'social genomes' with group genetic tool kits exceeding limited individual plasticity. To assess the possibility of social heterosis in viral infections and cancer progression, we conducted extensive literature searches for examples consistent with general and specific predictions from the social heterosis hypothesis. Numerous studies found supportive patterns in cancers across multiple tissues and in several families of RNA viruses. In viruses, social heterosis mechanisms probably result from long coevolutionary histories of competition between pathogen and host. Conversely, in cancers, social heterosis is a by-product of recent mutations. Investigating how social genomes arise and function in viral quasi-species swarms and cancer tumours may lead to new therapeutic approaches.

A small number of extraordinary “Major Evolutionary Transitions” (METs) have attracted attention among biologists. They comprise novel forms of individuality and information, and are defined in relation to organismal complexity, irrespective of broader ecosystem-level effects. This divorce between evolutionary and ecological consequences qualifies unicellular eukaryotes, for example, as a MET although they alone failed to significantly alter ecosystems. Additionally, this definition excludes revolutionary innovations not fitting into either MET type (e.g., photosynthesis). We recombine evolution with ecology to explore how and why entire ecosystems were newly created or radically altered – as Major System Transitions (MSTs). In doing so, we highlight important morphological adaptations that spread through populations because of their immediate, direct-fitness advantages for individuals. These are Major Competitive Transitions, or MCTs. We argue that often multiple METs and MCTs must be present to produce MSTs. For example, sexually-reproducing, multicellular eukaryotes (METs) with anisogamy and exoskeletons (MCTs) significantly altered ecosystems during the Cambrian. Therefore, we introduce the concepts of Facilitating Evolutionary Transitions (FETs) and Catalysts as key events or agents that are insufficient themselves to set a MST into motion, but are essential parts of synergies that do. We further elucidate the role of information in MSTs as transitions across five levels: (I) Encoded; (II) Epigenomic; (III) Learned; (IV) Inscribed; and (V) Dark Information. The latter is ‘authored’ by abiotic entities rather than biological organisms. Level IV has arguably allowed humans to produce a MST, and V perhaps makes us a FET for a future transition that melds biotic and abiotic life into one entity. Understanding the interactive processes involved in past major transitions will illuminate both current events and the surprising possibilities that abiotically-created information may produce.

A multitude of factors may determine reproductive skew among cooperative breeders. One explanation, derived from inclusive fitness theory, is that groups can partition reproduction such that subordinates do at least as well as noncooperative solitary individuals. The majority of recent data, however, fails to support this prediction; possibly because inclusive fitness models cannot easily incorporate multiple factors simultaneously to predict skew. Notable omissions are antagonistic selection (across generations, genes will be in both dominant and subordinate bodies), constraints on the number of sites suitable for successful reproduction, choice in which group an individual might join, and within-group control or suppression of competition. All of these factors and more are explored through agent-based evolutionary simulations. The results suggest the primary drivers for the initial evolution of cooperative breeding may be a combination of limited suitable sites, choice across those sites, and parental manipulation of offspring into helping roles. Antagonistic selection may be important when subordinates are more frequent than dominants. Kinship matters, but its main effect may be in offspring being available for manipulation while unrelated individuals are not. The greater flexibility of evolutionary simulations allows the incorporation of species-specific life histories and ecological constraints to better predict sociobiology.

While strict vertical transmission insures the durability of intracellular symbioses, phylogenetic incongruences between hosts and endosymbionts suggest horizontal transmission must also occur. These horizontal acquisitions can have important implications for the biology of the host. Wolbachia is one of the most ecologically successful prokaryotes in arthropods, infecting an estimated 50-70% of all insect species. Much of this success is likely due to the fact that, in arthropods, Wolbachia is notorious for manipulating host reproduction to favor transmission through the female germline. However, its natural potential for horizontal transmission remains poorly understood. Here we evaluate the fundamental prerequisites for successful horizontal transfer, including necessary environmental conditions, genetic potential of bacterial strains, and means of mediating transfers. Furthermore, we revisit the relatedness of Wolbachia strains infecting the Panamanian leaf-cutting ant, Acromyrmex echinatior, and its inquiline social parasite, Acromyrmex insinuator, and compare our results to a study published more than 15 years ago by Van Borm et al. (2003). The results of this pilot study prompt us to reevaluate previous notions that obligate social parasitism reliably facilitates horizontal transfer and suggest that not all Wolbachia strains associated with ants have the same genetic potential for horizontal transmission.

