Social Insect Colonies Research Articles

Autonomous tracking of honey bee behaviors over long-term periods with cooperating robots.

Digital and mechatronic methods, paired with artificial intelligence and machine learning, are transformative technologies in behavioral science and biology. The central element of the most important pollinator species-honey bees-is the colony's queen. Because honey bee self-regulation is complex and studying queens in their natural colony context is difficult, the behavioral strategies of these organisms have not been widely studied. We created an autonomous robotic observation and behavioral analysis system aimed at continuous observation of the queen and her interactions with worker bees and comb cells, generating behavioral datasets of exceptional length and quality. Key behavioral metrics of the queen and her social embedding within the colony were gathered using our robotic system. Data were collected continuously for 24 hours a day over a period of 30 days, demonstrating our system's capability to extract key behavioral metrics at microscopic, mesoscopic, and macroscopic system levels. Additionally, interactions among the queen, worker bees, and brood were observed and quantified. Long-term continuous observations performed by the robot yielded large amounts of high-definition video data that are beyond the observation capabilities of humans or stationary cameras. Our robotic system can enable a deeper understanding of the innermost mechanisms of honey bees' swarm-intelligent self-regulation. Moreover, it offers the possibility to study other social insect colonies, biocoenoses, and ecosystems in an automated manner. Social insects are keystone species in all terrestrial ecosystems; thus, developing a better understanding of their behaviors will be invaluable for the protection and even the restoration of our fragile ecosystems globally.

Ecological principles for the evolution of communication in collective systems.

Communication allows members of a collective to share information about their environment. Advanced collective systems, such as multicellular organisms and social insect colonies, vary in whether they use communication at all and, if they do, in what types of signals they use, but the origins of these differences are poorly understood. Here, we develop a theoretical framework to investigate the evolution and diversity of communication strategies under collective-level selection. We find that whether communication can evolve depends on a collective's external environment: communication only evolves in sufficiently stable environments, where the costs of sensing are high enough to disfavour independent sensing but not so high that the optimal strategy is to ignore the environment altogether. Moreover, we find that the evolution of diverse signalling strategies-including those relying on prolonged signalling (e.g. honeybee waggle dance), persistence of signals in the environment (e.g. ant trail pheromones) and brief but frequent communicative interactions (e.g. ant antennal contacts)-can be explained theoretically in terms of the interplay between the demands of the environment and internal constraints on the signal. Altogether, we provide a general framework for comparing communication strategies found in nature and uncover simple ecological principles that may contribute to their diversity.

The Evolution of Queen Pheromone Production and Detection in the Reproductive Division of Labor in Social Insect Colonies.

Structurally diverse queen pheromones and fertility signals regulate the reproductive division of labor of social insects, such as ants, termites, some bees, and some wasps. The independent evolution of sociality in these taxa allows for the exploration of how natural history differences in sender and receiver properties led to the evolution of these complex communication systems. While describing the different effects and the structural diversity of queen pheromones, we identify two major syndromes that mostly separate ants and wasps from bees and termites in their use of different pheromone classes. We compare olfactory receptor evolution among these groups and review physiological and hormonal links to fecundity and pheromone production. We explore the cases in which queen pheromone evolution is conserved, convergent, or parallel and those in which queen pheromone responses are more likely to be learned or innate. More mechanistic information about the pathways linking fecundity to queen pheromone production and perception could help close major knowledge gaps.

The refuse dump provides information that influences the foraging preferences of leaf‐cutting ants

Abstract The ability to respond to local stimuli individually and propagate information among colony members has allowed social organisms to generate collective responses. Leaf‐cutting ants are an excellent model to study this because, apart from exchanging information among groups assigned to different tasks (foragers, gardeners, and midden workers—ants working in refuse dumps), they also do so with their symbiotic fungus. Here, we tested the role of refuse dumps as an information centre influencing the foraging decisions of Acromyrmex lobicornis under field conditions. To simulate the presence of discarded resources, oat flakes were added to the refuse dumps of treated colonies, while the refuse dumps of controls remained untreated. After 24 h and for 6 weeks, we offered new clean oat flakes on the foraging trails simultaneously with cornflakes, another palatable resource that was never experienced by the ants on the dump. Although the oat flakes were suitable for the ants and harmless to the fungus, the acceptance index—a ratio of oat flakes foraged to total oat and cornflakes offered—decreased by 55% 24 h after the ants interacted with oat flakes in the refuse dump. After 6 weeks, the acceptance index recovered to levels that did not differ from initial values. Therefore, ants might use waste to obtain information about the suitability of resources found in foraging trails. We explore potential mechanisms underlying the transmission of information from refuse dumps to foraging trails, shedding light on the intricate dynamics of collective decision‐making in social insect colonies.

