Close relatedness has long been considered crucial to the evolution of eusociality. However, it has recently been suggested that close relatedness may be a consequence, rather than a cause, of eusociality. We tested this idea with a comparative analysis of female mating frequencies in 267 species of eusocial bees, wasps, and ants. We found that mating with a single male, which maximizes relatedness, is ancestral for all eight independent eusocial lineages that we investigated. Mating with multiple males is always derived. Furthermore, we found that high polyandry (>2 effective mates) occurs only in lineages whose workers have lost reproductive totipotency. These results provide the first evidence that monogamy was critical in the evolution of eusociality, strongly supporting the prediction of inclusive fitness theory.

Summary 1.Waggle dances of honey-bees ( Apis mellifera L . ) were decoded to determine where and how far the bees foraged during the blooming of heather ( Calluna vulgaris L.) in August 1996 using a hive located in Sheffield, UK, east of the heather moors. The median distance foraged was 6·1 km, and the mean 5·5 km. Only 10% of the bees foraged within 0·5 km of the hive whereas 50% went more than 6 km, 25% more than 7·5 km and 10% more than 9·5 km from the hive. 2. These results are in sharp contrast with previous studies in which foraging distances were much closer to the hive. In May 1997 the mean foraging distance was 1 km, showing that long-range dancing is not the rule in Sheffield. 3. The observed foraging distances described in this study may not be exceptional in a patchy environment where differences in patch size and patch quality are large. When travel distances to patches are large, distant patches can probably be utilized only by individuals that live in groups and recruit foragers to the patches found. Only then are the benefits of scouting for distant patches high enough to enable the exploitation of these patches.

Recent evidence for genetic effects on royal and worker caste differentiation from diverse social insect taxa has put an end to the view that these phenotypes stem solely from a developmental switch controlled by environmental factors. Instead, the relative influences of genotypic and environmental effects on caste vary among species, ranging from largely environmentally controlled phenotypes to almost purely genetic systems. Disentangling the selective forces that generate variation for caste predisposition will require characterizing the genetic mechanisms underlying this variation, and identifying particular life-history strategies and kin structures associated with strong genetic effects on caste.

A fundamental question in nutritional biology is how distributed systems maintain an optimal supply of multiple nutrients essential for life and reproduction. In the case of animals, the nutritional requirements of the cells within the body are coordinated by the brain in neural and chemical dialogue with sensory systems and peripheral organs. At the level of an insect society, the requirements for the entire colony are met by the foraging efforts of a minority of workers responding to cues emanating from the brood. Both examples involve components specialized to deal with nutrient supply and demand (brains and peripheral organs, foragers and brood). However, some of the most species-rich, largest, and ecologically significant heterotrophic organisms on earth, such as the vast mycelial networks of fungi, comprise distributed networks without specialized centers: How do these organisms coordinate the search for multiple nutrients? We address this question in the acellular slime mold Physarum polycephalum and show that this extraordinary organism can make complex nutritional decisions, despite lacking a coordination center and comprising only a single vast multinucleate cell. We show that a single slime mold is able to grow to contact patches of different nutrient quality in the precise proportions necessary to compose an optimal diet. That such organisms have the capacity to maintain the balance of carbon-and nitrogen-based nutrients by selective foraging has considerable implications not only for our understanding of nutrient balancing in distributed systems but for the functional ecology of soils, nutrient cycling, and carbon sequestration. acellular slime mold | complexity | geometrical framework | nutrition | Physarum polycephalum P lasmodia of Physarum polycephalum are single multinucleate cells extending up to hundreds of square centimeters. Cytoplasm streams rhythmically back and forth through a network of tubular elements, circulating nutrients and chemical signals and forming pseudopods that allow the organism to navigate around and respond to its environment. Plasmodia are distributed information processors, which, for example, can find the shortest route through a maze to locate food (1), anticipate the timing of periodic events (2), and solve multiobjective foraging problems (3).Under adequate nutrition, P. polycephalum plasmodia are completely sedentary and grow steadily (4, 5), but on nonnutrient substrates, they migrate a few centimeters per hour (6), directed by external stimuli, including gradients of nutrients such as sugars and proteins (7-12). When two or more identical food sources are presented at various positions to a starved plasmodium, it optimizes the shape of the network to facilitate effective absorption of nutrients (1), and plasmodia select the higher concentration patch of two patches differing in nutrient concentration (3). Can it solve complex nutrient balancing problems by altering its growth form and movement to maintain an optimal ratio of macronutrients in the face of variati...

Spatial memory enhances an organism's navigational ability. Memory typically resides within the brain, but what if an organism has no brain? We show that the brainless slime mold Physarum polycephalum constructs a form of spatial memory by avoiding areas it has previously explored. This mechanism allows the slime mold to solve the U-shaped trap problem-a classic test of autonomous navigational ability commonly used in robotics-requiring the slime mold to reach a chemoattractive goal behind a U-shaped barrier. Drawn into the trap, the organism must rely on other methods than gradient-following to escape and reach the goal. Our data show that spatial memory enhances the organism's ability to navigate in complex environments. We provide a unique demonstration of a spatial memory system in a nonneuronal organism, supporting the theory that an externalized spatial memory may be the functional precursor to the internal memory of higher organisms. extracellular slime | protist | reactive navigation | amoeboid organism

The complex collective behavior seen in many insect societies strongly suggests that a minimum number of workers are required for these societies to function effectively. Here we investigated the transition between disordered and ordered foraging in the Pharaoh's ant. We show that small colonies forage in a disorganized manner, with a transition to organized pheromone-based foraging in larger colonies. We also show that when food sources are difficult to locate through independent searching, this transition is first-order and exhibits hysteresis, comparable to a first-order phase transition found in many physical systems. To our knowledge, this is the first experimental evidence of a behavioral phase transition between a maladaptive (disorganized) and an adaptive (organized) state.

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Honey bee workers maintain the brood nest of their colony within a narrow temperature range of 34.5+/-1.5 degrees C, implying that there are significant fitness costs if brood is reared outside the normal range. However, the effects of abnormal incubation temperatures are subtle and not well documented. Here we show that short-term learning and memory abilities of adult workers are affected by the temperature they experienced during pupal development. In contrast, long-term learning and memory is not significantly affected by rearing temperature. Furthermore, we could detect no effects of incubation temperature on fluctuating asymmetry, as a measure of developmental stability, in workers, queens or drones. We conclude that the most important consequence of abnormal rearing temperatures are subtle neural deficiencies affecting short-term memory rather than physical abnormalities.