The influence of brood type on activity cycles inLeptothorax allardycei (Hymenoptera: Faruicidae)

We investigated the effect that different categories of brood had on the production of periodic activity within artificial aggregates of the ant,Leptothorax allardycei. Colonies of this species exhibit periodic patterns of movement activity with a period of approximately 20–30 min. Artificial aggregates of this species show that periodic activity appears gradually as the number of workers in the aggregate increases to 15 workers. In addition, it has been noted that the presence of brood with the workers makes the periodic activity more pronounced. In this paper we investigate the effect that four categories of brood had on the periodic activity of worker ants. Eggs, small larvae, large larvae, and pupae were tested with four different-sized aggregates of workers. We hypothesized that brood care is responsible for the increase in periodic activity and therefore that larvae (which require more tending) would be more effective at increasing periodic activity than eggs or pupae. Contrary to our expectations, eggs and both size categories of larvae were equally effective in enhancing periodicity in our experimental aggregates. Pupae, in contrast, were completely ineffective at enhancing periodic activity. We discuss some possible reasons for the differential effects of eggs, larvae, and pupae on the behavior of worker ants.     

Aggressive interactions between the introduced Argentine ant, Linepithema humile and the native odorous house ant, Tapinoma sessile

The Argentine ant (Linepithema humile) is an invasive species that disrupts the balance of natural ecosystems by displacing indigenous ant species throughout its introduced range. The mechanisms by which Argentine ants effectively compete against native ant species have been previously addressed in field studies that centered on interference and exploitation competition at baits and mainly examined the colony-level performance of Argentine ants. Detailed behavioral observations explaining the basis for the strong competitive ability of L. humile are comparatively rare. To gain a better understanding of the mechanisms by which Argentine ants displace native ants we examined the aggressive interactions between the Argentine ants and the odorous house ant, Tapinoma sessile in four different aggression assays: (1) worker dyad interactions, (2) symmetrical group interactions, (3) intruder introductions into an established resident colony, and (4) a resource competition assay which focused on competition for food and nesting space. Our results demonstrate a clear disparity between worker-level and colony-level fighting ability of Argentine ants and provide behavioral evidence to explain the superior interference ability of Argentine ants in group assays. Argentine ants experienced mixed success in fighting against odorous house ants in dyad interactions, but gradually gained a numerical advantage in symmetrical group interactions by active cooperation among nestmates. Results of the resource competition assay indicate that Argentine ants recruit rapidly, numerically dominate food and nesting sites, and aggressively displace T. sessile from baits. Taken together, the results of these assays allow us to pinpoint the behavioral mechanisms responsible for the remarkable competitive ability of Argentine ants.     

Herbivory-induced extrafloral nectar increases native and invasive ant worker survival

Ascertaining the costs and benefits of mutualistic interactions is important for predicting their stability and effect on community dynamics. Despite widespread designation of the interaction between ants and extrafloral nectaries (EFNs) as a mutualism and over 100 years of studies on ant benefits to plants, the benefits to ants have never been experimentally quantified. The success of invasive ants is thought to be linked to the availability of carbohydrate-rich resources, though reports of invasive ant visits to EFNs are mixed. In two laboratory experiments, we compared worker survival of one native (Iridomyrmex chasei) and two invasive ant species (Linepithema humile and Pheidole megacephala) exposed to herbivorized or non-herbivorized EFN-bearing plants (Acacia saligna) or positive and negative controls. We found that non-herbivorized plants did not produce any measurable extrafloral nectar, and ants with access to non-herbivorized plants had the same survival as ants with access to an artificial plant and water (unfed ants). Ants given herbivorized plants had 7–11 times greater worker survival relative to unfed ants, but there were no differences in survival between native and invasive ants exposed to herbivorized plants. Our results reveal that ants cannot induce A. saligna extrafloral nectar production, but workers of both native and invasive ant species can benefit from extrafloral nectar as much as they benefit from sucrose.     