Reverse social contagion as a mechanism for regulating mass behaviors in highly integrated social systems.

Mass behavior is the rapid adoption of similar conduct by all group members, with potentially catastrophic outcomes such as mass panic. Yet, these negative consequences are rare in integrated social systems such as social insect colonies, thanks to mechanisms of social regulation. Here, we test the hypothesis that behavioral deactivation between active individuals is a powerful social regulator that reduces energetic spending in groups. Borrowing from scaling theories for human settlements and using behavioral data on harvester ants, we derive ties between the hypermetric scaling of the interaction network and the hypometric scaling of activity levels, both relative to the colony size. We use elements of economics theory and metabolic measurements collected with the behavioral data to link activity and metabolic scalings with group size. Our results support the idea that metabolic scaling across social systems is the product of different balances between their social regulation mechanisms.

The genomic and cellular basis of biosynthetic innovation in rove beetles

How evolution at the cellular level potentiates macroevolutionary change is central to understanding biological diversification. The >66,000 rove beetle species (Staphylinidae) form the largest metazoan family. Combining genomic and cell type transcriptomic insights spanning the largest clade, Aleocharinae, we retrace evolution of two cell types comprising a defensive gland-a putative catalyst behind staphylinid megadiversity. We identify molecular evolutionary steps leading to benzoquinone production by one cell type via a mechanism convergent with plant toxin release systems, and synthesis by the second cell type of a solvent that weaponizes the total secretion. This cooperative system has been conserved since the Early Cretaceous as Aleocharinae radiated into tens of thousands of lineages. Reprogramming each cell type yielded biochemical novelties enabling ecological specialization-most dramatically in symbionts that infiltrate social insect colonies via host-manipulating secretions. Our findings uncover cell type evolutionary processes underlying the origin and evolvability of a beetle chemical innovation.

Inter-clonal competition over queen succession imposes a cost of parthenogenesis on termite colonies.

In social insect colonies, selfish behaviour due to intracolonial conflict among members can result in colony-level costs despite close relatedness. In certain termite species, queens use asexual reproduction for within-colony queen succession but rely on sexual reproduction for worker and alate production, resulting in multiple half-clones of a single primary queen competing for personal reproduction. Our study demonstrates that competition over asexual queen succession among different clone types leads to the overproduction of parthenogenetic offspring, resulting in the production of dysfunctional parthenogenetic alates. By genotyping the queens of 23 field colonies of Reticulitermes speratus, we found that clone variation in the queen population reduces as colonies develop. Field sampling of alates and primary reproductives of incipient colonies showed that overproduced parthenogenetic offspring develop into alates that have significantly smaller body sizes and much lower survivorship than sexually produced alates. Our results indicate that while the production of earlier and more parthenogenetic eggs is advantageous for winning the competition for personal reproduction, it comes at a great cost to the colony. Thus, this study highlights the evolutionary interplay between individual-level and colony-level selection on parthenogenesis by queens.

Dynamics of Information Flow and Task Allocation of Social Insect Colonies: Impacts of Spatial Interactions and Task Switching.

Models of social interaction dynamics have been powerful tools for understanding the efficiency of information spread and the robustness of task allocation in social insect colonies. How workers spatially distribute within the colony, or spatial heterogeneity degree (SHD), plays a vital role in contact dynamics, influencing information spread and task allocation. We used agent-based models to explore factors affecting spatial heterogeneity and information flow, including the number of task groups, variation in spatial arrangements, and levels of task switching, to study: (1) the impact of multiple task groups on SHD, contact dynamics, and information spread, and (2) the impact of task switching on SHD and contact dynamics. Both models show a strong linear relationship between the dynamics of SHD and contact dynamics, which exists for different initial conditions. The multiple-task-group model without task switching reveals the impacts of the number and spatial arrangements of task locations on information transmission. The task-switching model allows task-switching with a probability through contact between individuals. The model indicates that the task-switching mechanism enables a dynamical state of task-related spatial fidelity at the individual level. This spatial fidelity can assist the colony in redistributing their workforce, with consequent effects on the dynamics of spatial heterogeneity degree. The spatial fidelity of a task group is the proportion of workers who perform that task and have preferential walking styles toward their task location. Our analysis shows that the task switching rate between two tasks is an exponentially decreasing function of the spatial fidelity and contact rate. Higher spatial fidelity leads to more agents aggregating to task location, reducing contact between groups, thus making task switching more difficult. Our results provide important insights into the mechanisms that generate spatial heterogeneity and deepen our understanding of how spatial heterogeneity impacts task allocation, social interaction, and information spread.

