Which task a social insect worker engages in is influenced by the worker's age, genotype and the ... more Which task a social insect worker engages in is influenced by the worker's age, genotype and the colony's needs. In the honeybee, Apis mellifera, genotype influences both the age a worker switches tasks and its propensity of engaging in specialist tasks, such as water collecting, which only some workers will perform. In this study, we used colonies with natural levels of genetic diversity and manipulated colony age demography to drastically increase the stimuli for the generalist tasks of foraging and nursing, which all workers are thought to engage in at some point in their lives. We examined the representation of worker patrilines engaged in nursing and foraging before and after the perturbation. The representation of patrilines among foragers and nurses differed from that of their overall colony's population. In the case of foraging, over-and underrepresentation of some patrilines was not simply due to differences in rates of development among patrilines. We show that replacement foragers tend to be drawn from patrilines that were overrepresented among foragers before the perturbation, suggesting that there is a genetic component to the tendency to engage in foraging. In contrast, the representation of patrilines in replacement nurses differed from that in the unperturbed nursing population. Our results show that there is a genetic influence on even the generalist tasks of foraging and nursing, and that the way patrilines in genetically diverse colonies respond to increases in task stimuli depends upon the task. The possible significance of this genetic influence on task allocation is discussed.
Animals frequently have to decide between alternative resources and in social insects these indiv... more Animals frequently have to decide between alternative resources and in social insects these individual choices produce a colony-level decision. The choice of nest site is a particularly critical decision for a social insect colony to make, but the decision making process has still only been studied in a few species. In this study, we investigated nest selection by the Pharaoh's ant, Monomorium pharaonis, a species renowned for its propensity to migrate and its use of multi-component trail pheromones to organise decision-making in other contexts. When presented with the choice of familiar and novel nests of equal quality in a Y setup, colonies preferentially migrated towards the familiar nest, suggesting a form of colony-level 'memory' of potential nest sites. However, if the novel nest was superior to the familiar nest, then colonies began migrating initially to the familiar nest, but then redirected their migration to the superior quality novel nest. This may be an effective method of reducing colony exposure while searching for an optimum nest site. Branches that had previously led to a selected nest were attractive to ants in subsequent migrations, suggesting that trail pheromones mediate the decision making process. The adaptive, pheromone-based organisation of nest-site selection by Pharaoh's ants matches their ephemeral environment and is likely to contribute to their success as a 'tramp' species.
1. Parasites are an important selective force for almost all organisms and drive the evolution by... more 1. Parasites are an important selective force for almost all organisms and drive the evolution by hosts of defence mechanisms that are energetically costly. The strength of parasitism will vary between host species according to their specific ecology and life history, and so the optimal investment in costly resistance mechanisms is also likely to vary between host species with differing ecologies. 2. Parasites are particularly important for social species such as ants, but very little is known about the strength of selection in different species. It has been suggested that, because arboreality reduces exposure to soil-borne fungal pathogens, arboreal ant species may invest less in disease resistance. However, testing hypotheses such as this requires data on disease resistance in multiple species, and such studies have not previously been attempted. 3. Here we examine the arboreality hypothesis by comparing the disease resistance of seven Neotropical ant species with different degrees of arboreality. We exposed ants to controlled doses of the generalist, virulent fungal parasite, Metarhizium anisopliae (Metchnikoff) Sorokin. We then monitored survival, parasite sporulation, and the antifungal grooming response of the ants. 4. Contrary to the hypothesis, we found that arboreal species were not less resistant to M. anisopliae than species that were ground-dwelling, and that the species that inhabited both arboreal and ground habitats had the greatest resistance. Surprisingly, the most resistant species was one that lacked the antibiotic-producing metapleural gland, previously considered the lynchpin of disease resistance in ants. 5. The results suggest that it may be the diversity of parasites encountered that is the greatest selection pressure. Further experimental studies with other parasites are needed to confirm the generality of the results, and similar comparative studies of other taxa are needed to understand the relationship between host ecology and the evolution of disease resistance.