Parents often face an investment trade-off between either producing many small or fewer large offspring. When environments vary predictably, the fittest parental solution matches available resources by varying only number of offspring and never optimal individual size. However when mismatches occur often between parental expectations and true resource levels, dynamic models like multifaceted parental investment (MFPI) and parental optimism (PO) both predict offspring size can vary significantly. MFPI is a "realist" strategy: parents assume future environments of average richness. When resources exceed expectations and it is too late to add more offspring, the best-case solution increases investment per individual. Brood size distributions therefore track the degree of mismatch from right-skewed around an optimal size (slight underestimation of resources) to left-skewed around a maximal size (gross underestimation). Conversely, PO is an "optimist" strategy: parents assume maximally good resource futures and match numbers to that situation. Normal or lean years do not affect "core" brood as costs primarily fall on excess "marginal" siblings who die or experience stunted growth (producing left-skewed distributions). Investment patterns supportive of both MFPI and PO models have been observed in nature, but studies that directly manipulate food resources to test predictions are lacking. Ant colonies produce many offspring per reproductive cycle and are amenable to experimental manipulation in ways that can differentiate between MFPI and PO investment strategies. Colonies in a natural population of a harvester ant (Pogonomyrmex salinus) were protein-supplemented over 2 years, and mature sexual offspring were collected annually prior to their nuptial flight. Several results support either MFPI or PO in terms of patterns in offspring size distributions and how protein differentially affected male and female production. Unpredicted by either model, however, is that supplementation affected distributions more strongly across years than within (e.g., small females are significantly rarer in the year after colonies receive protein). Parental investment strategies in P. salinus vary dynamically across years and conditions. Finding that past conditions can more strongly affect reproductive decisions than current ones, however, is not addressed by models of parental investment.

Game theory is used in biology to understand why otherwise rational individuals make nonintuitive decisions regarding cooperation and competition. Recently, biology teachers engaged their students in game theory curricula by presenting them with a real-world game theory challenge: the opportunity to cheat on a game theory exam. Here we present a guide for other teachers to employ this provocative and educational classroom exercise, and discuss the results of the Cheating to Learn exercise in a biology class.

One of the hallmarks of eusociality is that workers forego their own reproduction to assist their mother in raising siblings. This seemingly altruistic behaviour may benefit workers if gains in indirect fitness from rearing siblings outweigh the loss of direct fitness. If worker presence is advantageous to mothers, however, eusociality may evolve without net benefits to workers. Indirect fitness benefits are often cited as evidence for the importance of inclusive fitness in eusociality, but have rarely been measured in natural populations. We compared inclusive fitness of alternative social strategies in the tropical sweat bee, Megalopta genalis, for which eusociality is optional. Our results show that workers have significantly lower inclusive fitness than females that found their own nests. In mathematical simulations based on M. genalis field data, eusociality cannot evolve with reduced intra-nest relatedness. The simulated distribution of alternative social strategies matched observed distributions of M. genalis social strategies when helping behaviour was simulated as the result of maternal manipulation, but not as worker altruism. Thus, eusociality in M. genalis is best explained through kin selection, but the underlying mechanism is likely maternal manipulation.

The role of the ant colony largely consists of non-reproductive tasks, such as foraging, tending brood, and defense. However, workers are vitally linked to reproduction through their provisioning of sexual offspring, which are produced annually to mate and initiate new colonies. Gynes (future queens) have size-associated variation in colony founding strategy (claustrality), with each strategy requiring different energetic investments from their natal colony. We compared the per capita production cost required for semi-claustral, facultative, and claustral gynes across four species of Pogonomyrmex harvester ants. We found that the claustral founding strategy is markedly expensive, costing approximately 70% more energy than that of the semi-claustral strategy. Relative to males, claustral gynes also had the largest differential investment and smallest size variation. We applied these investment costs to a model by Brown and Bonhoeffer (2003) that predicts founding strategy based on investment cost and foraging survivorship. The model predicts that non-claustral foundresses must survive the foraging period with a probability of 30-36% in order for a foraging strategy to be selectively favored. These results highlight the importance of incorporating resource investment at the colony level when investigating the evolution of colony founding strategies in ants.

Smaldino proposes emergent properties of human groups, arising when individuals display both differentiation and organization, constitute a novel unit of cultural selection not addressed by current evolutionary theory. We propose existing theoretical frameworks for maintenance of genetic diversity - social heterosis and social genomes - can similarly explain the appearance and maintenance of human cultural diversity (i.e., group-level traits) and collaborative interdependence.

