Alien Colonies Research Articles

There and Back Again: A Commentary on Social Welfare Policy in the Wake of 2020

There and Back Again: A Commentary on Social Welfare Policy in the Wake of 2020

Population and colony structure of an ant with territorial males, Cardiocondyla venustula

BackgroundMany species of social insects have large-scale mating and dispersal flights and their populations are therefore often relatively homogenous. In contrast, dispersal on the wing appears to be uncommon in most species of the ant genus Cardiocondyla, because its males are wingless and the winged queens mate in their natal nests before dispersing on foot. Here we examine the population structure of C. venustula from South Africa. This species is of particular interest for the analysis of life history evolution in Cardiocondyla, as it occupies a phylogenetic position between tropical species with multi-queen (polygynous) colonies and fighting males and a Palearctic clade with single-queen colonies and mutually peaceful males. Males of C. venustula exhibit an intermediate strategy between lethal fighting and complete tolerance – they mostly engage in non-lethal fights and defend small territories inside their natal nests. We investigated how this reproductive behavior influences colony and population structure by analyzing samples on two geographic scales in South Africa: a small 40 × 40m2 plot and a larger area with distances up to 5 km between sampling sites in Rietvlei Nature Reserve near Pretoria.ResultsColonies were found to be facultatively polygynous and queens appear to mate only with a single male. The extraordinarily high inbreeding coefficient suggests regular sib-mating. Budding by workers and young queens is the predominant mode of colony-founding and leads to high population viscosity. In addition, some queens appear to found colonies independently or through adoption into foreign nests.ConclusionWhile C. venustula resembles tropical Cardiocondyla in queen number and mating frequency, it differs by the absence of winged disperser males. Dispersal by solitary, mated queens on foot or by short flights and their adoption by alien colonies might promote gene flow between colonies and counteract prolonged inbreeding. The abundance of suitable habitat and the high density of nests facilitate the spread of this species by budding and together with the apparent resistance against inbreeding make it a highly successful pioneer species and invader of degraded and man-made habitats.

Discriminatory abilities of facultative slave-making ants and their slaves.

Intra-colony odor variability can disturb ants’ ability to discriminate against intruders. The evolutionary relevance of this phenomenon can be revealed by studies on colonies of slave-making ants in which the parasite, and not the host, is subject to selection pressures associated with living in a mixed colony. We examined how the European facultative slave-making species Formica sanguinea and its F. fusca slaves perform in discriminating ants from alien colonies. Results of behavioral assays showed that slave-maker ants respond with hostility to conspecific individuals from alien colonies but are relatively tolerant to alien slaves. Furthermore, the behavior of slaves indicated a limited ability to discriminate ants from alien parasitic colonies. The subdivision of colony fragments into mixed and species-separated groups demonstrated that contact with the parasite is necessary for F. fusca slaves to be re-accepted by former nestmates after a period of separation from the stock colony. The results presented in this paper are consistent with the following hypotheses: (1) F. sanguinea ants, as opposed to their slaves, are adapted to discriminate alien individuals in the conditions of odor variability found in a mixed-species colony, (2) the recognition of slaves by F. sanguinea ants involves a dedicated adaptive mechanism that prevents aggression toward them, (3) the odor of slaves is strongly influenced by the parasite with beneficial effect on the colony integrity.Electronic supplementary materialThe online version of this article (doi:10.1007/s00040-016-0493-z) contains supplementary material, which is available to authorized users.

Post-mortem Changes in Chemical Profile and their Influence on Corpse Removal in Ants

In order to decrease the risk of pathogen transmission, ants remove corpses from the vicinity of nests, but little is known about the underlying mechanisms. In particular, it is unclear how the odor profile of corpses changes with time since death and how any changes might relate to behavior. We have addressed these questions in the red ant Myrmica rubra, where we asked how the time since death determines the ability of workers to discriminate a dead individual from a live one, and whether dead workers are removed in a similar way when they originate from the same or an alien colony. We found that ants could discriminate alien from nestmate corpses up to 2 days after death, since the former continued to elicit aggressive behavior over that period. For dead nestmates, only 15 % of corpses were removed when freshly killed but this rises to 80 % for corpses between 1 to 6 days post mortem. Using gas chromatography, we found that oleic and linoleic acids, which are absent on freshly killed corpses, appeared post mortem and were in higher quantities on those corpses that were ejected from the nest vicinity. When added to fresh corpses, linoleic and oleic acids, alone or blended, enhanced removal to levels observed for corpses of 2-6 days post-mortem. Thus, oleic and linoleic acids appear to be important cues involved in corpse recognition and necrophoresis over a long timeframe, and we advance the hypothesis that these fatty acids in combination with other cues may also trigger other behaviors such as prey retrieval.