The evolution of polyandry is a general problem in behavioral ecology that has attracted particul... more The evolution of polyandry is a general problem in behavioral ecology that has attracted particular interest in the social insects. Most social insects are monandrous (i.e., females are inseminated by a single male), but approximately a third of species have evolved polyandry (i.e., females are inseminated by multiple males), which can reach extreme levels. One of the leading explanations for polyandry is that genetically diverse colonies may have improved division of labor because of genotypic variation in the propensity of workers to engage in particular tasks. Here we investigate whether there may also be genotypic variation in the ability to carry out a task, by examining the effect of genotype, as well as caste and age, on the foraging performance of workers of the leaf-cutting ant Acromyrmex subterraneus. Larger ants cut larger fragments of orange-impregnated parafilm, and did so more quickly than smaller ants, whereas older ants also cut larger fragments but were slower to transport them. Most importantly, we found that patrilines within colonies differed significantly in the size of fragment cut, and the speed of cutting and transporting fragments when controlling for both ant size and fragment size in the analyses. Certain patrilines were better foragers, often in multiple ways, whereas other patrilines were significantly worse at foraging. Genotype can therefore affect the ability of social insect workers to carry out tasks as well as their previously shown propensity to engage in them, providing an additional mechanism by which genetic diversity may be beneficial to social insect colonies.
1. Over a million commercially produced bumblebee colonies are imported annually on a global scal... more 1. Over a million commercially produced bumblebee colonies are imported annually on a global scale for the pollination of greenhouse crops. After importation, they interact with other pollinators, with an associated risk of any parasites they carry infecting and harming native bees. National and supranational regulations are designed to prevent this, and commercially produced bumblebee colonies are accordingly now often sold and imported as being parasite-free. 2. Here, we used molecular methods to examine the occurrence of parasites in bumblebee colonies that were commercially produced in 2011 and 2012 by three producers. We then used controlled experiments to determine whether any parasites present were infectious. 3. We found that 77% of the commercially produced bumblebee colonies from the three producers, which were imported on the basis of being free of parasites, in fact carried microbial parasites, with five different parasites being detected across the total sample of bumblebees and a further three in the pollen supplied with the colonies as food. 4. Our controlled experiments demonstrated that at least three of these parasites were infectious to bumblebees with significant negative effects on their health. Furthermore, we also found that at least four of the parasites carried by commercially produced bumblebees were infectious to honeybees, indicating that they pose a risk to other pollinators as well. 5. Synthesis and applications. The results demonstrate that commercially produced bumblebee colonies carry multiple, infectious parasites that pose a significant risk to other native and managed pollinators. More effective disease detection and management strategies are urgently needed to reduce the pathogen spillover threat from commercially produced bumblebees.
The effective communication of alarm can be critical for social animals so that they are able to ... more The effective communication of alarm can be critical for social animals so that they are able to deal with threats posed by predators and competitors. In the case of many of the most ecologically dominant, large-colony ant species, these alarm responses are aggressive and coordinated by alarm pheromones, produced generally from the mandibular glands. In the present study, the alarm behaviour of two Neotropical army ant species is examined, the swarm raiding Eciton burchellii (Westwood) and the column raiding Eciton hamatum (Fabricius). Both species exhibit aggressive alarm responses in response to crushed heads, suggesting that the alarm pheromone is indeed produced by the mandibular glands in these ants. The most abundant component of the mandibular gland secretion, 4-methyl-3-heptanone (10 μL on a rubber septum), stimulates a substantial alarm response, although this is less than the response to a single crushed head. This suggests that 4-methyl-3-heptanone may be an alarm-stimulating compound in Eciton. The alarm response of E. burchellii involves more workers than that of E. hamatum, although major workers play a much greater role in the response of the latter species. The differences in the alarm response of the two closely-related species may relate to their foraging strategies, with E. burchellii relying more on quantity rather than the caste of ants responding and possibly using alarm pheromones for offensive as well as defensive functions.
The enhancement of bait for the control of grass-cutting ants was investigated using two species ... more The enhancement of bait for the control of grass-cutting ants was investigated using two species of grass-cutting ant, Atta bisphaerica (Forel) and Atta capiguara (Gonçalves) (Hymenoptera: Formicidae). Bait was applied in loose piles to obtain a direct relationship between ant attraction and bait harvest. Enhancement with alarm pheromone compounds significantly increased the attractiveness and harvest of bait under certain conditions. A large proportion of the ants attracted to the enhanced bait were minor workers. These ants rarely transport bait because of their small size, and so it may be possible to increase the effect of bait enhancement by using smaller bait granules. Foragers of A. capiguara were less inclined to transport citrus-pulp bait than were those of Atta laevigata (Fr. Smith), a species that also harvests dicotyledonous plants. This emphasizes the importance of developing a bait matrix that is more acceptable to grass-cutting species. Nevertheless, the results suggest that alarm pheromone compounds have significant potential to improve the efficacy of baits for the control of grass-cutting ants.