The rediscovery of the putative ant social parasite Manica parasitica syn. nov. (Hymenoptera: Formicidae) reveals an unexpected endoparasite syndrome.

Parasitism is ubiquitous across the tree of life, and parasites comprise approximately half of all animal species. Social insect colonies attract many pathogens, endo- and ectoparasites, and are exploited by social parasites, which usurp the social environment of their hosts for survival and reproduction. Exploitation by parasites and pathogens versus social parasites may cause similar behavioural and morphological modifications of the host. Ants possess two overlapping syndromes: the endo- and social parasite syndromes. We rediscovered two populations of the putative social parasite Manica parasitica in the Sierra Nevada, and tested the hypothesis that M. parasitica is an independently evolving social parasite. We evaluated traits used to discriminate M. parasitica from its host Manica bradleyi, and examined the morphology of M. parasitica in the context of ant parasitic syndromes. We find that M. parasitica is not a social parasite. Instead, M. parasitica represents cestode-infected M. bradleyi. We propose that M. parasitica should be regarded as a junior synonym of M. bradleyi. Our results emphasize that an integrative approach is essential for unravelling the complex life histories of social insects and their symbionts.

Precise tactile stimulation of worker ants by a robotic manipulator reveals that individual responses are density‐ and context‐dependent

Abstract Ant workers are often specialized in specific tasks, and it is well‐established that the main task an ant performs in the colony can be used to predict its sensitivity and responses to task‐associated stimuli. An often‐overlooked aspect of ants' task specialization is that individuals often switch tasks throughout the day and are not always engaged in functional tasks. Furthermore, the tasks individuals engage in are often correlated with other context‐specific factors, such as worker density, which can independently influence individuals' behaviour. Given this intra‐individual variation in task engagement and its correlation with density, it is currently unknown how these two factors interact to modulate ants' sensitivity and responses to stimuli. To address this question, we built a robotic manipulation system that allowed us to teleoperate a dummy inside ant colonies and to provide simulated antennations to ants when performing different tasks in areas with different worker densities. We coupled this manipulation system with a custom‐built automated tracking system (FORT) that allowed us to track individual identities and locations as well as to record the ants' responses to the dummy stimulation. We found independent effects of task and worker density on ants' responsiveness and alarm towards the dummy. Ants were less likely to respond and be alarmed by the dummy when stimulated in areas with high worker density. Responsiveness but not alarm was further influenced by the task being performed, with ants doing broodcare being the least responsive. Our results suggest that ants' behaviour is density‐dependent and that ants experience a process of habituation to tactile stimulation. Additionally, ants' responsiveness is modulated by the task they are performing at a given time, showing that sensitivity to stimuli is context‐dependent. Our robotic set‐up constitutes a valuable tool to systematically investigate social insect behaviour under unprecedented experimental control to unravel the individual‐level behavioural rules that underpin the organization of social insect colonies. The integrated system presented here opens new research avenues to empirically investigate the effects of more complex stimuli on social insect behaviour and has the potential to significantly further our understanding of decentralized collective systems.

A modeling framework for adaptive collective defense: crisis response in social-insect colonies.