Comparative morphology of Van der Vecht's organ inPolistessocial parasites: host ecology and adaptation of the parasite

Camouflage strategies are common in insect social parasites. Being accepted into an alien colony as a dominant nestmate favours behavioural and morphological adaptations to mimic a specific odour. In Polistes social parasites, abdominal tegumental glands are involved in this camouflage strategy. These glands secreting cuticular hydrocarbons are connected with a modified cuticular area of the last gastral sternite of female wasps, named Van der Vecht’s organ, whose secretion is involved in rank and dominance recognition. The size of this exocrine area has been demonstrated to be under selective pressure in Polistes, as a response to an efficient dominance recognition. Because chemical and behavioural integration differs between parasitic species, we carried out a comparison of Van der Vecht’s organ size between the three Polistes social parasites and their respective hosts. The parasites Polistes sulcifer and Polistes semenowi, capable of a rapid chemical mimicry and specialized to exploit a lowland host, also show an enlarged Van der Vecht’s organ. Conversely, the parasite Polistes atrimandibularis, specialized on a mountain species and showing a slow chemical integration, has a smaller organ. The time available for the parasite to tune up its chemical mimicry, before the emergence of workers to be accepted as a dominant nestmate, appears to be the most important selective pressure acting on the size of this abdominal organ. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, , ••‐••.

The differential response of workers and queens of the ant Lasius niger to an environment marked by workers: Ants dislike the unknown

It is well known that ants can use cuticular hydrocarbons (CHCs) as a specific recognition cue. Most previous studies addressed the perception of CHCs occurring on the cuticle. However, the presence of CHCs in the environment (e.g., on the substrate) and the role of these compounds as a signal cue are less clear.In this study of the ant L. niger, CHCs were extracted from corpses of workers, from the legs of workers and from filter paper marked by workers. Behavioural choice tests were then performed to study the reactions of workers and queens to CHC passively deposited on the substrate by walking workers.A GC–MS analysis of the CHCs revealed a complex mixture of 28 peaks composed of linear alkanes, alkenes and mono-, di- or trimethylalkane. This mixture was consistently present in the three sources tested (corpses of workers, the tarsus of workers and paper marked by walking workers). Nevertheless, discriminant analyses distinguished clearly among the three types of profiles.The results of our behavioural experiments showed that the workers were able to distinguish between marked and unmarked papers but that they exhibited no preferences for different types of marked papers (papers marked by workers from their own colony or papers marked by workers from a different colony).In queens, the perception of marked papers depended on age or physiological state. Just-mated queens did not discriminate between unmarked papers and papers marked by an alien colony. In contrast, the egg-laying queens 3days after mating significantly favoured the marked papers. We discuss the ecological relevance of the differences in decision-making between castes.

Does lack of intraspecific aggression or absence of nymphs determine acceptance of foreign reproductives in Macrotermes?

The rejection or acceptance of a foreign reproductive by an alien colony may not always be as straightforward as cue recognition between worker termites. This paper aims to determine whether adoption of foreign reproductives is caused simply by lack of intraspecific aggression or is contingent on the reproductive status of the host colony. In the fungus-culturing termites, Macrotermes gilvus (Hagen) and Macrotermes carbonarius (Hagen), major workers showed low intraspecific aggression towards non-nestmates irrespective of geographic distance between source colonies. Our results indicated that workers were hardly aggressive towards non-nestmates. In royal cell-swapping experiments, both species responded in a similar way: (1) in host colonies with nymphs present, the foreign reproductives were rejected; while (2) in host colonies without nymphs the foreign reproductives were either accepted and breeding resumed or the host colonies died eventually. Workers from the host colonies preferentially maintained offspring nymphs from which adultoid replacement reproductives develop rather than accepting foreign reproductives. There is no fitness gain for the queenless workers in accepting foreign reproductives; however, there is overall benefit to the newly born population.