Which task a social insect worker engages in is influenced by the worker's age, genotype and the ... more Which task a social insect worker engages in is influenced by the worker's age, genotype and the colony's needs. In the honeybee, Apis mellifera, genotype influences both the age a worker switches tasks and its propensity of engaging in specialist tasks, such as water collecting, which only some workers will perform. In this study, we used colonies with natural levels of genetic diversity and manipulated colony age demography to drastically increase the stimuli for the generalist tasks of foraging and nursing, which all workers are thought to engage in at some point in their lives. We examined the representation of worker patrilines engaged in nursing and foraging before and after the perturbation. The representation of patrilines among foragers and nurses differed from that of their overall colony's population. In the case of foraging, over-and underrepresentation of some patrilines was not simply due to differences in rates of development among patrilines. We show that replacement foragers tend to be drawn from patrilines that were overrepresented among foragers before the perturbation, suggesting that there is a genetic component to the tendency to engage in foraging. In contrast, the representation of patrilines in replacement nurses differed from that in the unperturbed nursing population. Our results show that there is a genetic influence on even the generalist tasks of foraging and nursing, and that the way patrilines in genetically diverse colonies respond to increases in task stimuli depends upon the task. The possible significance of this genetic influence on task allocation is discussed.
Animals frequently have to decide between alternative resources and in social insects these indiv... more Animals frequently have to decide between alternative resources and in social insects these individual choices produce a colony-level decision. The choice of nest site is a particularly critical decision for a social insect colony to make, but the decision making process has still only been studied in a few species. In this study, we investigated nest selection by the Pharaoh's ant, Monomorium pharaonis, a species renowned for its propensity to migrate and its use of multi-component trail pheromones to organise decision-making in other contexts. When presented with the choice of familiar and novel nests of equal quality in a Y setup, colonies preferentially migrated towards the familiar nest, suggesting a form of colony-level 'memory' of potential nest sites. However, if the novel nest was superior to the familiar nest, then colonies began migrating initially to the familiar nest, but then redirected their migration to the superior quality novel nest. This may be an effective method of reducing colony exposure while searching for an optimum nest site. Branches that had previously led to a selected nest were attractive to ants in subsequent migrations, suggesting that trail pheromones mediate the decision making process. The adaptive, pheromone-based organisation of nest-site selection by Pharaoh's ants matches their ephemeral environment and is likely to contribute to their success as a 'tramp' species.
1. Parasites are an important selective force for almost all organisms and drive the evolution by... more 1. Parasites are an important selective force for almost all organisms and drive the evolution by hosts of defence mechanisms that are energetically costly. The strength of parasitism will vary between host species according to their specific ecology and life history, and so the optimal investment in costly resistance mechanisms is also likely to vary between host species with differing ecologies. 2. Parasites are particularly important for social species such as ants, but very little is known about the strength of selection in different species. It has been suggested that, because arboreality reduces exposure to soil-borne fungal pathogens, arboreal ant species may invest less in disease resistance. However, testing hypotheses such as this requires data on disease resistance in multiple species, and such studies have not previously been attempted. 3. Here we examine the arboreality hypothesis by comparing the disease resistance of seven Neotropical ant species with different degrees of arboreality. We exposed ants to controlled doses of the generalist, virulent fungal parasite, Metarhizium anisopliae (Metchnikoff) Sorokin. We then monitored survival, parasite sporulation, and the antifungal grooming response of the ants. 4. Contrary to the hypothesis, we found that arboreal species were not less resistant to M. anisopliae than species that were ground-dwelling, and that the species that inhabited both arboreal and ground habitats had the greatest resistance. Surprisingly, the most resistant species was one that lacked the antibiotic-producing metapleural gland, previously considered the lynchpin of disease resistance in ants. 5. The results suggest that it may be the diversity of parasites encountered that is the greatest selection pressure. Further experimental studies with other parasites are needed to confirm the generality of the results, and similar comparative studies of other taxa are needed to understand the relationship between host ecology and the evolution of disease resistance.