Living systems, from cells to superorganismic insect colonies, have an organizational boundary between inside and outside and allocate resources to defend it. Whereas the micro-scale dynamics of cell walls can be difficult to study, the adaptive allocation of workers to defense in social-insect colonies is more conspicuous. This is particularly the case for Tetragonisca angustula stingless bees, which combine different defensive mechanisms found across other colonial animals: (1) morphological specialization (distinct soldiers(majors) are produced over weeks); (2) age-based polyethism (young majors transition to guarding tasks over days); and (3) task switching (small workers(minors) replace soldiers within minutes under crisis). To better understand how these timescales of reproduction, development, and behavior integrate to balance defensive demands with other colony needs, we developed a demographic Filippov ODE system to study the effect of these processes on task allocation and colony size. Our results show that colony size peaks at low proportions of majors, but colonies die if minors are too plastic or defensive demands are too high or if there is a high proportion of quickly developing majors. For fast maturation, increasing major production may decrease defenses. This model elucidates the demographic factors constraining collective defense regulation in social insects while also suggesting new explanations for variation in defensive allocation at smaller scales where the mechanisms underlying defensive processes are not easily observable. Moreover, our work helps to establish social insects as model organisms for understanding other systems where the transaction costs for component turnover are nontrivial, as in manufacturing systems and just-in-time supply chains.

Chemical Resemblance of Egg Surface Compounds and Dufour's Gland in Two Neotropical Polistinae Wasps Polistes versicolor (Olivier) and Mischocyttarus metathoracicus (de Saussure, 1854).

Chemical communication plays a major role in regulating social dynamics in social insect colonies. The most studied class of chemical compounds are the cuticular hydrocarbons (CHCs), compounds with high molecular weight that cover the insect body. CHCs are used in nestmate recognition and to signal reproductive status. Brood, in the form of larvae and eggs, is known to participate in chemical communication and social dynamics by performing hunger behaviour and inducing interaction with adults and conferring nest and maternity identity. CHCs of adults and egg surface compounds are similar in composition in social insect species. The main source of egg compounds is proposed to be Dufour's gland, an accessory reproductive gland found in several Hymenoptera females. There is still a lack of information about the level of similarity among CHCs, compounds of egg surface and Dufour's gland for several wasp species, which could provide correlational evidence about the origins of egg-marking compounds. Thus, we investigated whether egg surface compounds were more similar to CHCs or Dufour's gland secretions in two Neotropical primitively eusocial wasp species, Polistes versicolor (Olivier) and Mischocyttarus metathoracicus (de Saussure, 1854). As expected, there was a higher chemical similarity between eggs and Dufour's gland secretions in both studied species, supporting the hypothesis that this gland is the source of chemical compounds found over the eggs in these two primitively eusocial species.

Indirect exposure to insect growth disruptors affects honey bee (Apis mellifera) reproductive behaviors and ovarian protein expression.

Pesticide exposure and queen loss are considered to be major causes of honey bee colony mortality, yet little is known regarding the effects of regularly encountered agrochemicals on honey bee reproduction. Here, we present the results of a two-generational study using specialized cages to expose queens to commonly used insect growth disrupting pesticides (IGDs) via their retinue of worker bees. Under IGD exposure, we tracked queen performance and worker responses to queens, then the performance of the exposed queens' offspring was assessed to identify patterns that may contribute to the long-term health and stability of a social insect colony. The positive control, novaluron, resulted in deformed larvae hatching from eggs laid by exposed queens, and methoxyfenozide, diflubenzuron, and novaluron caused a slight decrease in daily egg laying rates, but this was not reflected in the total egg production over the course of the experiment. Curiously, eggs laid by queens exposed to pyriproxyfen exhibited increased hatching rates, and those larvae developed into worker progeny with increased responsiveness to their queens. Additionally, pyriproxyfen and novaluron exposure affected the queen ovarian protein expression, with the overwhelming majority of differentially expressed proteins coming from the pyriproxyfen exposure. We discuss these results and the potential implications for honey bee reproduction and colony health.

Influence of juvenile hormone analog on behavior in the red imported fire ant, Solenopsis invicta

Division of labor is a hallmark characteristic of social insect colonies. While it is understood that worker differentiation is regulated through either the queen or her brood, the understanding of the physiology behind task regulation varies within social species. Studies in eusocial insects have shown that juvenile hormone (JH) is associated with division of labor and the onset of foraging tasks. Although, outside of a few key species, this interaction has yet to be elucidated in the red imported fire ant, Solenopsis invicta. In this study, we evaluated the role of a JH analog, S-hydroprene in worker task transition in Solenopsis invicta. S-hydroprene was applied to nurses to observe behavioral changes. S-hyroprene application to nurses did not affect phototaxis, but there was a shift in behavior from internal, nest-based behaviors to external, foraging-based behaviors. These results show that JH may be implicated in worker task transition in S. invicta and may function similarly as it does in other eusocial insects.