The acceptance rate of young wasps by alien colonies depends on colony developmental stages in the swarm-founding wasp, Polybia paulista von ihering (Hymenoptera: Vespidae)

In social insects, newly emerged individuals learn the colony-specific chemical label from their natal comb shortly after their emergence. These labels help to identify each individual's colony of origin and are used as a recognition template against which individuals can discriminate nestmates from non-nestmates. Our previous studies with Polybia paulista von Ihering support this general pattern, and the acceptance rate of young female and male wasps decreased as a function of their age. Our study also showed in P. paulista that more than 90% of newly emerged female wasps might be accepted by conspecific unrelated colonies. However, it has not been investigated whether the acceptance rate of newly emerged female wasps depends on colony developmental stage of recipient colonies. We introduced newly emerged female wasps of P. paulista into different colony developmental stags of recipient colonies, i.e., worker-producing and male-producing colonies. We found that the acceptance rate of newly emerged female wasps by alien colonies was pretty lower by male-producing colonies than worker-producing colonies. This is the first study to show that the acceptance rate of young female wasps depends on stages of recipient colonies.

The defensive behaviour of two Formica slave‐ant species: coevolutive implications with their parasite Polyergus rufescens (Hymenoptera, Formicidae)†

The cooperative defence in ant societies is a suitable system that can keep the integrity of the colonies from threats represented by competitors, predators and parasites. This is due to the presence of a well‐organized alarm communication system that induces defensive behaviours. Alarm reaction of Formica cunicularia and F. rufibarbis ant species was investigated by means of laboratory tests presenting different kinds of threatening stimuli (ants from alien colonies, homospecific and heterospecific) to colony fragments. Results show that both species react to a potential threat to the colony with an aggressive alarm instead of a panic alarm, characterized by a prompt exit of workers from the nest, attraction towards the stimulus and aggressive behaviours. Moreover they seem to feature the phenomenon of the ‘Enemy Specification’ towards their obligate parasite Polyergus rufescens. This could be considered as a result of a reciprocal adaptation between host and parasite. Furthermore, the comparison between the alarm reaction of the two studied species shows that F. rufibarbis seems to be more aggressive than F. cunicularia towards the parasite; this result could add a further clarification about the preference of the parasite towards F. cunicularia as host species.

Nestmate recognition in a Neotropical swarm-founding wasp: no effect of seasonality on tolerance of alien conspecifics

Previous study revealed that the swarm-founding wasp Polybia paulista is accurately able to distinguish nestmates from non-nestmates in the summer. However, the risk of accepting alien intruders is considered to be low in winter colonies, and additionally brood production is limited in 30- 40% of colonies during the winter in this species. Thus, it is expected that colonies might lower their acceptance threshold and accept some conspecific wasps from alien colonies in winter. We conducted field experiments to examine tolerance of conspecific (nestmate and non-nestmate) females in winter. In contrast to our prediction, our colonies did not accept any individuals from alien colonies. We suggest that P. paulista exhibits the colony-specific acceptance threshold in winter, and colonies that produced brood in their nests may have raised the acceptance threshold even if the risk of accepting alien intruders is low in winter.

Nest volatiles as modulators of nestmate recognition in the ant Camponotus fellah

When ants from alien colonies encounter each other the stereotypic reaction is usually one of aggressive behavior. It has been shown that factors such as queen-derived cues or nest-odors modulate this reaction. Here we examined whether nest volatiles affect nestmate recognition by observing the reaction of nestmates towards individual workers under one of four regimes: completely isolated; isolated but receiving a constant airflow from the mother colony; as previous but with the passage of nest volatiles towards the isolated ants blocked by adsorption on a SuperQ column; or reversed airflow direction-from the isolated ants to the nest interior.Ants that had been completely isolated for three weeks were subjected to aggressive behavior, but not those that had continued to receive airflow from the mother colony. Adsorbing the nest volatiles from the airflow by SuperQ abolished this difference, with these ants now also being subjected to aggression, indicating that nest volatiles can modulate nestmate recognition. Reverse airflow also reduced the level of aggression but to a lesser extent than airflow directed from the mother colony. In queenless colonies the overall aggression was reduced under all regimes, and there was no effect of flow, suggesting that the volatiles involved are queen-borne. The SuperQ adsorbed volatiles originated from Dufour's gland secretions of both workers and queen, implicating them in the process. Cuticular hydrocarbon profiles were not affected by exposure to nest volatiles, suggesting that the latter either constitute part of the recognition cues or affect worker behavior via a different, as yet elusive mechanism.

