Blending of heritable recognition cues among ant nestmates creates distinct colony gestalt odours but prevents within‐colony nepotism

The evolution of sociality is facilitated by the recognition of close kin, but if kin recognition is too accurate, nepotistic behaviour within societies can dissolve social cohesion. In social insects, cuticular hydrocarbons act as nestmate recognition cues and are usually mixed among colony members to create a Gestalt odour. Although earlier studies have established that hydrocarbon profiles are influenced by heritable factors, transfer among nestmates and additional environmental factors, no studies have quantified these relative contributions for separate compounds. Here, we use the ant Formica rufibarbis in a cross‐fostering design to test the degree to which hydrocarbons are heritably synthesized by young workers and transferred by their foster workers. Bioassays show that nestmate recognition has a significant heritable component. Multivariate quantitative analyses based on 38 hydrocarbons reveal that a subset of branched alkanes are heritably synthesized, but that these are also extensively transferred among nestmates. In contrast, especially linear alkanes are less heritable and little transferred; these are therefore unlikely to act as cues that allow within‐colony nepotistic discrimination or as nestmate recognition cues. These results indicate that heritable compounds are suitable for establishing a genetic Gestalt for efficient nestmate recognition, but that recognition cues within colonies are insufficiently distinct to allow nepotistic kin discrimination.     

Early learning of volatile chemical cues leads to interspecific recognition between two ant species

Nestmate recognition in social insects generally involves matching a label to the template that is acquired through the early learning of non-volatile cuticular hydrocarbon cues. However, a possible role of the volatile chemical cues that exist in the nest, and which may also affect template formation, has not been studied. We investigated this possibility using experimental mixedspecies groups composed of the two ant species Manica rubida and Formica selysi. The experimental set-up either allowed full contact between workers of the two species or interspecific contact was hindered or prohibited by a single or a double mesh. After three months, workers of M. rubida ants were selected as focal ants for aggression tests including the following target ants: F. selysi workers from the same mixed-species group (for each of the three rearing conditions) or from a single-species group (control). Workers of M. rubida were always amicable towards their group-mates, irrespective of the experimental group (contact, single or double mesh). However, M. rubida that were not imprinted on F. selysi, expressed high levels of aggression towards the non-familiar F. selysi workers. The finding that F. selysi workers in the mixed-species groups appeared familiar to their M. rubida group-mates even without physical contact between them, suggests that the volatile cues produced by F. selysi affected nestmate recognition in M. rubida. In an attempt to identify these volatile cues we performed SPME analysis of the head space over groups of F. selysi workers. The findings revealed that F. selysi Dufour’s gland constituents, with undecane as the major product, are released into the head space, rendering them likely candidates to affect template formation in M. rubida. Analysis of Dufour’s gland secretion of F. selysi revealed a series of volatile alkanes, including undecane as a major product. These alkanes were not present in the glandular secretion of M. rubida, whose secretion was mainly composed of isomers of farnesene. We therefore hypothesize that callow M. rubida workers in the mixed-species groups had become imprinted by the above alkanes (in particular undecane, being the major heterospecific volatile in the head space) and incorporated them into their own template. Received 18 October 2007; revised 2 January 2008; accepted 7 January 2008.     

The scent of supercolonies: the discovery, synthesis and behavioural verification of ant colony recognition cues

Background  

Ants form highly social and cooperative colonies that compete, and often fight, against other such colonies, both intra- and interspecifically. Some invasive ants take sociality to an extreme, forming geographically massive 'supercolonies' across thousands of kilometres. The success of social insects generally, as well as invasive ants in particular, stems from the sophisticated mechanisms used to accurately and precisely distinguish colonymates from non-colonymates. Surprisingly, however, the specific chemicals used for this recognition are virtually undescribed.     

