Why do male Wool-carder bees defend a flower patch?

I found the first male Wool Carder Bee, Anthidium manicatum of the year in the garden today, patrolling and basking on the large sage patch that is a favourite if many species of bee. If you have a flower-rich garden with a sunny aspect, chances are that in warm, sunny summer days you will see this chunky bee attacking other flower visitors, including large bumblebees. Male bees from most bee species often just hang around nests where they can mate as females emerge, or patrol flowers favoured by females. Occasionally they 'jump' onto or chase other bees or insects, possibly checking if they are females. For example, Male Hairy-footed flower bees, Anthophora plumipes, who emerge a week or so before females, will jump onto bluebottles, the only insect that roughly resemble the black females of his species. No other male bee is as aggressive actively defending a patch of flowers as Wool carder bees. They even have sharp spines at the end of their abdomen with they can use to harm, or even kill, other bees, so they rapidly become the only bees using the flowers

The abdominal spines of male Wool carder bees

 Most female bees mate once shortly after emergence, so males only had a chance of mating successfully early in the season. Females store the sperm of this early mating and use it to fertilise their eggs throughout the nesting season. Females will repel or avoid courting males vigorously once they've been mated. In contrast, Wool-carder bees are unusual in that females both mate multiple times and accept matings throughout their flight season. Witnessing solitary bees mating is quite rare, but with Wool carder bees, matings are quite frequent. Many females visiting a defended flower patch will mate with the resident male. The male defending a good flower patch will likely gain many matings with several visiting females, but, given what we know about sperm usage in solitary bees, will he benefit from fathering the female's offspring?

Mating Wool carder bees

 In a recent study, Kathrin Lampert and colleagues from Ruhr-University Bochum carried out some experiments to investigate why Wool Carder bee mating behaviour is so different from other bees. They in particular were interested in testing the hypothesis that Wool-carder bees might show what is known as 'late male sperm preference' in which the last male mating with a female has a disproportionally higher chance of fathering the females offspring.
 To find out 'who is the daddy' they used genetic testing in a similar way to how paternity is tested in humans, using genetic markers that have many variants, and therefore are likely to be different between individuals. In order to be able to capture the likely mothers and fathers of particular nests they constructed large flight cages containing Betony plants (for pollen and nectar) and Stachys byzantina (for plant wool, which the females collect to line their nest cells). Wool carder bees nest readily in sections of bamboo sticks, and they fixed artificial nesting sites inside the flight cages. Offspring of a female are found in a linear nest in consecutive order, the deeper cells in the bamboo stick contain the earlier laid eggs, while the cells closer to the entrance contain the later brood, which is likely to be male offspring. They carried out three experiments, in the first one, to test the feasibility of these experiments wild males and females (likely to have mated before) were captured, individually labelled with a dot of paint in the thorax, and released in the flight cage. In the second experiment, males in the cage were swapped to test if males with later access to females would father offspring. In the third experiment males were removed after a few days and females left to nest with no males, to investigate if females are able to store sperm. After the females had completed their nests in all the experiments, DNA samples were taken from the males and females, and them and the offspring found in the artificial nests screened for genetic paternity analyses.
A problem with their experiment were cleptoparasitic wasps, which destroyed the bee larvae and prevented genotyping. Another problem, which stems from bee's sex determination system, is that only female offspring have a dad: males develop from unfertilised eggs, so they only have a mum. They could only find out if offspring were male or female after genetic screening. Despite this, they obtained a number of female offspring from many females.
 The results revealed that the males flying with females at the time the females were provisioning a cell, where they are close to laying the egg, were the most likely fathers of the offspring (84% of the time), whereas most of the remaining fathers were males present in the cage in previous days, supporting the hypothesis of late male sperm preference. Males that were dominant, that is, chased away other males from flowers, tended to father more offspring, as most matings happened on flowers.
 The male removal experiment showed that females were able to store sperm for at least 11 days, and possibly much longer.
 This research suggests that a predisposition for late male sperm precedence in the ancestors of wool carder bees might have been what favoured the aggressive territorial behaviour seen in males, as they can benefit from monopolising floral resources visited by females.

More information
Lampert, K. P., Pasternak, V., Brand, P., Tollrian, R., Leese, F., & Eltz, T. (2014). ‘Late’male sperm precedence in polyandrous wool-carder bees and the evolution of male resource defence in Hymenoptera. Animal Behaviour, 90, 211-217.