Effect of a foreign male’s age on acceptance into a conspecific colony in a Neotropical wasp

AbstractWe conducted field experiments to examine whether young males of different age cohorts were accepted from alien colonies in the swarm‐founding wasp Polybia paulista. We showed, as for young females in other social wasp species, that newly emerged males were frequently accepted from alien colonies. Our study suggests that young males of P. paulista acquire colony‐specific chemical odors shortly after emergence.

“Little Alien Colonies”: Representations of Immigrants and Their Neighborhoods in Social Work Discourse, 1875–1924

The “vexed problem of immigration,” as Jane Addams (1909, 214) termed it, became a central issue for the emerging social work profession. Through a close reading of social work’s public documents in the period from 1875 to 1924, this article analyzes the complex discursive mechanisms by which early social workers made immigrants and their environments legible as objects of intervention and advocacy. In particular, it examines the parallel social and moral calculi employed in social work’s analysis of immigrants and their “little alien colonies” (e.g., Greenleigh 1941, 208). Despite good intentions, social workers often viewed immigrants as dependent, abject, and exotic subjects; at a deeper register than their expressed interest in “the needs and possibilities of the immigrant” (Abbott 1917, 282), social work representations underscored and supported the problematization of immigrants in the public discourse. Contemporary social work faces many of the same dilemmas in fashioning a response to immigration.

Preferences and differences in the trail pheromone of the leaf-cutting ant Atta sexdens sexdens (Hymenoptera: Formicidae)

The amount of the trail pheromone substance, 3-ethyl-2,5-dimethylpyrazine, of the leaf-cutting ant Atta sexdens sexdens (Hymenoptera: Formicidae) increases with increasing size of the ant from 0 to 35 ng per individual. The compounds 2,5-dimethylpyrazine and trimethylpyrazine are also consistently present. Trails of venom gland extracts are followed with unequal fidelity by different castes. Minor workers follow best, mediums least well. Workers walk sinuously on narrow trails and less sinu- ously on wider trails up to 9 mm. Trails wider than 9 mm are not followed. The optimum concentration of pheromone on synthetic trails lies between 15 and 150 pg cm -1 . Given a choice of concentrations at a trail branch, workers always choose the more concen- trated. Workers showed slight preference for a trail made with venom gland secretion from their own, over that from an alien colony, but there is no preference for a trail that contains Dufour gland secretion additionally.

Behavioural and chemical studies of discrimination processes in the leaf-cutting ant Acromyrmex laticeps nigrosetosus (Forel, 1908)

Leaf-cutting ants live in symbiosis with a basidiomycete fungus that is exploited as a source of nutrients for ant larvae. Tests of brood transport revealed that Acromyrmex laticeps nigrosetosus workers did not discriminate a concolonial brood from an alien brood. The same result was observed with tests of fungus transport. Adult workers showed no aggressive behaviour to workers from other alien colonies (non-nestmates). There was no qualitative variation in the chemical profiles of larvae, pupae and adult workers from the different colonies. However, quantitative differences were observed between the different colonies. Hypotheses about the lack of intraspecific aggression in this subspecies of ants are discussed.

Comparison of Genetic Diversities in Native and Alien Populations of Hoary Mustard (Hirschfeldia incana[L.] Lagreze‐Fossat)

Increased selfing and inbreeding and, consequently, depauperate genetic diversities are commonly expected for alien colonies. We compared RAPDs data for native (southern Europe) and alien (British Isles) populations of hoary mustard (Hirschfeldia incana). This species is normally out‐breeding, but it is capable of self‐fertilization. Contrary to the common expectations, genetic diversities in native and alien populations were similar, without any strong evidence of decreased levels of genetic diversities in alien populations. A variety of factors may have contributed to this observation, including high variation in founding groups, founders originating from multiple H. incana source populations, and high rates of past and/or current gene flow. A review of other studies showed that this pattern of similar genetic diversities in native and alien populations was not unusual but has been regularly observed in other invasive plant species.