Many social Hymenoptera species have morphologically sterile worker castes. It is proposed that the evolutionary routes to this obligate sterility must pass through a 'monogamy window', because inclusive fitness favours individuals retaining their reproductive totipotency unless they can rear full siblings. Simulated evolution of sterility, however, finds that 'point of view' is critically important. Monogamy is facilitating if sterility is expressed altruistically (i.e. workers defer reproduction to queens), but if sterility results from manipulation by mothers or siblings, monogamy may have no effect or lessen the likelihood of sterility. Overall, the model and data from facultatively eusocial bees suggest that eusociality and sterility are more likely to originate through manipulation than by altruism, casting doubt on a mandatory role for monogamy. Simple kin selection paradigms, such as Hamilton's rule, can also fail to account for significant evolutionary dynamics created by factors, such as population structure, group-level effects or non-random mating patterns. The easy remedy is to always validate apparently insightful predictions from Hamiltonian equations with life-history appropriate genetic models.

The reproductive (queen) and nonreproductive (worker) castes of eusocial insect colonies are a classic example of insect polyphenism. A complementary polyphenism may also exist entirely among females in the reproductive caste. Although less studied, reproductive females may vary in behavior based on size-associated attributes leading to the production of daughter workers. We studied a bee with flexible social behavior, Megalopta genalis, to better understand the potential of this polyphenism to shape the social organization of bee colonies and, by extension, its role in the evolution of eusociality. Our experimental design reduced variation among nest foundresses in life history variables that could influence reproductive decisions, such as nesting quality and early adulthood experience. Within our study population, approximately one third of M. genalis nests were eusocial and the remaining nests never produced workers. Though they do not differ in survival, nest-founding females who do not attempt to produce workers (which we refer to as the solitary phenotype) are significantly smaller and become reproductive later than females who attempt to recruit workers (the social phenotype). Females with the social phenotype are more likely to produce additional broods but at a cost of having some of their first offspring become nonreproductive workers. The likelihood of eusocial organization varies with body size across females of the social phenotype. Thus, fitness consequences associated with size-based plasticity in foundress behavior has colony level effects on eusociality. The potential for size-based polyphenisms among reproductive females may be an important factor to consider in the evolutionary origins of eusociality. © 2012 Springer-Verlag Berlin Heidelberg.

Social castes of eusocial insects may have arisen through an evolutionary modification of an ancestral reproductive ground plan, such that some adults emerge from development physiologically primed to specialize on reproduction (queens) and others on maternal care expressed as allo-maternal behaviour (workers). This hypothesis predicts that variation in reproductive physiology should emerge from ontogeny and underlie division of labour. To test these predictions, we identified physiological links to division of labour in a facultatively eusocial sweat bee, Megalopta genalis. Queens are larger, have larger ovaries and have higher vitellogenin titres than workers. We then compared queens and workers with their solitary counterparts-solitary reproductive females and dispersing nest foundresses-to investigate physiological variation as a factor in caste evolution. Within dyads, body size and ovary development were the best predictors of behavioural class. Queens and dispersers are larger, with larger ovaries than their solitary counterparts. Finally, we raised bees in social isolation to investigate the influence of ontogeny on physiological variation. Body size and ovary development among isolated females were highly variable, and linked to differences in vitellogenin titres. As these are key physiological predictors of social caste, our results provide evidence for developmental caste-biasing in a facultatively eusocial bee.

Infestations of buildings by Argentine ants, Linepithema humile (Mayr), were monitored on the campus of the University of California, Los Angeles. Foraging ant activity peaked during the hotter months of the year. The mean monthly maximum temperature, but not rainfall, positively correlated with indoor infestation frequency. Neither garden size nor the predominant groundcover vegetation correlated with the number of foraging ants at baits within gardens. Although the number of foraging ants outside a building varied over 40-fold, ant density in gardens did not predict the likelihood of infestation within the building. Also, the type of vegetative groundcover employed did not predict infestation frequency. There was, however, a significant negative relationship between the size of the garden outside of a building and the number of infestations. Given the large foraging area of L. humile workers, buildings next to small gardens may be infested simply because they lie within the "normal" foraging area of a colony. The best predictor of which rooms were infested within buildings was the presence of a water source. Thus providing water for ant colonies outside and away from buildings may be one method of integrated pest management to reduce the proclivity of ants to infest structures. Copyright © 2012 Smadar Gilboa et al.