Timekeeping through social contacts: social synchronization of circadian locomotor activity rhythm in the carpenter ant Camponotus paria

In ant colonies a large proportion of individuals remain inside nests for most of their lives and come out only when necessary. It is not clear how, in a nest of several thousand individuals, information about local time is communicated among members of the colony. Central to this seem to be circadian clocks, which have an intrinsic ability to keep track of local time by entraining to environmental light-dark, temperature, and social cycles. Here, the authors report the results of their study aimed at understanding the role of cyclic social interactions in circadian timekeeping of a day-active species of carpenter ant Camponotus paria. The authors found that daily social interactions with visitors (worker ants) was able to synchronize the circadian locomotor activity rhythm of host worker ants and queens, in one-on-one (pair-wise) and multi-individual (group-wise) interactions. Interestingly, the outcome of cyclic social interactions was context specific; when visitor workers socially interacted with host workers one-on-one, host workers considered the time of interaction as subjective day, but when visitor workers interacted with a group of workers and queens, the hosts considered the time of interaction as subjective night. These results can be taken to suggest that members of the ant species C. paria keep track of local time by socially interacting with workers (foragers) who shuttle in and out of the colony in search of food. (Author correspondence: vsharma@jncasr.ac.in ).     

Respiration,worker body size,tempo and activity in whole colonies of ants

Ants are social, and their metabolism should be measured on at least two levels: (i) the individual workers and brood of which the colony is composed and (ii) the colony in its entirety. Whole colony respiration, tempo (size‐free running speed in body lengths per second) and whole colony activity were simultaneously measured for 15 species of ants in four subfamilies, and these data are related to average worker and whole‐colony weight, activity, percentage brood and percentage fat. Across all 15 species, whole colony respiration rate (μL CO₂ h^?1) is linearly related to whole colony live weight (log–log slope = 1.0). Colonies composed of large workers respire less than colonies composed of an equal live weight of small workers, and colonies with high tempos respire more than lower tempo colonies of equal weight. The tempos and respiration rates of smaller ants tend to be higher, and a higher tempo exacts a cost in higher respiration independent of the effect of small body size. Individual worker respiration (μL CO₂ h^?1) scales to worker live weight with an exponent of 0.76. Whole colony specific respiration rate (μL CO₂ g^?1 h^?1) is unrelated to colony live weight. The regressions of respiration rates against colony and worker dry weight, lean weight and metabolic weight have similar slopes to those of live weight but different intercepts. Respiration is not related to worker percentage fat, percentage brood or activity. Ant ecology, tempo, body size, polymorphism and colony size are discussed in relation to respiration.     

Reciprocally beneficial interactions between introduced plants and ants are induced by the presence of a third introduced species

Interspecific interactions play an important role in the success of introduced species. For example, the ‘enemy release’ hypothesis posits that introduced species become invasive because they escape top–down regulation by natural enemies while the ‘invasional meltdown’ hypothesis posits that invasions may be facilitated by synergistic interactions between introduced species. Here, we explore how facilitation and enemy release interact to moderate the potential effect of a large category of positive interactions – protection mutualisms. We use the interactions between an introduced plant (Japanese knotweed Fallopia japonica), an introduced herbivore (Japanese beetle Popillia japonica), an introduced ant (European red ant Myrmica rubra), and native ants and herbivores in riparian zones of the northeastern United States as a model system. Japanese knotweed produces sugary extrafloral nectar that is attractive to ants, and we show that both sugar reward production and ant attendance increase when plants experience a level of leaf damage that is typical in the plants’ native range. Using manipulative experiments at six sites, we demonstrate low levels of ant patrolling, little effect of ants on herbivory rates, and low herbivore pressure during midsummer. Herbivory rates and the capacity of ants to protect plants (as evidenced by effects of ant exclusion) increased significantly when plants were exposed to introduced Japanese beetles that attack plants in the late summer. Beetles were also associated with greater on‐plant foraging by ants, and among‐plant differences in ant‐foraging were correlated with the magnitude of damage inflicted on plants by the beetles. Last, we found that sites occupied by introduced M. rubra ants almost invariably included Japanese knotweed. Thus, underlying variation in the spatiotemporal distribution of the introduced herbivore influences the provision of benefits to the introduced plant and to the introduced ant. More specifically, the presence of the introduced herbivore converts an otherwise weak interaction between two introduced species into a reciprocally beneficial mutualism. Because the prospects for facilitation are linked to the prospects for enemy release in protection mutualisms, species introductions can have complex effects on existing species interactions, between both native and introduced species.     