Social wasp parasites affect the nestmate recognition abilities of their hosts ( Polistes atrimandibularis and P. biglumis , Hymenoptera, Vespidae)

Queens of the parasitic social wasp, Polistes atrimandibularis, temporarily mimic the odor of their host species, Polistes biglumis, but their offspring have parasite-specific odors. As a consequence, in parasitized colonies individuals with different odors co-inhabit the colony and host workers, who are responsible for colony defense, accept wasps with different odors. In order to verify whether this particular condition causes a change in recognition abilities of hosts, we tested nestmate/non-nestmate discrimination in field colonies invaded by social parasites (and in non-parasitized colonies as controls). Results show that parasitized colonies distinguish between nestmates and non-nestmates, distinguish their parasite queen from those that usurped alien colonies, and accept their parasite's non-mimetic offspring but make more recognition errors than non-parasitized colonies. The optimal acceptance threshold model predicts that when the frequency of encountering non-kin increases, residents become less permissive towards intruders. However, my data show that parasitized colonies are more permissive towards non-nestmates with respect to non-parasitized colonies but they are also more aggressive towards nestmates, suggesting that host workers' learning abilities are impaired.

Collective exploration and area marking in the ant Lasius niger

The aphid tending ant species, Lasius niger collectively explore new areas. Scouts are mobilised within the first five minutes of exploration with activity thereafter decreasing and stabilising after 40 minutes. Collective path choices of colonies underline the existence of chemical recruitment and area marking during exploration. To identify the territorial or home-range role of exploratory marking, we examined its influence on agonistic behaviour during intraspecific encounters. Ten nestmates and ten alien conspecifics were confronted: (i) on a paper marked by their own colony, (ii) on a paper marked by the alien colony or (iii) on a paper not chemically marked by ants. Residents and intruders showed a weak level of aggression and behaved similarly whatever the type of area marking. As no colony-specificity was observed in the odds of eliciting aggression, exploratory area marking in Lasius niger seems to be rather a home-range than a true territorial marking. We discuss the role of collective exploration, home-range marking and intraspecific interactions in the structuring and use of foraging space by neighbour Lasius niger colonies.

Chemical trails used for orientation in nest cavities by two vespine wasps, Vespa crabro and Vespula vulgaris

Foragers of the two cavity-nesting vespines, Vespa crabro and Vespula vulgaris, lay and follow chemical trails in their nest cavities. In our experiments 75% of V. crabro and 72% of V. vulgaris foraging workers followed a trail arranged in a direction to which they were not accustomed. In experiments with trails taken in from conspecific alien colonies, 21% of V. crabro foragers and 65% of V. vulgaris foragers followed the trails. These behavioural responses indicate that the trail of V. crabro is colony-specific whereas that of V. vulgaris is not.

Oxidative stress induces cytotoxicity during rejection reaction in the compound ascidian Botryllus schlosseri

When genetically incompatible colonies of the compound ascidian Botryllus schlosseri contact each other, a rejection reaction occurs, characterised by the appearance of cytotoxic foci along the touching borders. In the course of this reaction, morula cells, a common haemocyte-type in ascidians, release their vacuolar content, mainly phenoloxidase and its polyphenol substrata, upon the recognition of soluble factors diffusing from the alien colony through the partially fused tunic. In a previous paper, we demonstrated the relationship between phenoloxidase and cytotoxicity. Here, we investigated the effects of superoxide dismutase, catalase and sorbitol (scavengers of superoxide anions, peroxides and hydroxyl radicals, respectively) on the cytotoxicity observed in haemocyte cultures incubated with heterologous blood plasma. Although the above compounds have no effects on morula cell degranulation and phenoloxidase activity, they suppress cell death, suggesting that oxidative stress plays a key role in in vitro cytotoxicity. In addition, sorbitol reduces the extent of the cytotoxicity occurring in the rejection reaction between incompatible colonies, which stresses the important role of hydroxyl radicals in this process. The observation of a decrease in total and non-protein thiols in haemocytes previously incubated with heterologous blood plasma fits the hypothesis of oxidative stress as the main cause of phenoloxidase-related cytotoxicity.