戈壁生态系统蚁穴微生境对大型土壤动物多样性的影响

收获蚁蚁穴是戈壁生态系统中重要的微生境, 它通过汇集凋落物和改善土壤环境强烈影响动植物的分布及多样性。鉴于此, 本文选择戈壁荒漠收获蚁(Messor desertus)蚁穴为研究对象, 于2020年5月、6月和10月利用陷阱法调查了蚁穴及毗邻裸地大型土壤动物的种类组成及数量变化, 并分析其影响要素。结果表明: (1) 10月, 蚁穴和裸地间大型土壤动物群落组成存在显著差异, 而5月和6月二者间相差较小, 10月(62.9%)蚁穴和裸地大型土壤动物的平均相异性高于5月(34.8%)和6月(39.3%); (2) 5月, 蚁穴大型土壤动物均匀度指数显著低于裸地, 6月, 蚁穴大型土壤动物活动密度和类群丰富度均显著高于裸地, 10月, 蚁穴大型土壤动物类群丰富度和多样性指数均显著高于裸地; (3)荒漠收获蚁蚁穴显著提高了其他食性土壤动物的活动密度及类群丰富度, 还增强了捕食性和非捕食性土壤动物的相互作用关系并改变了荒漠收获蚁与蚁穴大型土壤动物类群的种间相互作用关系; (4) pRDA排序结果表明, 土壤电导率、全氮和粉粒含量是影响蚁穴和裸地大型土壤动物分布的主要土壤因子。总之, 戈壁生态系统荒漠收获蚁蚁穴微生境提高了大型土壤动物多样性, 改变了大型土壤动物类群间的营养和非营养关系, 这会影响大型土壤动物的营养结构及其功能。     

Recruitment in starved nests: the role of direct and indirect interactions between scouts and nestmates in the ant <Emphasis Type=Italic>Lasius niger</Emphasis>

In social insects, the foraging activity usually increases with the length of food deprivation. In Lasius niger, a mass-recruiting ant species, the foraging adjustment to the level of food deprivation is regulated by the scout that fed at the food source and by the response of the nestmates to signals performed by the scout inside the nest. In this study, we look at the role of these direct interactions (antennations or trophallaxes) and indirect interactions (pheromonal emission) and how they are influenced by the level of food deprivation. At the beginning of recruitment, the relative number of nestmates leaving the nest to forage increases with the level of deprivation. The nestmates’ propensity to exit the nest is not influenced by a previous trophallactic and/or antennal contact with a scout. Our results strongly suggest that the exit of nestmates is triggered by a chemical signal emitted by a scout. Deprivation lowers the response threshold of nestmates to this chemical signal resulting in a more important exit from the nest. Surprisingly, 27% of starved nestmates that receive food from the scout relay the information by depositing a chemical signal before having discovered and drunk the food source. Both phenomena boost the recruitment process. Though successful foragers returning to the nest have a significant role in the recruitment to the food source, we observed that the response of the nestmates inside the nest also greatly influence regulation of the foraging activity.     

Mate preference for multiple cues: interplay between male and nest size in the sand goby, Pomatoschistus minutus

Female mating preferences are often based on more than one cue.In empirical studies, however, different mate choice cues aretypically treated separately ignoring their possible interactions.In the current work, we studied how male body size and sizeof the male's nest jointly affect mate preferences of femalesand gobies, Pomatoschistus minutus. The females were givena binary choice between males that differed either in body sizeor size of their nest or both. We found that neither body sizenor size of the nest alone affected male attractiveness, buttogether these 2 cues had a significant effect. Specifically,large males were more popular among females when they had alarge nest than when they occupied a small nest. The resultssuggest that if interaction effects between multiple mate choicecues are not considered, there is a danger of ignoring or underestimatingthe importance of these cues in sexual selection by female choice.     

Sources of nestmate recognition cues in the imported fire ant Solenopsis invicta buren (Hymenoptera: Formicidae)

Aggression bioassays were conducted to investigate nestmate (worker-worker) recognition in monogyne colonies of the imported fire ant. Environmentally correlated cues dominated the recognition cue hierarchy of laboratory and field colonies, anddiet alone significantly modified recognition labels and templates of laboratory-reared workers. ‘Discriminators’ associated with worker genotype also affected recognition, but ‘queen discriminators’ did not significantly affect either labels or templates of laboratoryreared workers exposed as adults to alien queens for 28 days. Factors contributing to the importance of environmentally derived recognition cues in this species and the potential implications of such cues for the formation of polygynous fire ant populations are discussed.     