Evolution of the neuronal substrate for kin recognition in social Hymenoptera.

In evolutionary terms, life is about reproduction. Yet, in some species, individuals forgo their own reproduction to support the reproductive efforts of others. Social insect colonies for example, can contain up to a million workers that actively cooperate in tasks such as foraging, brood care and nest defence, but do not produce offspring. In such societies the division of labour is pronounced, and reproduction is restricted to just one or a few individuals, most notably the queen(s). This extreme eusocial organisation exists in only a few mammals, crustaceans and insects, but strikingly, it evolved independently up to nine times in the order Hymenoptera (including ants, bees and wasps). Transitions from a solitary lifestyle to an organised society can occur through natural selection when helpers obtain a fitness benefit from cooperating with kin, owing to the indirect transmission of genes through siblings. However, this process, called kin selection, is vulnerable to parasitism and opportunistic behaviours from unrelated individuals. An ability to distinguish kin from non-kin, and to respond accordingly, could therefore critically facilitate the evolution of eusociality and the maintenance of non-reproductive workers. The question of how the hymenopteran brain has adapted to support this function is therefore a fundamental issue in evolutionary neuroethology. Early neuroanatomical investigations proposed that social Hymenoptera have expanded integrative brain areas due to selection for increased cognitive capabilities in the context of processing social information. Later studies challenged this assumption and instead pointed to an intimate link between higher social organisation and the existence of developed sensory structures involved in recognition and communication. In particular, chemical signalling of social identity, known to be mediated through cuticular hydrocarbons (CHCs), may have evolved hand in hand with a specialised chemosensory system in Hymenoptera. Here, we compile the current knowledge on this recognition system, from emitted identity signals, to the molecular and neuronal basis of chemical detection, with particular emphasis on its evolutionary history. Finally, we ask whether the evolution of social behaviour in Hymenoptera could have driven the expansion of their complex olfactory system, or whether the early origin and conservation of an olfactory subsystem dedicated to social recognition could explain the abundance of eusocial species in this insect order. Answering this question will require further comparative studies to provide a comprehensive view on lineage-specific adaptations in the olfactory pathway of Hymenoptera.

Biology of the southern giant hornet, Vespa soror: nest architecture, morphological differences among castes, and the genetic structure of colonies.

Giant hornets in the genus Vespa are apex predators that are known throughout Asia for their exceptional size and devastating group attacks on social insect colonies. The giant hornets include Vespa mandarinia, a well-studied and widespread temperate species, and Vespa soror, a poorly known sister species that is limited to subtropical and tropical regions of Southeast Asia. Both species have been recently documented on the west coast of North America, raising urgent questions about their potential impact in novel ecosystems. To better understand the biology of V. soror, we describe the nest architecture, caste morphology, and genetic structure of colonies collected in Vietnam. Comparisons of colony metrics between the two giant hornet species suggest important differences that are likely a consequence of the relatively warmer climate in which V. soror occurs. Like V. mandarinia, V. soror constructs large, underground nests of partially enveloped horizontal combs. However, compared to temperate V. mandarinia colonies, the longer nesting period of subtropical V. soror colonies likely resulted in relatively larger colony sizes and nests by the end of their annual cycle. Vespa soror workers and gynes were larger than males, distinguishable based on wing shape and body size (total length and measures of six body parts), and equivalent in size to female castes of V. mandarinia. We genotyped colony members from three mature nests, which revealed that males and females were offspring of singly mated queens. Two colonies were monogynous, but one colony was comprised of two unrelated matrilines. Polygyny has not been observed for V. mandarinia, but is more common in tropical hornet species. Our study sheds light on essential details about the biology of an understudied species of giant hornet, whose populous colonies and long nesting period suggest the potential for substantial ecological impact wherever they occur.