Social Hymenoptera have played a leading role in development and testing of kin selection theory. Inclusive fitness models, following from Hamilton's rule, successfully predict major life history characteristics, such as biased sex investment ratios and conflict over parentage of male offspring. However, kin selection models poorly predict patterns of caste-biasing nepotism and reproductive skew within groups unless kin recognition constraints or group-level selection is also invoked. These successes and failures mirror the underlying kin recognition mechanisms. With reliable environmental cues, such as the sex of offspring or the origin of male eggs, predictions are supported. When only genetic recognition cues are potentially available, predictions are not supported. Mathematical simulations demonstrate that these differing mechanisms for determining kinship produce very different patterns of behavior. Decisions based on environmental cues for relatedness result in a robust mixture of cooperation and noncooperation depending on whether or not Hamilton's rule is met. In contrast, cooperation evolves under a wider range of conditions and to higher frequencies with genetic kin recognition as shared greenbeard traits. This "excess of niceness" matches the existing patterns in caste bias and reproductive skew; individuals often help others at an apparent cost to their inclusive fitness. The results further imply a potential for greenbeard-type kin recognition to create arbitrary runaway social selection for shared genetic traits. Suggestive examples in social evolution may be alloparental care and unicoloniality in ants. Differences in kin recognition mechanisms also can have consequences for maintenance of advantageous genetic diversity within populations.

Developmental maternal effects are a potentially important source of phenotypic variation, but they can be difficult to distinguish from other environmental factors. This is an important distinction within the context of social evolution, because if variation in offspring helping behavior is due to maternal manipulation, social selection may act on maternal phenotypes, as well as those of offspring. Factors correlated with social castes have been linked to variation in developmental nutrition, which might provide opportunity for females to manipulate the social behavior of their offspring. Megalopta genalis is a mass-provisioning facultatively eusocial sweat bee for which production of males and females in social and solitary nests is concurrent and asynchronous. Female offspring may become either gynes (reproductive dispersers) or workers (non-reproductive helpers). We predicted that if maternal manipulation plays a role in M. genalis caste determination, investment in daughters should vary more than for sons. The mass and protein content of pollen stores provided to female offspring varied significantly more than those of males, but volume and sugar content did not. Sugar content varied more among female eggs in social nests than in solitary nests. Provisions were larger, with higher nutrient content, for female eggs and in social nests. Adult females and males show different patterns of allometry, and their investment ratio ranged from 1.23 to 1.69. Adult body weight varied more for females than males, possibly reflecting increased variation in maternal investment in female offspring. These differences are consistent with a role for maternal manipulation in the social plasticity observed in M. genalis.

A major evolutionary question is how reproductive sharing arises in cooperatively breeding species despite the inherent reproductive conflicts in social groups. Reproductive skew theory offers one potential solution: each group member gains or is allotted inclusive fitness equal to or exceeding their expectation from reproducing on their own. Unfortunately, a multitude of skew models with conflicting predictions has led to confusion in both testing and evaluating skew theory. The confusion arises partly because one set of models (the 'transactional' type) answer the ultimate evolutionary question of what ranges of reproductive skew can yield fitness-enhancing solutions for all group members. The second set of models ('compromise') give an evolutionarily proximate, game-theoretic evolutionarily stable state (ESS) solution that determines reproductive shares based on relative competitive abilities. However, several predictions arising from compromise models require a linear payoff to increased competition and do not hold with non-linear payoffs. Given that for most species it may be very difficult or impossible to determine the true relationship between effort devoted to competition and reproductive share gained, compromise models are much less predictive than previously appreciated. Almost all skew models make one quantitative prediction (e.g. realized skew must fall within ranges predicted by transactional models), and two qualitative predictions (e.g. variation in relatedness or competitive ability across groups affects skew). A thorough review of the data finds that these three predictions are relatively rarely supported. As a general rule, therefore, the evolution of cooperative breeding appears not to be dependent on the ability of group members to monitor relatedness or competitive ability in order to adjust their behaviour dynamically to gain reproductive share. Although reproductive skew theory fails to predict within-group dynamics consistently, it does better at predicting quantitative differences in skew across populations or species. This suggests that kin selection can play a significant role in the evolution of sociality. To advance our understanding of reproductive skew will require focusing on a broader array of factors, such as the frequency of mistaken identity, delayed fitness payoffs, and selection pressures arising from across-group competition. We furthermore suggest a novel approach to investigate the sharing of reproduction that focuses on the underlying genetics of skew. A quantitative genetics approach allows the partitioning of variance in reproductive share itself or that of traits closely associated with skew into genetic and non-genetic sources. Thus, we can determine the heritability of reproductive share and infer whether it actually is the focus of natural selection. We view the 'animal model' as the most promising empirical method where the genetics of reproductive share can be directly analyzed in wild populations. In the quest to assess whether skew theory can provide a framework for understanding the evolution of sociality, quantitative genetics will be a central tool in future research.