A meta‐analysis of leaf‐cutting ant nest effects on soil fertility and plant performance

1. Leaf‐cutting ants (LCAs) are considered as one of the most important agents of soil disturbances that affect vegetation patterns, but these assertions are based on isolated studies or anecdotal data. In this study, meta‐analysis techniques were used to quantitatively analyse the generality of these effects and determine some of their sources of variation. 2. The results reveal the following: (i) LCA nest sites showed higher levels of soil fertility than control sites, but the key source of these nutrients is the refuse material rather than the nest soil itself; (ii) refuse material from external piles tended to be richer in nutrient content than refuse material from internal refuse chambers; (iii) nest sites from temperate habitats showed higher cation content than those located in tropical/subtropical habitats; and (iv) nest sites showed higher plant growth than adjacent non‐nest sites (especially if plants have access to the refuse) but similar plant density and plant richness. 3. As LCAs improve nutrient availability in nest sites through the accumulation of refuse material, the location of the refuse will have a relevant role affecting vegetation. LCA species with external refuse dumps could benefit herbs, early vegetation stages and short‐living plants, whereas those with internal refuse chambers could benefit long‐living, large trees. However, the positive effect on individual plants does not extend to population and community levels. The foraging preferences of ants and the changes in microclimatic conditions around nests could act as selective ecological filters. 4. As refuse material from external piles and nest sites in temperate habitats tend to show higher fertility than refuse material from internal nest chambers and nest sites in tropical/subtropical habitats, LCA species with external refuse dumps in temperate regions could be of particular relevance for nutrient cycling and vegetation patterns.     

红火蚁蚁巢表层各虫态数量与蚁丘大小的关系

采用挖巢取样法调查广州市番禺区6个地点的红火蚁蚁巢表层各虫态数量,分析各虫态数量与蚁丘大小的相互关系。结果表明,在调查的180个蚁巢中,占蚁巢总数57.78%的红火蚁蚁丘体积5000 cm~3,占蚁巢总数20.56%的红火蚁蚁丘体积介于5000~10000 cm~3,占蚁巢总数21.67%的红火蚁蚁丘体积10000 cm~3;占蚁巢总数32.78%的红火蚁蚁巢蚁丘表面基部面积500 cm~2,占蚁巢总数42.78%的红火蚁蚁巢蚁丘表面基部面积在500~1000 cm~2,占蚁巢总数24.44%的红火蚁蚁巢蚁丘表面基部面积1000 cm~2。平均1cm~3有效蚁丘体积中有3头红火蚁,1 cm~2蚁丘表面基部面积有22头红火蚁。红火蚁各虫态数量和蚁群总数均明显地随着蚁丘体积和表面基部面积的增大而增大,工蚁数量与蚁丘体积及表面基部面积的正相关性较其他虫态强;红火蚁数量与蚁丘体积的相关性强于该蚁数量与蚁丘表面基部面积的相关性。     

红火蚁巢穴土壤中生物碱成分分析

【目的】红火蚁Solenopsis invicta栖息在真菌和细菌较丰富的土壤环境中,容易受多种真菌和细菌性病原体的侵染,因红火蚁的毒液具有很好的防御和抗菌作用,致使红火蚁能生存于此类土壤环境中。为了探索这一奥秘,本研究旨在建立红火蚁巢穴土壤中的毒液生物碱的最佳提取方法,并对毒液生物碱成分进行定量分析。【方法】采集蚁巢周边3 m处的土壤,进行添加、回收红火蚁毒液实验。采用抽滤法提取土壤中的毒液生物碱,利用GC-FID对毒液生物碱成分进行定量分析,筛选最佳的提取溶剂,并且确定添加三乙胺的最佳体积比。然后用最优方法提取蚁巢土壤中的毒液生物碱,并进行定量分析。【结果】正己烷、二氯甲烷、乙酸乙酯、丙酮和甲醇5种提取溶剂中,正己烷处理的生物碱回收率略优。当添加的三乙胺的体积为1 mL及以上时,提取效果最佳。红火蚁巢穴土壤中的毒液生物碱成分中,trans-C15∶1的含量最高,trans-C13∶1的含量次之。巢穴土壤中总生物碱含量约为22μg/g。【结论】三乙胺有助于提取蚂蚁巢穴土壤中的毒液生物碱。红火蚁巢穴土壤中的生物碱浓度较高,有可能对巢穴土壤微生物群落产生重要影响。     