Aethina tumida

Social insect colonies live in nests and are superorganisms consisting of individuals analogous to cells in metazoans. Small hive beetles, Aethina tumida, are free-flying parasites of social bee colonies, where they feed and reproduce. The kleptoparasitic adults exploit social feeding between hosts. Adults and larvae feed on honey, pollen, and host brood, which can damage colonies. Damage rarely occurs in the endemic range of A. tumida in sub-Saharan Africa. They have become a widespread invasive species and can cause severe damage to managed colonies in the new ranges, raising concern about the possible impact on wild bees. However, this complex host–parasite system, with multiple host species and high parasite opportunism, imposes challenges for our understanding, and respective mitigation, for example, via traps, RNAi, and adjusted beekeeping management. Accordingly, there are considerable gaps in our knowledge of this parasite, calling for more research in all areas of its biology.

Significant colony differences in thermal tolerances and decoupling of high and low critical temperatures in the army ant Eciton burchellii parvispinum

Abstract Differences in behaviour and physiology among social insect colonies provide opportunities for evolutionary responses to environmental selection. Colony differences in thermal physiology can indicate potential for social insect populations to accommodate temperature extremes and determine how populations will respond to changing climates. We tested whether army ant (Eciton burchellii parvispinum) colonies differed in their distributions of worker tolerances of extreme low temperatures (CTmin) and extreme high temperatures (CTmax). We sampled workers across a range of body sizes in each colony, and we collected data from colonies at both warmer low elevation sites and cooler high elevation sites. Colonies differed significantly in the distributions of both CTmin and CTmax; colony mean CTmin and CTmax were not correlated. The colony differences were not associated with worker body size variation. Some thermal physiology differences were associated with local climates: colonies from warmer lower elevation sites had significantly higher CTmin and higher CTmax, but much of the inter‐colony variation was unexplained. If significant colony variation in thermal physiology is associated with genetic differences, thermal environments could drive evolutionary changes in thermal physiology.

Social Learning versus Individual Learning in the Division of Labour.

Division of labour, or the differentiation of the individuals in a collective across tasks, is a fundamental aspect of social organisations, such as social insect colonies. It allows for efficient resource use and improves the chances of survival for the entire collective. The emergence of large inactive groups of individuals in insect colonies sometimes referred to as laziness, has been a puzzling and hotly debated division-of-labour phenomenon in recent years that is counter to the intuitive notion of effectiveness. It has previously been shown that inactivity can be explained as a by-product of social learning without the need to invoke an adaptive function. While highlighting an interesting and important possibility, this explanation is limited because it is not yet clear whether the relevant aspects of colony life are governed by social learning. In this paper, we explore the two fundamental types of behavioural adaptation that can lead to a division of labour, individual learning and social learning. We find that inactivity can just as well emerge from individual learning alone. We compare the behavioural dynamics in various environmental settings under the social and individual learning assumptions, respectively. We present individual-based simulations backed up by analytic theory, focusing on adaptive dynamics for the social paradigm and cross-learning for the individual paradigm. We find that individual learning can induce the same behavioural patterns previously observed for social learning. This is important for the study of the collective behaviour of social insects because individual learning is a firmly established paradigm of behaviour learning in their colonies. Beyond the study of inactivity, in particular, the insight that both modes of learning can lead to the same patterns of behaviour opens new pathways to approach the study of emergent patterns of collective behaviour from a more generalised perspective.

Aliens in the society: foreign arthropods and small vertebrates associated with the social spider Stegodyphus sarasinorum Karsch, 1892 (Araneae: Eresidae)

Animals use nests for various functions including laying eggs, raising young and gaining protection from predators. Social insect colonies provide rich microhabitats for various foreign inhabitants. Social spiders in the family Eresidae live in large, long-lasting colonies with a dense silken nest where spiders reside in complex capture webs. Social spider colonies contain several organisms apart from the host spiders, yet little is known about the foreign species and their roles. In this study, we cataloged foreign animals found in the nests and webs of the Indian social spider Stegodyphus sarasinorum Karsch, 1892 from different sites in India. We then examined the abundance and diversity of foreign spiders in adult and juvenile colonies, and the nature of interaction of foreign spiders with host S. sarasinorum spiders. We found spiders classified into nine families and insects classified into five orders and a few small vertebrate taxa associated with S. sarasinorum colonies. Adult S. sarasinorum nests and webs contained significantly more foreign spiders than juvenile colonies. However, diversity of foreign spiders did not increase with the number of host spiders. Additionally, we found that foreign spiders found inside the nests preyed on individual S. sarasinorum spiders. Our study sheds light onto the intriguing biology of nest- and web-associates of social spiders.