The benefits of being big and diverse: early colony survival in harvester ants.

In sessile organisms such as plants and benthic invertebrates, founding propagules typically suffer extremely high rates of mortality due to both extrinsic and intrinsic factors. Many social insect species share similarities with these groups, but factors influencing early colony survival are relatively unstudied. We used a field experiment to measure the importance of environmental quality relative to intrinsic colony properties in the harvester ant, Pogonomyrmex occidentalis, by monitoring the survival of 584 experimental colonies.

Offspring Size and Reproductive Allocation in Harvester Ants.

A fundamental decision that an organism must make is how to allocate resources to offspring, with respect to both size and number. The two major theoretical approaches to this problem, optimal offspring size and optimistic brood size models, make different predictions that may be reconciled by including how offspring fitness is related to size. We extended the reasoning of Trivers and Willard (1973) to derive a general model of how parents should allocate additional resources with respect to the number of males and females produced, and among individuals of each sex, based on the fitness payoffs of each.

Inclusive fitness theory and eusociality.

Arising from M. A. Nowak, C.

Patriline shifting leads to apparent genetic caste determination in harvester ants.

The harvester ant, Pogonomyrmex occidentalis, is characterized by high levels of intracolonial genetic diversity resulting from multiple mating by the queen. Within reproductively mature colonies, the relative frequency of different male genotypes (patrilines) is not stable. The difference between samples increases with time, nearing an asymptote after a year.

Dioecy and the evolution of sex ratios in ants.

Split sex ratios, when some colonies produce only male and others only female reproductives, is a common feature of social insects, especially ants. The most widely accepted explanation for split sex ratios was proposed by Boomsma and Grafen, and is driven by conflicts of interest among colonies that vary in relatedness. The predictions of the Boomsma-Grafen model have been confirmed in many cases, but contradicted in several others.

Mating for variety increases foraging activity in the harvester ant, Pogonomyrmex occidentalis.

Multiple mating by females characterizes most insect species, but is relatively uncommon in social insects. Females may mate with multiple mates because they experience the direct benefits of increased survival or fecundity, to acquire high quality mates, or to lower the risk of reduced fecundity by mating with incompatible males. We used the extensive natural variation in mating frequency in the western harvester ant, Pogonomyrmex occidentalis, to test the hypothesis that increased mating by the queen leads to an increase in colony performance.

Polyandry and fitness in the western harvester ant, Pogonomyrmex occidentalis.

Using four highly polymorphic microsatellite markers (12-28 alleles), we gentoyped workers from 63 colonies of Pogonomyrmex occidentalis. Colonies have a single, multiply mated queen, and an average number of 6.3 patrilines per colony.

Queen size mediates queen survival and colony fitness in harvester ants.

We examined the effect of queen size on the probability of new colony establishment in the ant Pogonomyrmex occidentalis. Large queens are significantly more likely to survive than small queens through the initial stages of colony founding. These differences in individual fitness correlates have corresponding effects on colony fitness.

Recruitment in the harvester ant, Pogonomyrmex occidentalis: effects of experimental removal.

We examined the importance of experimental removal of mature colonies on colony recruitment in the western harvester ant Pogonomyrmex occidentalis. To test the common assumption that established colonies suppress the establishment of new colonies we removed all colonies in ten 0.25 ha plots in 1996 and an additional five plots in 1997 and measured new colony recruitment in 1997, 1998, and 1999.

SEXUAL SELECTION ON BODY SIZE AND SHAPE IN THE WESTERN HARVESTER ANT, POGONOMYRMEX OCCIDENTALIS CRESSON.

Mating in social insects has generally been studied in relation to reproductive allocation and relatedness. Despite the tremendous morphological diversity in social insects, little is known about how individual morphology affects mating success. We examined the correlation of male size and shape with mating success in the western harvester ant, Pogonomyrmex occidentalis.

DEVELOPMENT, FUNCTION, AND THE QUANTITATIVE GENETICS OF WING MELANIN PATTERN IN PIERIS BUTTERFLIES.

Do genetic correlations among phenotypic characters reflect developmental organization or functional coadaptation of the characters? We test these hypotheses for the wing melanin pattern of Pieris occidentalis butterflies, by comparing estimated genetic correlations among wing melanin characters with a priori predictions of the developmental organization and the functional (thermoregulatory) organization of melanin pattern. There were significant broad-sense heritabilities and significant genetic correlations for most melanin characters. Matrix correlation tests revealed significant agreement between the observed genetic correlations and both developmental and functional predictions in most cases; this occurred even when the overlap between developmental and functional predictions was eliminated.

FEMALE CHOICE AND SEXUAL SELECTION OF MALE WING MELANIN PATTERN IN PIERIS OCCIDENTALIS (LEPIDOPTERA).

Using field experiments, I examined the role of 13 melanin-pattern elements in mate choice by female Pieris occidentalis butterflies. Males that mated successfully differed significantly from unsuccessful males in the extent of marginal forewing melanization but not on the basis of the entire forewing pattern. Deletion of the marginal forewing melanin characters significantly reduced the mating success of manipulated males relative to controls under field conditions.

DISSECTING CORRELATED CHARACTERS: ADAPTIVE ASPECTS OF PHENOTYPIC COVARIATION IN MELANIZATION PATTERN OF PIERIS BUTTERFLIES.

In this study we address the question of how much of the covariation among phenotypic characters observed in natural populations is adaptive. We examine covariation among a set of phenotypic characters that describe the wing-melanization pattern of Pieris butterflies. Previous functional analyses of thermoregulatory performance allow us to predict a priori whether and how different wing melanic characters should be correlated.