The nest architecture of the ant, Pheidole oxyops Forel, 1908 （Hymenoptera： Formicidae）

Pheidole oxyops builds subterranean nests, with an external architecture that is distinctive and easily recognizable by its wide and specific entrance hole, measuring up to 12.2 cm in diameter, denoting a pitfall-trap. In order to study the nests＇ internal architecture, seven nests were excavated; four were identified with neutral talc, while the others were cast in cement and then excavated. Measurements were made in order to gain a better understand- ing of their structures, and a photographic documentation was obtained as well. The excavations revealed that the nests are perpendicular relative to the ground, beginning with a cylindrical channel with a mean length of 13.5 cm, containing irregular formations, and whose diameter becomes progressively narrower until the first chamber is formed. As the channel continues, dish-like chambers appear, interconnected by channels that become progressively narrower and longer, while the chambers are arranged at greater distances from each other as nest depth increases. Both channels and chambers are located on the vertical projection of the entrance hole. Nests may reach a depth of up to 5.09 m, with a number of chambers ranging between 4 and 14.     

Of mice and mallards: positive indirect effects of coexisting prey on waterfowl nest success

Coexisting prey species interact indirectly via their shared predators when one prey type influences predation rates of the second prey type. In a temperate system where the predominant shared predator is a generalist, I studied the indirect effects of rodent populations on waterfowl nest success, both within the nesting season among sites and among years. Among six to ten upland fields (14 to 27 ha), mallard ( Anas platyrhynchos ) nest success was positively correlated with rodent abundance in all three years of the study. After removing year effects, mallard nest success remained positively correlated with the relative abundance of rodents. Of the rodent species present, California voles ( Microtus californicus ) were the most important coexisting prey type influencing nest success. Among years, mallard nest success was positively correlated with vole abundance; the asymptotic relationship suggests a threshold response to vole abundance, beyond which predators become satiated and additional voles do little to affect nest success. I tested and rejected three alternative explanations for the observed positive correlation between mallard nest success and rodent abundance that do not involve an indirect effect of coexisting prey populations. The influences of dense nesting cover, nesting density, and predator activity did not explain the observed patterns of nest success. These results suggest that rodent populations buffer predation on waterfowl nests, both within and among years, via the behavioral responses of shared predators to coexisting prey.     

Cheating on a mutualism: indirect benefits of ant attendance to a coccidophagous coccinellid

Coccinellids (Coleoptera: Coccinellidae) are generally unable to prey on ant-tended prey. However, particular coccinellid species have morphological, behavioral, or chemical characteristics that render them immune to ant attacks, and some species are even restricted to ant-tending areas. The benefit gained from living in close association with ants can be twofold: (1) gaining access to high-density prey areas and (2) gaining enemy-free space. Here, the myrmecophily of Azya orbigera Mulsant (Coleoptera: Coccinellidae), an important predator of the green coffee scale, Coccus viridis (Green) (Hemiptera: Coccidae), is reported. In this paper, three main questions were studied. (1) Are the waxy filaments of A. orbigera larvae effective as defense against attacks of the mutualistic ant partner of C. viridis, Azteca instabilis F. Smith (Hymenoptera: Formicidae)? (2) Does A. instabilis reduce the rate at which A. orbigera larvae prey on scales? (3) Do A. orbigera larvae gain enemy-free space by living in close association with A. instabilis? Laboratory and field experiments were conducted to answer these questions. We found that, because of the sticky waxy filaments of A. orbigera larvae, A. instabilis is incapable of effectively attacking them and, therefore, the predation rate of A. orbigera on C. viridis does not decrease in the presence of ants. Furthermore, A. instabilis showed aggressive behavior toward A. orbigera's parasitoids, and the presence of ants reduced the parasitism suffered by A. orbigera. This is the first time that this kind of indirect positive effect is reported for an ant and a coccidophagous coccinellid. Furthermore, this indirect positive effect may be key to the persistence of A. orbigera's populations.     