Timekeeping Through Social Contacts: Social Synchronization of Circadian Locomotor Activity Rhythm in the Carpenter Ant Camponotus paria

In ant colonies a large proportion of individuals remain inside nests for most of their lives and come out only when necessary. It is not clear how, in a nest of several thousand individuals, information about local time is communicated among members of the colony. Central to this seem to be circadian clocks, which have an intrinsic ability to keep track of local time by entraining to environmental light-dark, temperature, and social cycles. Here, the authors report the results of their study aimed at understanding the role of cyclic social interactions in circadian timekeeping of a day-active species of carpenter ant Camponotus paria. The authors found that daily social interactions with visitors (worker ants) was able to synchronize the circadian locomotor activity rhythm of host worker ants and queens, in one-on-one (pair-wise) and multi-individual (group-wise) interactions. Interestingly, the outcome of cyclic social interactions was context specific; when visitor workers socially interacted with host workers one-on-one, host workers considered the time of interaction as subjective day, but when visitor workers interacted with a group of workers and queens, the hosts considered the time of interaction as subjective night. These results can be taken to suggest that members of the ant species C. paria keep track of local time by socially interacting with workers (foragers) who shuttle in and out of the colony in search of food. (Author correspondence: vsharma@jncasr.ac.in)     

Difference in seasonal activity pattern between non-native and native ants in subtropical forest of Okinawa Island,Japan

Biological invasions by non-native tramp ants are reported throughout the world, particularly in island ecosystems. In Okinawa Island, a subtropical island in south-western Japan, many tramp ants including the invasive ant, Anoplolepis gracilipes, already occur. In this study activity patterns of the ground-foraging tramp and native ants were investigated for one year by using food bait traps at the forest edges where both categories of ants coexist. On a seasonal time scale, activity patterns were different between exotic and native ants. Native ants are active and seemingly more dominant from spring to summer whereas tramp ants become more dominant from autumn to winter. These results suggest there might be temporal niche partitioning between tramp and native ants, and native ants might be able to dominate tramp ants in their suitable seasons.     

Linking nutrition and behavioural dominance: carbohydrate scarcity limits aggression and activity in Argentine ants

Predicting the outcome of competitive interactions is a fundamental goal in ecology. Ecological stoichiometry, which relates nutrient balance to ecological processes, provides a framework for identifying mechanistic links among macronutrient availability, nutritional physiology and competitive performance. Because carbohydrates serve as a principal metabolic fuel, carbohydrate scarcity may impinge upon behaviours affecting competitive dominance (e.g. aggression, activity) to a greater extent than deficiencies of protein or other nutrients used preferentially for growth. Here, we tested this prediction with a diet manipulation study involving laboratory colonies of Argentine ants (Linepithema humile), a widespread and aggressive invasive species. The availability of both sucrose and insect prey influenced brood production and worker survival after three months. However, colonies became less aggressive and less active only when deprived of sucrose (but not prey). Scarcity of sucrose (but not prey) was also associated with reduced fat mass in individual workers. These data provide the first experimental support that carbohydrate scarcity compromises aggression and activity in ants, and illustrate, in principle, how access to carbohydrate-rich resources (e.g. plant exudates, hemipteran honeydew) might influence behavioural investments that contribute to competitive performance. Such investments might be especially important for invasive ants, given their aggressiveness and tendency to interact with honeydew-producing Hemiptera.     

Comparative studies of tidal rhythms. VI. Several clocks govern the activity of two species of fiddler crabs

The tide‐associated persistent activity rhythms of the ocypod crabs, Uca pugnax and U. pugilator were observed in constant conditions, some for as long as three months. In some animals the twice/day peaks of activity were found to scan the day at significantly different rates from one another. Individual peaks in some animals split into two fragments. And at times, one peak would fade and then return, or vanish completely. These examples of the independence between tide‐related activity peaks have become the basis for a hypothesis that each peak is controlled by its own clock(s). The basic period of these clocks is “circalunidian”, with periods varying up to about 4–8% on either side of 24.8 hours.     