Abstract Background Phylogenetic analyses strongly associate nonsocial ancestors of cooperatively-breeding or eusocial species with monogamy. Because monogamy creates high-relatedness family groups, kin selection has been concluded to drive the evolution of cooperative breeding (i.e., the monogamy hypothesis). Although kin selection is criticized as inappropriate for modeling and predicting the evolution of cooperation, there are no examples where specific inclusive fitness-based predictions are intrinsically wrong. The monogamy hypothesis may be the first case of such a flawed calculation. Results A simulation model mutated helping alleles into non-cooperative populations where females mated either once or multiply. Although multiple mating produces sibling broods of lower relatedness, it also increases the likelihood that one offspring will adopt a helper role. Examining this tradeoff showed that under a wide range of conditions polygamy, rather than monogamy, allowed helping to spread more rapidly through populations. Further simulations with mating strategies as heritable traits confirmed that multiple-mating is selectively advantageous. Although cooperation evolves similarly regardless of whether dependent young are close or more distant kin, it does not evolve if they are unrelated. Conclusions The solitary ancestral species to cooperative breeders may have been predominantly monogamous, but it cannot be concluded that monogamy is a predisposing state for the evolution of helping behavior. Monogamy may simply be coincidental to other more important life history characteristics such as nest defense or sequential provisioning of offspring. The differing predictive outcome from a gene-based model also supports arguments that inclusive fitness formulations poorly model some evolutionary questions. Nevertheless, cooperation only evolves when benefits are provided for kin: helping alleles did not increase in frequency in the absence of potential gains in indirect fitness. The key question, therefore, is not whether kin selection occurs, but how best to elucidate the differing evolutionary advantages of genetic relatedness versus genetic diversity.

Models of reproductive skew assume reproductive shares are either conceded, competed over, or both. Previous mathematical evaluations found that simultaneous concessions and contests are evolutionarily unstable. Recently, Shen and Reeve (2010) challenged these conclusions and developed a series of sub-models they argued to be a unified approach to reproductive skew: the general bordered tug-of-war (BTOW). However, BTOW fails as a general model for two reasons: (1) the BTOW strategy cannot invade populations where individuals either only compete for or only concede reproductive shares and (2) contrary to Shen and Reeve's assertion, BTOW populations are easily invaded by strategies with fewer or no concessions, but competing at lower levels. The failure of BTOW as a general model has major implications for interpreting experiments on reproductive skew. A large number of studies have measured the effects of genetic relatedness and competitive ability on reproductive skew, with a great majority finding no significant correlation between variation in within-group relatedness or competitive ability and across-group differences in skew. No model of reproductive skew except one variant of the BTOW predicts such results. With the rejection of BTOW as a valid general model, it is clear that these results are contradictory to reproductive skew theory rather than supportive of it.

Ants exhibit a size-associated colony founding trait that is characterized by the degree to which foundresses rely on internal reserves to raise their first brood of workers (claustrality). The reliance on stored reserves is positively correlated with degree of claustrality (claustral > facultative > semi-claustral) and is variable across species of Pogonomyrmex harvester ants. Three species of harvester ant foundresses that differ in degree of claustrality were observed initiating nests under laboratory conditions over 2 years. P. rugosus is fully claustral, P. salinus is facultative, and P. californicus is semi-claustral. Across species, degree of claustrality was positively associated with mean digging rate and nest depth over the first 3 days of nest initiation, total nest depth, and degree of nest closure. Branching and abundance of peripheral nodes were higher in semi-claustral and facultative nests than in claustral nests. The facultative species dug for the longest time and achieved the greatest tunnel length. Within each species, there were trends associating mass with digging rate, but these were not consistent in all species. There were no intraspecific trends of mass with nest depth. Also within species, a foundress’s mass did not affect her tendency to open or close her nest. These results reveal degree of claustrality is correlated across species with several nest initiation characteristics that together may represent different colony founding syndromes.