Predation of the oriental fruit fly,Bactrocera dorsalis puparia by the red imported fire ant,Solenopsis invicta: role of host olfactory cues and soil depth

Predation by red imported fire ants, Solenopsis invicta on oriental fruit fly, Bactrocera dorsalis puparia was evaluated. No significant olfactory response of the workers was observed at 0, 2 and 4 days after fly pupation, whereas the workers were significantly attracted by the 6th day old puparia. We found S. invicta that predated on puparia of B. dorsalis in the field. The predation rate was negatively correlated with the depth of puparia in the soil. The predation rate was 70% at 4 cm depth; whereas, zero predation rate was observed at 6 cm depth. The predation rate was also significantly affected by soil moisture. The predation rate was 66.5% and 72.1% at soil moisture values of 40% and 80%, respectively, and no predation occurred at soil moisture value of 0%.     

Below-ground nitrogen transfer between different grassland species: Direct quantification by 15N leaf feeding compared with indirect dilution of soil 15N

Nitrogen (N) transfer from one species to another is important for the N cycling in low-input grassland. In the present work, estimates obtained by an indirect ¹⁵N dilution technique were compared with estimates obtained by a direct ¹⁵N leaf feeding technique over two complete growing seasons in red clover-ryegrass and white clover-ryegrass mixtures under field conditions. The direct technique confirmed that N transfer between clovers and ryegrass is a bi-directional process. The transfer of N from both clovers to ryegrass occurred within 25 days upon the first labelling event. A very high N transfer occurred from white clover to the associated ryegrass, 4.5 and 7.5 g m⁻² in the 1st and 2nd production year, respectively. The corresponding values for transfer from red clover to the associated ryegrass were 1.7 and 3.6 g m⁻². Quantified relatively to the total above-ground N content of white clover- ryegrass and red clover-ryegrass mixtures, the N transfer exceeded 50% and 10%, respectively, in three out of seven harvests. The N transfer from ¹⁵N labelled grass to associated clovers constituted a relatively constant proportion of approx. 8% of the above-ground N content of the mixtures. Estimates based on the soil ¹⁵N dilution technique generally underestimated the net N transfer by more than 50% compared to the direct ¹⁵N labelling technique. Furthermore, the indirect ¹⁵N dilution technique estimated only marginal differences between red and white clover in the quantities of N transferred, whereas the direct ¹⁵N labelling technique showed the N transfer from white clover to the associated ryegrass to be significantly higher than that involving red clover. It is concluded that N transfer is a much more dynamic and quantitatively important process in grassland than previously recognised. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effect of aromatic cedar mulch on Argentine ant (Hymenoptera: Formicidae) foraging activity and nest establishment

In the laboratory, Argentine ant, Linepithema humile (Mayr), mortality was positively correlated to the length of an aromatic cedar mulch section that had to be crossed before food could be reached. When ants could access food without crossing the mulch, mortality was not correlated to mulch section length. In the field, Argentine ants showed a tendency to avoid aromatic cedar mulch as a nesting substrate. In plant beds alongside buildings the number of ant nests (pockets containing brood) found was not significantly different between aromatic cedar and cypress mulch. However, when pine straw mulch around oak trees was replaced with aromatic cedar or cypress mulch, a similar number of ant nests was found in the cypress mulch as in the original pine straw, whereas numbers in aromatic cedar mulch were significantly lower. Also, fewer ants were trailing on the trees surrounded by aromatic cedar mulch compared with cypress mulch or the original pine straw. The number of ants attracted to apple jelly baits placed alongside the buildings did not differ between mulch types; neither did the number of ant trails crossing the mulch beds around the buildings. We suggest that aromatic cedar mulch may help control Argentine ants and reduce insecticide input when applied in combination with conventional control methods.     

