Restricted antennal movement impacts the tandem running dynamics in a ponerine ant.

BACKGROUND: Tandem running is a recruitment method found in some species of ants where one ant follows another ant to reach a destination having maintained a physical contact with its antennae, throughout the journey. It is considered that the exchange of information regarding the destination among the nestmates happened during the process of tandem running. We examined the impact of restricting antennal movement on tandem running by using Diacamma indicum, a tandem-running ponerine ant by following 480 tandem runs across 9 treatment colonies and comparing it with 10 control relocating colonies. RESULT: Though all the 19 colonies relocated successfully, treatment colonies took significantly longer time to do so. Restricted antennal movement did not influence the ability to become tandem leaders, initiate tandem runs or the work organization significantly. However, antennae-restricted ants performed fewer tandem runs and took significantly longer time. Followers with single or both antennae-restriction performed significantly higher number of interruptions and the alignment between the leader and follower was impacted as antenna-restricted followers subtended a greater angle and walked more to the side of the leader as compared to the control followers. CONCLUSION: This study showed unhindered movement of the followers' antennae is important for tandem-running ants. In the next step, to gain a comprehensive understanding of this recruitment method, it is essential to individually delineate different sensory modalities.

Using pupae as appetitive reinforcement to study visual and tactile associative learning in the Ponerine ant Diacamma indicum.

Associative learning is of great importance to animals, as it enhances their ability to navigate, forage, evade predation and improve fitness. Even though associative learning abilities of Hymenopterans have been explored, many of these studies offered food as appetitive reinforcement. In the current study, we focus on tactile and visual cue learning in an ant Diacamma indicum using a Y-maze setup with pupa as a positive reinforcement. Using pupa as a reward resulted in a significantly higher proportion of ants completing the training in a shorter time as compared to using food as reinforcement. Ants spent significantly more time in the conditioned arm for both visual cues (white dots or black dots) and tactile cues (rough or smooth surfaces) presented on the floor when associated with pupa, thus showing that they were capable of associative learning. On encountering a conflict between visual and tactile cues during the test, ants chose to spend significantly more time on the arm with the tactile cues indicating that they had made a stronger association between pupa and the tactile cue as compared to the visual cue during training. Using pupa as an ecologically relevant reward, we show that these solitary foraging ants living in small colonies are capable of visual and tactile associative learning and are likely to learn tactile cues over visual cues in association with pupa.

Functional and mechanistic diversity in ant tandem communication.

Communication is fundamental to the organization of animal societies, often resulting in the convergent evolution of similar social behavior across lineages. However, this similarity may conceal underlying functional and mechanistic differences. Here we combined network and information-theoretic analysis to quantify how tandem recruitment is distinguishable between two ant genera, Temnothorax and Diacamma. We show that Temnothorax uses tandem running to recruit additional recruiters, while Diacamma uses it principally to move the passive majority of their colony, a task that Temnothorax accomplishes with a different behavior, social carrying. Accordingly, the network structure of Diacamma tandems was dissimilar to that of Temnothorax, instead resembling the social-carrying networks in Temnothorax. Furthermore, our information-theoretical analysis on movement trajectories revealed that Diacamma tandem runs lack bidirectional information transfer, the signature of route learning in Temnothorax. By quantifying the diversity of similar communication systems, this study increases the resolution of our understanding of animal societies.

Characterization of Nest Architecture of an Indian Ant Diacamma indicum (Hymenoptera: Formicidae).

Nests are physical entities that give shelter to the inhabitants from natural adversities, predators and act as a platform for organization of tasks particularly in social insects. Social insect nests can range from simple structures consisting of a single entrance leading to a chamber to complex nests containing hundreds of connected shafts and chambers. This study characterizes nest architecture of a tropical ponerine ant Diacamma indicum (Hymenoptera: Formicidae), which has small colony sizes and is known to be a scavenger. We also examined if these nests vary seasonally. By examining the microhabitat in the vicinity of the nest, the nest entrance characteristics and casting 77 natural nests of D. indicum across a year, we found that this species occupies relatively simple nests consisting of a single entrance that leads to a single chamber. This chamber progressed to a secondary tunnel that terminated at a greater depth than the chamber. The nest volume was not correlated to the number of adult members in the colony. Even though the microhabitat around the nest and the entrance itself change across seasons, principle component analysis showed that the nest architecture remained similar. Only one parameter, the entrance tunnel showed significant difference and was longer during postmonsoon. Nests of colonies living in the immediate vicinity of human habitation were comparable to other nests. We conclude that D. indicum found in the Gangetic plains live in relatively simple nests that do not vary across seasons.

Path minimization in a tandem running Indian ant in the context of colony relocation.