The velvety tree ant (Liometopum occidental Emery) is a dominant ant species found in Californian pine and oak woodlands. We performed a series of experiments to examine the degree of intraspecific aggression between nests. Within a site locality, ants collected from distances over a kilometer apart showed no aggression. However, ants from separate sites (more than 150 km apart) were strongly aggressive towards each other. Observation of food collection from baits further showed that L. occidentale trails can exceed over 70m. These combined observations suggest that L. occidentale has a natural life history that exhibits unicoloniality with perhaps the formation of large, habitat-dominating supercolonies. We also report an observation of a mating flight of L. occidentale that suggests females do disperse from their natal nests.

Cooperative breeding often results in unequal reproduction between dominant and subordinate group members. Transactional skew models attempt to predict how unequal reproduction can be before the groups themselves become unstable. A number of variants of transactional models have been developed, with a key difference being whether reproduction is controlled by one party or contested by all. It is shown here that ESS solutions for all situations of contested control over reproduction are given by the original tug-of-war model (TOW). Several interesting results follow. First, TOW can escalate enough to destabilize some types of groups. Particularly vulnerable are those that have low relatedness and gain little from cooperative breeding relative to solitary reproduction. Second, TOW can drastically reduce group productivity and especially the inclusive fitness of dominant individuals. Third, these results contrast strongly with those from variants of TOW models that include concessions to maintain group stability. Such models are shown to be special cases of the general and simpler TOW framework, and to have assumptions that may be biologically suspect. Finally, the overall analysis suggests that there is no mechanism within existing TOW framework that will prevent a costly struggle for reproductive control. Because social species rarely exhibit the high levels of aggression predicted by TOW models, alternative evolutionary mechanisms are considered that can limit conflict and produce more mutually beneficial outcomes. The further development of alternative models to predict patterns of reproductive skew are highly recommended.

In many eusocial Hymenoptera, workers prevent each other from producing male offspring by destroying worker-laid eggs. Kin selection theory predicts that such 'worker policing' behaviour can evolve by increasing the average relatedness between workers and their male brood. Alternatively, if worker-laid eggs are of low relative viability, their replacement would increase the developmental reliability of the brood. Less colony investment in terms of time and resources would be lost on poor males. This gain is independent of the relatedness of the males. Unfortunately, both nepotistic and group efficiency benefits can simultaneously accrue with the replacement of worker-laid eggs. Therefore, worker behaviour towards eggs cannot completely resolve whether both processes have been equally evolutionarily important. Adequate resolution requires the presentation of worker-produced brood of various ages. The stage at which brood are replaced can discriminate whether worker policing occurs owing to a preference for closer genetic kin, a preference for the more reliable brood or both.

Ants are social and are haplodiploid. This combination may allow the evolution of a variety of unusual genetic pathways to achieve reproductive success. These include hybridizing across species, differential use of sperm to create a hybrid worker population, and reproductively isolated gene pools that depend on each other for their survival. Although there are demonstrable costs for colony development and reproduction, these phenomena may nevertheless be relatively common in nature. The specific ecological advantages that favor the evolution of these reproductive modes remain to be discovered.

We observed aggregations of the sand wasp, Steniolia nigripes, in the Granite Mountains of Southern California. Males gathered in open areas between bushes in a sun-dancing behavior very close to the ground. They did not hold exclusive territories. Aggregations were highly visible, but occurred at low densities in the area. Males often chased each other and females, who would approach the aggregation, then leave with males following closely or attached to her. Females often burrowed around the edges of the aggregation. We captured and measured male and female head width, dry weight and thorax length. Males were significantly larger than females in all of these measurements. Intrasexual selection resulting from scramble competition for access to mates may drive this dimorphism. As these aggregations did not occur at obvious landmarks, it is possible that the aggregations themselves serve as a landmarks for females.

Although theories of parental investment and sex ratio generally assume that a single resource limits reproduction, many organisms invest two or more qualitatively different types of resources in the production of offspring. We examine the consequences of multifaceted parental investment for offspring provisioning and sex allocation, building our argument around a study of the nest-building Hymenoptera (wasps, bees, and ants). We review empirical studies that demonstrate that lifetime reproductive success may be constrained not only by resources used to provision offspring but also by the supply of mature oocytes or, in some cases, by the availability of space within nest sites or the time required to defend nests. Under multifaceted parental investment, the factor limiting parental fitness determines the currency of the optimization problem; parents are predicted to adjust reproductive behavior to maximize fitness returns per unit of the limiting resource. We develop simple models that predict that a greater availability of resources used for provisions will lead to an increase in the amount provisioned per offspring and an increase in the numerical or biomass proportion of females produced. These predictions explain widely observed patterns of variation in offspring provisioning and sex allocation in the nest-building Hymenoptera.