Between a rock and a hard place: impact of nest selection behavior on the altitudinal range of an alpine ant, Formica neorufibarbis

Formica neorufibarbis Emery is a dominant alpine ant in North America. Above timberline, colonies nest under rocks, placing brood against the rock surface to warm during the day. We tested the hypothesis that nest rock preference represents a mechanism of behavioral thermoregulation and used a demographic model to explore its impact on the species' altitudinal range. Under sunny conditions rocks provide warm microclimates. Across a 400-m altitudinal gradient from below timberline to high alpine tundra, nest rocks warmed on average 5°C above ambient at mid-day, but cooled to near ambient before dawn. Soil was warmer at mid-day under rocks occupied by ants than under randomly chosen unoccupied rocks in the same area. Occupied nest rocks were painted white or black to manipulate temperature and test its impact on colony occupancy. In addition, black-painted and white-painted rocks of similar size were placed in the field to test for a temperature effect on nest site colonization. Likelihood of nest persistence and colonization decreased under cooler, white-painted rocks and depended strongly on rock warming capacity. Experimental results showed that nest site colonization and occupancy are favored in microsites maintaining mid-day temperature ≥22°C. Across the altitudinal gradient, 66-90% of nest microsites occupied by F. neorufibarbis met this temperature threshold. Conversely, none of the exposed microsites and only 15% of those microsites under randomly selected rocks warmed to 22°C at the high altitude site (3,900 m). Results support the conclusion that nest site preference contributes to the broad altitudinal distribution of F. neorufibarbis.     

Waste of leaf-cutting ants: disposal,nest structure,and abiotic soil factors around internal waste chambers

Leaf-cutting ants produce large quantities of waste that harbor bacteria and fungi that are harmful to the colony. To be protected from these pathogens, the workers of Atta species present a sophisticated organization to manage harmful material, which can be deposited outside the nest or in internal chambers. However, little is known about the behavior of Acromyrmex species in handling and disposal of waste. Due to some observations, we assume that the same species of Acromyrmex can deposit waste outside the nest and into internal chambers and raise the following question: what determines the occurrence of internal waste chambers in Acromyrmex? To address this question, we verified whether nest depth influences the waste-chamber occurrence. We also verified the nest structure and the abiotic factors of soil beside each waste-chamber: pH and water content of the soil. For this, eight nests were excavated for Acromyrmex balzani and Acromyrmex rugosus rugosus. We verified that not only can the same leaf-cutting ant species deposit debris both outside and inside the nest but also the same nest can present internal chambers and external waste deposit. The soil beside the waste chamber always presented an acidic pH, while the humidity varied widely. Our results showed that the nest depth was highly correlated with the depth of the waste chamber (p = 0.0003) and probably has some influence on waste disposal. The characteristics of the nest and the role of depth in the choice of waste chamber location are discussed.     

蚂蚁扰动对土壤有机碳循环过程的影响研究进展

蚂蚁作为生态系统的消费者和分解者,其对土壤有机碳库的影响一直是学术界研究的热点。目前研究主要从蚂蚁对土壤宏量元素储量、理化性质、微生物群落活动等方面探究蚂蚁对土壤有机碳库的影响。本文综述了蚂蚁扰动对土壤有机碳循环过程特征的影响。蚂蚁筑巢改变了蚁穴土壤的微生境、微气候与土壤理化性质,并通过重构土壤微生物群落结构特征、调控地表植被演替过程与格局等方式,直接或间接的影响蚁巢中土壤有机碳来源、碳库分配过程、有机碳库稳定性、有机质微观分子特征等,在微域、局地乃至景观尺度上影响土壤有机碳的循环过程。未来研究应着重从量化蚂蚁扰动及其导致的环境因子波动对土壤碳通量变化的贡献、建立定量模型联系并统一蚂蚁影响下土壤碳循环过程、厘清蚂蚁影响土壤有机碳库稳定性的机制等方面开展深入研究,揭示蚂蚁作为“生态工程师”在调控土壤碳循环过程中的作用机制。