The other insect societies: overview and new directions.

The diversity of societies and forms of social interaction across the Arthropoda is commensurate with the great taxonomic diversity within this pylum. Social evolution research has, however, largely focused on a small subset of social forms; namely, those deemed to be 'eusocial'-groups exhibiting overlapping generations, cooperative brood care, and reproductive division of labor. Here I provide a brief overview of the 'other', non-eusocial, societies of insects and allies, defining the main social traits of interest and summarizing recent work. Four active and emerging fields of inquiry in the other insect societies are discussed.

Social nutrition: an emerging field in insect science.

Nutrition is thought to be a major driver of social evolution, yet empirical support for this hypothesis is scarce. Here we illustrate how conceptual advances in nutritional ecology illuminate some of the mechanisms by which nutrition mediates social interactions in insects. We focus on experiments and models of nutritional geometry and argue that they provide a powerful means for comparing nutritional phenomena across species exhibiting various social ecologies. This approach, initially developed to study the nutritional behaviour of individual insects, has been increasingly used to study insect groups and societies, leading to the emerging field of social nutrition. We discuss future directions for exploring how these nutritional mechanisms may influence major social transitions in insects and other animals.

Adaptations: Using Darwin's Origin to teach biology and writing.

Charles Darwin's On the Origin of Species is at once familiar and unfamiliar. Everyone knows that the Origin introduced the world to the idea of evolution by natural selection, but few of us have actually read it. We suggest that it is worth taking the time not only to read what Darwin had to say, but also to use the Origin to teach both biology and writing. It provides scientific lessons in areas beyond evolutionary biology, such as ecology and biogeography. In addition, it provides valuable rhetorical lessons-how to construct an argument, write persuasively, make use of evidence, know your audience, and anticipate counterarguments. We have been using the Origin in various classes for several years, introducing new generations to Darwin, in his own words.

Hamiltonian inclusive fitness: a fitter fitness concept.

In 1963-1964 W. D. Hamilton introduced the concept of inclusive fitness, the only significant elaboration of Darwinian fitness since the nineteenth century. I discuss the origin of the modern fitness concept, providing context for Hamilton's discovery of inclusive fitness in relation to the puzzle of altruism. While fitness conceptually originates with Darwin, the term itself stems from Spencer and crystallized quantitatively in the early twentieth century. Hamiltonian inclusive fitness, with Price's reformulation, provided the solution to Darwin's 'special difficulty'-the evolution of caste polymorphism and sterility in social insects. Hamilton further explored the roles of inclusive fitness and reciprocation to tackle Darwin's other difficulty, the evolution of human altruism. The heuristically powerful inclusive fitness concept ramified over the past 50 years: the number and diversity of 'offspring ideas' that it has engendered render it a fitter fitness concept, one that Darwin would have appreciated.

Fitness effects of group merging in a social insect.

Animal social groups often consist of non-relatives, a condition that arises in many cases because of group merging. Although indirect fitness contributions are reduced in such groups compared with those in groups composed of close kin, the genetic-heterogeneity hypothesis suggests that these groups may benefit from increased intracolony genetic variation, which may boost group performance through increased task efficiency or parasite resistance. We confirm one prediction of the task-efficiency explanation by demonstrating a genetic basis for task thresholds of socially important behaviours in eastern tent caterpillars. However, we found no evidence that the expanded range of task thresholds in mixed colonies translates into improved individual or colony performance in the field. By contrast, increased group size, a less commonly considered correlate of group mixing, was found to enhance individual fitness through its effects on larval growth. We conclude that fitness benefits offsetting the dilution of relatedness in heterogeneous social groups may often stem from augmented group size rather than increased genotypic diversity.

HIERARCHICAL GENETIC STRUCTURE AND GENE FLOW IN MACROGEOGRAPHIC POPULATIONS OF THE EASTERN TENT CATERPILLAR (MALACOSOMA AMERICANUM).

Genetic structure and inferred rates of gene flow in macrogeographic populations of the eastern tent caterpillar Malacosoma americanum were analyzed at two hierarchical scales: local demes and regional subpopulations. Wright's F-statistics were used to estimate population genetic structure using multilocus genotypic data generated electrophoretically. Estimated values of FST and the distribution of private alleles were then used to obtain indirect estimates of gene flow. We found modest, though significant, genetic structure at both spatial scales, a pattern consistent with high rates of gene flow over the large distances involved. Modest values obtained for Nei's genetic distance also suggested high levels of gene flow across the range of this species, although some gene-flow restriction resulting from isolation by distance was suggested by a positive regression of genetic distance on geographic distance. The observed homogeneity at enzyme loci across the range of M. americanum parallels the reported uniformity in morphology, suggesting a general absence of local genetic differentiation in this widely distributed species. The genetic homogeneity observed in this wide-ranging insect is discussed in terms of organism-specific environmental experience at different spatial scales. Some organisms occupying apparently heterogeneous environments may ameliorate unsuitable local conditions through microhabitat selection or behavioral modification of their microenvironment. This may be accomplished in M. americanum through group shelter construction and behavioral thermoregulation, closely tying thermoregulation to social biology in this species. If in this way the tent helps produce an effectively homogeneous environment for this species across its extensive range, this system may provide a unique example of how social behavior can influence the distribution of genetic variation in a population.