The phenomenon of minimizing the path length to a target site in order to increase transport efficiency is described as path optimization, and it has been observed in many mammals, birds and some invertebrates such as honeybees and ants. It has been demonstrated that ants can optimize their foraging path through an emergent process, involving the trail pheromone concentration, without individual ants having to measure and compare distances. In the current study, we investigated whether ants that use only tandem running to recruit their nestmates can minimize their path while relocating their entire colony into a new nest. As colony relocation directly impacts the survival of the whole colony, it would be particularly important to optimize their path to the new nest. Using the ponerine ant Diacamma indicum, we conducted relocation experiments, in which ants had to choose between different defined paths, and contrasted our findings with open arena experiments, as they navigated to their new nest. After following 4100 unique transports by 450 different transporters, we found that these ants do minimize their path. Individual leaders, as well as colonies, chose the shorter path significantly more than the longer path, and they showed a significant preference for the shorter arm at multiple decision points on encountering a combination of paths. Thus, we concluded that tandem leaders are capable of path minimization based on the information they themselves collect. Further investigation into the proximate mechanisms by which they achieve this is required.

Caught red-handed: behaviour of brood thieves in an Indian ant.

Theft of resources is ubiquitous in the animal kingdom. An evolutionary arms race between thieves and their victims is expected. Although several studies have documented inter- and intraspecific theft of resources in different taxa, studies that delve into the behaviour of thieves and the factors that influence their behaviour have not been undertaken. In the current study on the primitively eusocial ant Diacamma indicum, we caught brood thieves red-handed: we observed them in the act of stealing brood and examined their behaviour. Thieves were persistent in their attempts despite facing aggression in the victim colony. Experiencing aggression or failure to steal in the previous attempt negatively impacted a thief's drive to reattempt. To avoid the risks associated with theft, successful thieves exited from victim nests about three times faster than others who were procuring brood from unguarded nests. In a series of experiments examining factors that caused thieves to increase their exit speed, we found that indirect cues of a foreign colony's presence, such as odour or the presence of foreign ants, did not induce these changes in thieves. Thus, we conclude that these ant thieves only respond to the direct threat posed by aggressive foreign ants. In this comprehensive study using behavioural experiments, we reveal the simple rules of engagement between victims and brood thieves.

To reunite or not: A study of artificially fragmented Diacamma indicum ant colonies.

Social insects live together in groups and maintain cohesion to enhance their chances of survival and productivity. Colony cohesion is severely challenged during relocation. We examined the dynamics of colony reunification and the factors affecting nest choice of artificially fragmented colonies of the queenless ant Diacamma indicum. None of the twelve undisturbed colonies fragmented or relocated when a good nest was available in their neighbourhood. When colonies were artificially fragmented, they mostly (25/30) reunified into a single nest unlike in randomized time-ordered network models, indicating that reunification is not the result of random recruitment acts. When the reproductive individual was present in a good nest, the colonies reunified at this address. However, when she was present in a suboptimal nest, colonies relocated her to a better quality nest and reunified there, illustrating that quality of the new nest is more important. The work distribution and relocation dynamics of reunification were comparable to intact colonies relocating to a single new nest. This is made possible by enhanced exchange of information among tandem leaders in the form of increased number of tandem runs among them. We conclude that colony cohesion is very important and is maintained after incorporating the risks of relocation and preference for nest quality during decision making.

Tricks of the trade: Mechanism of brood theft in an ant.

Thievery is ubiquitous in the animal kingdom, social insects not being an exception. Brood is invaluable for the survival of social insect colonies and brood theft is well documented in ants. In many species the stolen brood act as slaves in the thief colony as they take up tasks related to foraging, defence and colony maintenance. Slave-making (dulotic) ants are at an advantage as they gain workforce without investing in rearing immature young, and several slave-making species have been recorded in temperate regions. In the current study we investigate brood theft in a primitively eusocial ponerine ant Diacamma indicum that inhabits the tropics. In the context of colony relocation we asked how thieves steal brood and what victim colonies do to prevent theft. While exposed nests increased colonies' vulnerability, the relocation process itself did not enhance the chances of theft. Various aggressive interactions, in particular immobilization of intruders helped in preventing theft. Thieves that acted quickly, stayed furtive and stole unguarded brood were found to be successful. This comprehensive study of behavioural mechanism of theft reveals that these are the 'tricks' adopted by thieves.

Network approach to understanding the organization of and the consequence of targeted leader removal on an end-oriented task.