Proximate Determinants of Reproductive Skew in Polygyne Colonies of the Ant Formica fusca

Understanding the determinants of reproductive skew (the partitioning of reproduction among co‐breeding individuals) is one of the major questions in social evolution. In ants, multiple‐queen nests are common and reproductive skew among queens has been shown to vary tremendously both within and between species. Proximate determinants of skew may be related to both queen and worker behaviour. Queens may attempt to change their reproductive share through dominance interactions, egg eating and by changing individual fecundity. Conversely, workers are in a position to regulate the reproductive output of queens when rearing the brood. This paper investigates queen behaviour at the onset of egg laying and the effect of queen fecundity and worker behaviour on brood development and reproductive shares of multiple queens in the ant Formica fusca. The study was conducted in two‐queen laboratory colonies where the queens produced only worker offspring. The results show that in this species reproductive apportionment among queens is not based on dominance behaviour and aggression, but rather on differences in queen fecundity. We also show that, although the queen fecundity at the onset of brood rearing is a good indicator of her final reproductive output, changes in brood composition occur during brood development. Our results highlight the importance of queen fecundity as a major determinant of her reproductive success. They furthermore suggest that in highly derived polygyne species, such as the Formica ants, direct interactions as a means for gaining reproductive dominance have lost their importance.     

Simple rules based on pile slope are used in the self organization of sand pile formation by <Emphasis Type="Italic">Pheidole oxyops</Emphasis> ants

We tested the hypothesis that slope influences where worker ants deposit excavated soil on piles near the nest entrance. We predicted that ants will deposit their load near the top of a pile where the slope changes from upward to downward, to prevent material rolling back towards the entrance. We tested this hypothesis by studying five natural colonies of Pheidole oxyops ants at a field site at S?o Sim?o, Brazil. At this site, each colony was dumping sandy soil excavated from its underground nest in a crescent-shaped pile c. 13 cm from and perpendicular to the nest entrance. Each nest was given an experimental sand pile of symmetrical curved cross section on a plywood platform that could be tilted 15 degrees up or down. From videos, the locations where individual ants dumped their soil loads were measured in relation to the inner (position = 0) and outer (position = 1) edges of the pile. When the platform was tilted down the ants deposited their loads significantly closer to the inner edge (0.458 ± 0.007) than when not tilted (0.530 ± 0.006). When the platform was tilted up the ants deposited their loads significantly further from the inner edge (0.626 ± 0.006) than when not tilted (0.522 ± 0.006). These results support the hypothesis that ants use pile slope in deciding where to dump their load. A similar rule is probably used in other ant species that place excavated soil in steep piles near the nest entrance. Received 5 February 2007; revised 10 June 2007; accepted 9 October 2007.     

广州地区红火蚁工蚁的巢外活动及有翅蚁的婚飞规律

为了阐明红火蚁Solenopsis invicta Buren的发生危害规律,为制定红火蚁的监测与防治措施提供科学依据,本研究调查了广州地区（113°45′E,22°43′N）红火蚁工蚁巢外活动日节律和季节性变化,以及有翅蚁的婚飞活动。结果表明,工蚁巢外活动日节律随季节或月份的不同而存在着明显差异：其中12-2月份的日活动节律为单峰型,即在中午温度较高时数量较大,而在5-10月份的日活动节律为双峰型,即在上午或下午工蚁数量较大,其余月份的活动为不明显的双峰型。工蚁巢外活动数量和时间随季节的不同而有显著差异。其中以6月和10月份的日活动数量最多,而在1月和2月份的活动数量最少,时间最短,其余月份活动数量居中。工蚁在阵雨前后的活动数量明显多于晴天,但处于降雨时刻的巢外工蚁数量极少。蚁巢受侵扰后出巢工蚁数量在30 s内最大,之后便随时间的延长而逐渐减少,该种现象可以用房室函数进行描述。另外,工蚁的巢外数量会随侵扰强度的加大而增加。在试验区内全年都可见到红火蚁婚飞,但婚飞活动主要集中在3-5月份,每日婚飞活动主要发生在下午,并主要发生在雨前或雨后。上述结果对于了解我国红火蚁的发生危害规律和提升其监控技术水平有较大参考价值。     