The evolution of polyandry is a general problem in behavioral ecology that has attracted particul... more The evolution of polyandry is a general problem in behavioral ecology that has attracted particular interest in the social insects. Most social insects are monandrous (i.e., females are inseminated by a single male), but approximately a third of species have evolved polyandry (i.e., females are inseminated by multiple males), which can reach extreme levels. One of the leading explanations for polyandry is that genetically diverse colonies may have improved division of labor because of genotypic variation in the propensity of workers to engage in particular tasks. Here we investigate whether there may also be genotypic variation in the ability to carry out a task, by examining the effect of genotype, as well as caste and age, on the foraging performance of workers of the leaf-cutting ant Acromyrmex subterraneus. Larger ants cut larger fragments of orange-impregnated parafilm, and did so more quickly than smaller ants, whereas older ants also cut larger fragments but were slower to transport them. Most importantly, we found that patrilines within colonies differed significantly in the size of fragment cut, and the speed of cutting and transporting fragments when controlling for both ant size and fragment size in the analyses. Certain patrilines were better foragers, often in multiple ways, whereas other patrilines were significantly worse at foraging. Genotype can therefore affect the ability of social insect workers to carry out tasks as well as their previously shown propensity to engage in them, providing an additional mechanism by which genetic diversity may be beneficial to social insect colonies.
1. Over a million commercially produced bumblebee colonies are imported annually on a global scal... more 1. Over a million commercially produced bumblebee colonies are imported annually on a global scale for the pollination of greenhouse crops. After importation, they interact with other pollinators, with an associated risk of any parasites they carry infecting and harming native bees. National and supranational regulations are designed to prevent this, and commercially produced bumblebee colonies are accordingly now often sold and imported as being parasite-free. 2. Here, we used molecular methods to examine the occurrence of parasites in bumblebee colonies that were commercially produced in 2011 and 2012 by three producers. We then used controlled experiments to determine whether any parasites present were infectious. 3. We found that 77% of the commercially produced bumblebee colonies from the three producers, which were imported on the basis of being free of parasites, in fact carried microbial parasites, with five different parasites being detected across the total sample of bumblebees and a further three in the pollen supplied with the colonies as food. 4. Our controlled experiments demonstrated that at least three of these parasites were infectious to bumblebees with significant negative effects on their health. Furthermore, we also found that at least four of the parasites carried by commercially produced bumblebees were infectious to honeybees, indicating that they pose a risk to other pollinators as well. 5. Synthesis and applications. The results demonstrate that commercially produced bumblebee colonies carry multiple, infectious parasites that pose a significant risk to other native and managed pollinators. More effective disease detection and management strategies are urgently needed to reduce the pathogen spillover threat from commercially produced bumblebees.
The effective communication of alarm can be critical for social animals so that they are able to ... more The effective communication of alarm can be critical for social animals so that they are able to deal with threats posed by predators and competitors. In the case of many of the most ecologically dominant, large-colony ant species, these alarm responses are aggressive and coordinated by alarm pheromones, produced generally from the mandibular glands. In the present study, the alarm behaviour of two Neotropical army ant species is examined, the swarm raiding Eciton burchellii (Westwood) and the column raiding Eciton hamatum (Fabricius). Both species exhibit aggressive alarm responses in response to crushed heads, suggesting that the alarm pheromone is indeed produced by the mandibular glands in these ants. The most abundant component of the mandibular gland secretion, 4-methyl-3-heptanone (10 μL on a rubber septum), stimulates a substantial alarm response, although this is less than the response to a single crushed head. This suggests that 4-methyl-3-heptanone may be an alarm-stimulating compound in Eciton. The alarm response of E. burchellii involves more workers than that of E. hamatum, although major workers play a much greater role in the response of the latter species. The differences in the alarm response of the two closely-related species may relate to their foraging strategies, with E. burchellii relying more on quantity rather than the caste of ants responding and possibly using alarm pheromones for offensive as well as defensive functions.
The enhancement of bait for the control of grass-cutting ants was investigated using two species ... more The enhancement of bait for the control of grass-cutting ants was investigated using two species of grass-cutting ant, Atta bisphaerica (Forel) and Atta capiguara (Gonçalves) (Hymenoptera: Formicidae). Bait was applied in loose piles to obtain a direct relationship between ant attraction and bait harvest. Enhancement with alarm pheromone compounds significantly increased the attractiveness and harvest of bait under certain conditions. A large proportion of the ants attracted to the enhanced bait were minor workers. These ants rarely transport bait because of their small size, and so it may be possible to increase the effect of bait enhancement by using smaller bait granules. Foragers of A. capiguara were less inclined to transport citrus-pulp bait than were those of Atta laevigata (Fr. Smith), a species that also harvests dicotyledonous plants. This emphasizes the importance of developing a bait matrix that is more acceptable to grass-cutting species. Nevertheless, the results suggest that alarm pheromone compounds have significant potential to improve the efficacy of baits for the control of grass-cutting ants.
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