Relocation is an important event in the lives of several social insects whereby all colony members have to be transferred to a new nest when conditions in the old nest become unfavorable. In the current study, network tools were used to examine the organization of this goal-oriented task in the Indian queenless ant Diacamma indicum which relocate their colonies by means of tandem running. Individual ants were used as nodes and tandem runs as directed edges to construct unweighted networks. Network parameters were characterized in control relocations (CRs) and in relocations where the node with the highest outdegree, that is, the Maximum tandem leader (Max TL) was experimentally removed. These were then compared to 1) randomized networks, 2) simulated networks in which Max TL was removed, and 3) simulated networks with removal of a random leader. Not only was there complete recovery of the task, but the manner in which it was organized when Max TL was removed was comparable to CRs. The results obtained from our empirical study were significantly different from the results predicted by simulations of leader removal. At an individual level, the Max TL had a significantly higher outdegree than expected by chance alone and in her absence the substitute Max TL did comparable work. In addition, the position of the Max TL in the pathway of information flow was conserved in control and experimentally manipulated conditions. Understanding the organization of this critical event as more than the sum of individual interactions using network parameters allows us to appreciate the dynamic response of groups to perturbations.

Deciphering a survival strategy during the interspecific competition between Bacillus cereus MSM-S1 and Pseudomonas sp. MSM-M1.

Interspecific competition in bacteria governs colony growth dynamics and pattern formation. Here, we demonstrate an interesting phenomenon of interspecific competition between Bacillus cereus MSM-S1 and Pseudomonas sp. MSM-M1, where secretion of an inhibitor by Pseudomonas sp. is used as a strategy for survival. Although B. cereus grows faster than Pseudomonas sp., in the presence of Pseudomonas sp. the population of B. cereus reduces significantly, whereas Pseudomonas sp. do not show any marked alteration in their population growth. Appearance of a zone of inhibition between growing colonies of two species on nutrient agar prevents the expanding front of the MSM-S1 colony from accessing and depleting nutrients in the region occupied by MSM-M1, thereby aiding the survival of the slower growing MSM-M1 colonies. To support our experimental results, we present simulations, based on a chemotactic model of colony growth dynamics. We demonstrate that the chemical(s) secreted by Pseudomonas sp. is responsible for the observed inhibition of growth and spatial pattern of the B. cereus MSM-S1 colony. Our experimental results are in excellent agreement with the numerical results and confirm that secreted inhibitors enable Pseudomonas sp. to survive and coexist in the presence of faster growing B. cereus, in a common niche.

Opportunistic brood theft in the context of colony relocation in an Indian queenless ant.

Brood is a very valuable part of an ant colony and behaviours increasing its number with minimum investment is expected to be favoured by natural selection. Brood theft has been well documented in ants belonging to the subfamilies Myrmicinae and Formicinae. In this study we report opportunistic brood theft in the context of nest relocation in Diacamma indicum, belonging to the primitively eusocial subfamily Ponerinae. Pupae was the preferred stolen item both in laboratory conditions and in natural habitat and a small percentage of the members of a colony acting as thieves stole about 12% of the brood of the victim colony. Stolen brood were not consumed but became slaves. We propose a new dimension to the risks of relocation in the form of brood theft by conspecific neighbours and speculate that examination of this phenomenon in other primitively eusocial species will help understand the origin of brood theft in ants.

Tight knit under stress: colony resilience to the loss of tandem leaders during relocation in an Indian ant.

The movement of colonies from one nest to another is a frequent event in the lives of many social insects and is important for their survival and propagation. This goal-oriented task is accomplished by means of tandem running in some ant species, such as Diacamma indicum. Tandem leaders are central to this process as they know the location of the new nest and lead colony members to it. Relocations involving targeted removal of leaders were compared with unmanipulated and random member removal relocations. Behavioural observations were integrated with network analysis to examine the differences in the pattern of task organization at the level of individuals and that of the colony. All colonies completed relocation successfully and leaders who substituted the removed tandem leaders conducted the task at a similar rate having redistributed the task in a less skewed manner. In terms of network structure, this resilience was due to significantly higher density and outcloseness indicating increased interaction between substitute leaders. By contrast, leader-follower interactions and random removal networks showed no discernible changes. Similar explorations of other goal-oriented tasks in other societies will possibly unveil new facets in the interplay between individuals that enable the group to respond effectively to stress.

Dual response to nest flooding during monsoon in an Indian ant.

Flooding causes destruction of shelter and disruption of activity in animals occupying subterranean nests. To ensure their survival organisms have evolved various responses to combat this problem. In this study we examine the response of an Indian ant, Diacamma indicum, to nest flooding during the monsoon season. Based on characterization of nest location, architecture and the response of these ants to different levels of flooding in their natural habitat as well as in the laboratory, we infer that they exhibit a dual response. On the one hand, the challenges presented by monsoon are dealt with by occupying shallow nests and modifying the entrance with decorations and soil mounds. On the other hand, inundated nests are evacuated and the ants occupy shelters at higher elevations. We conclude that focused studies of the monsoon biology of species that dwell in such climatic conditions may help us appreciate how organisms deal with, and adapt to, extreme seasonal changes.