Corrigendum: Shear adhesive performance of leaf-cutting ant workers (Atta cephalotes)

In our previous article, Shear adhesive performance of leaf-cutting ant workers (Atta cephalotes), a technical error occurred when calculating shear adhesive performance divided by tarsal pad area of worker ants carrying leaves (Stark, A. Y., Davis, H. R., & Harrison, W. K. (2019). Biotropica, 51(4), 572–580). Shear adhesive performance of worker ants carrying leaves was mistakenly divided by the tarsal pad area of workers riding on leaves. This has slightly changed our results. After correcting for this error, there is now no statistically significant difference in shear adhesive performance divided by tarsal pad area of worker ants riding on leaves and those carrying leaves. Shear adhesive performance of these groups remains statistically different than soldiers, and our original results (shear adhesive performance of all three groups differ from one another) are consistent with the reported trend in the original article using true pad area for each individual (rather than estimated). We list below the pages and figures in the original article that this correction applies to. All authors agree with this change, apologize for their error, and appreciate the opportunity granted by the editors and publisher to make this correction. Our original conclusion, that there is variation in shear adhesive performance per unit area among worker roles of Atta cephalotes, is still supported by these corrected results.     

Dominance and species co-occurrence in highly diverse ant communities: a test of the interstitial hypothesis and discovery of a three-tiered competition cascade

The role of competitive exclusion is problematic in highly diverse ant communities where exceptional species richness occurs in the face of exceptionally high levels of behavioural dominance. A possible non-niche–based explanation is that the abundance of behaviourally dominant ants is highly patchy at fine spatial scales, and subordinate species act as insinuators by preferentially occupying these gaps—we refer to this as the interstitial hypothesis. To test this hypothesis, we examined fine-scale patterns of ant abundance and richness according to a three-tiered competition hierarchy (dominants, subdominants and subordinates) in an Australian tropical savanna using pitfall traps spaced at 2 m intervals. Despite the presence of gaps in the fine-scale abundance of individual species, the combined abundance of dominant ants (species of Iridomyrmex, Papyirus and Oecophylla) was relatively uniform. There was therefore little or no opportunity for subordinate species to preferentially occupy gaps in the foraging ranges of dominant species, and we found no relationship between the abundance of dominant ants and nondominant species richness at fine spatial scales. However, we found a negative relationship between subdominant and subordinate ants, a negative relationship between dominant and subdominant ants, and a positive relationship between dominant and subordinate ants. These results suggest that dominant species actually promote species richness by neutralizing the effects of subdominant species on subordinate species. Such indirect interactions have very close parallels with three-tiered trophic cascades in food webs, and we propose a “competition cascade” where the interactions are through a competition rather than trophic hierarchy.     

Synchronization of the behaviour within nests of the antLeptothorax acervorum (Fabricius)—II. Modelling the phenomenon and predictions from the model

Observations on activity of ants of the speciesLeptothorax acervorum show that ants within the nest are inactive for about 72% of their time (Frankset al., 1990.Bull. math. Biol.,52, 597–612). By examination of the activity of individual ants it is demonstrated that activity bouts of individuals are highly synchronized. The bursts of activity detected by Frankset al. occurred three to four times per hour. In this paper we develop a model to describe the phenomenon. As a result of the interdependence of the number of active ants within the nest and the high level of community activity some predictions are made, which are supported by experimental data in a quantitative way. In case of starvation the number of active ants will increase and no rhythms should occur. When proportionally more brood is present the rhythms should occur with a higher frequency. Eventually the rhythm breaks down and a stable equilibrium is reached.     

Changes in the speed of ants as a result of aggressive interactions

Subordinate ant species utilize different tactics to reduce competition with the stronger, larger and more aggressive individuals of a dominant species. In our experimental study, we assessed the behavioral response of individual workers of 4 subordinate ant species during their co‐occurrence with workers of a single dominant species. Contrary to most classical experiments focused on aggressive interactions, we assessed workers’ speed as a crucial factor in the outcome of co‐occurrence. Generally, there was a large intraspecific variation in the speed of the studied species—each had slow and fast individuals. Workers of all studied species moved faster just after interaction, suggesting that contact between 2 hostile workers is a stressful stimulus, generating a behavioral reaction of increasing speed. Also, the number of aggressive contacts experienced by a given individual positively affected its speed. Moreover, workers which were fast when exploring territory were also fast after interspecific interactions. The duration of aggression was significantly reduced by the speed and body size of a subordinate species worker—the more quickly a worker reacted and bigger it was, the shorter was the time of cumulative aggression. To our knowledge, this is the first study of this type to be conducted on ants and we conclude that speed is an overlooked and important characteristic of species and also individuals, therefore it should be considered as a driver of patterns of co‐occurrence in ant assemblages.     

Juvenile hormone and circadian locomotor activity in the honey bee Apis mellifera

Age-related division of labor in honeybees is associated with plasticity in circadian rhythms. Young nest bees care for brood around the clock with no circadian rhythms while older foragers have strong circadian rhythms that are used for sun compass navigation and for timing visits to flowers. Since juvenile hormone (JH) is involved in the coordination of physiological and behavioral processes underlying age-related division of labor in honey bees, we tested the hypothesis that JH influences the ontogeny of circadian rhythms and other clock parameters in young worker bees. Treatments with the JH analog methoprene or allatectomy did not influence the onset of rhythmicity, overall locomotor activity, or the free-running period of rhythmic locomotor behavior. There were, however, significant differences in the onset of rhythmicity, overall locomotor activity, and longevity between bees from different source colonies, suggesting that there is significant genetic variation for these traits. Our results suggest that JH does not coordinate all aspects of division of labor in bees and that coordination of task performance with circadian rhythms is probably mediated by other regulatory systems.     

Polyethism in social interactions in ants

Worker ants, which perform various tasks in their society (division of labour or polyethism), also demonstrate different types of antennal activity in trophallactic interactions. These differences concern the variability in successions of behavioural units performed by the worker in the course of substance transmission as well as in the sequential organization of these units. The antennal activity of an aged worker varies according to whether the ant gives food to another aged worker or to a callow worker. In the latter case, the donor's behaviour does not differ according to its social function (brood-tending worker or forager). A parallel can be established between these results and those obtained in a previous study of aggressive behaviour. This behaviour is liable to appear when a callow worker extracted from its society before hatching and therefore deprived of relations with aged workers of the society, is therefore placed with an aged worker. Polyethism is then manifested in social interactions, and not only in the division of labour between members of the society.     

Competition for food between the exotic wasp <Emphasis Type="Italic">Vespula germanica</Emphasis> and the native ant assemblage of NW Patagonia: evidence of biotic resistance?

The success of a biological invasion may depend on the interactions between the invader and the native biota. However, little experimental evidence demonstrates whether local species can successfully compete with exotics. We experimentally determined the existence of competition for food between the exotic wasp Vespula germanica, one of the most recent Patagonian invaders, and the native ant assemblage. Both wasps and ants are generalist predators and scavengers, sharing habitat and food resources. We selected 30 sites within scrubland habitats where both ants and wasps were present. At each site, we placed containers with protein baits under three treatments: wasp exclusion, ant exclusion, and control (i.e., free access for wasps and ants). Ant exclusion increased the number of wasps (with regard to a control), but wasp exclusion did not affect ant abundance. This result suggests that native ants affect the foraging activity of exotic wasps but not vice versa. Aggressive behaviors and worker aggregation may explain the competitive advantage of ants. Ants bite wasp legs and massively aggregate on food sources, physically limiting the landing of wasps on baits. If the outcome of interactions at baits reported here influence wasp population-level parameters, this competitive interaction could be one of the factors explaining the low abundance of this exotic wasp in NW Patagonia in comparison with other invaded regions.