Termite Behavior Research Articles

Ecological correlates of chimpanzee termite fishing behavior in Mbam & Djerem National Park, Cameroon.

Chimpanzee insectivory is seasonally variable, with pronounced peaks or set seasonal periods of consumption observed in most chimpanzee communities. This variation is interesting given that chimpanzees invest considerable effort into complex tool-using behaviors to acquire insect prey. Evidence suggests this seasonal variation is related to insect behavior, but few studies have been done to empirically examine this relationship. In this study, we assessed whether a seasonal pattern of termite fishing by Nigeria-Cameroon chimpanzees (Pan troglodytes ellioti) in Mbam & Djerem National Park, Cameroon was driven by termite behavior. We measured termite presence and termite foraging activity monthly at seven termite mounds near Ganga Research Station from April 2022 to April 2023. Macroscopic fecal analysis and camera traps placed at each mound demonstrated termite fishing in this community occurred from March to June, with a rare smaller period of termite fishing in October 2021. Average monthly rainfall, average monthly temperature, and average monthly fruit availability were used to examine potential environmental factors that could impact termite fishing seasonality. Termite presence was significantly different between months with and without chimpanzee termite fishing (t-test, -6.569, p < .001). Termite presence was also significantly associated with average monthly rainfall (ANOVA, F = 13.9, p = .002, R 2 = .775). Termites in this region appear to respond to the transition from dry to wet seasons by moving closer to the soil surface. This corresponds with greater chimpanzee termite fishing, suggesting that termite accessibility may be driving seasonal variation in this behavior.

The function of OforOrco in the allogrooming behavior of Odontotermes formosanus (Shiraki) induced by Serratia marcescens Bizio (SM1)

Termites are consistently confronted with a complex microbial environment. In addition to the role of their innate immune system in resisting pathogen infection, social immune behavior also plays a significant role in helping termites withstand the stress caused by pathogenic microorganisms. The allogrooming behavior among different individuals is commonly observed in termites, and it plays a crucial role in the social immune interaction network. In the case of Odontotermes formosanus (Shiraki), Orco is specifically involved in detecting pheromones and volatile chemicals released by termites to communicate with each other. Nonetheless, the function of Orco in the social immunity remains unreported in O. formosanus. Consequently, in this study, we recorded the allogrooming behavior of O. formosanus workers under SM1 stress. The results indicated a significant increase in allogrooming behavior due to SM1 infection. The allogrooming behavior of workers under SM1 stress was significantly increased after the addition of soldiers. Compared with pronotum group treated by SM1, SM1 treatment of workers' heads significantly reduced the allogrooming behavior among workers. In addition, we found that SM1 could greatly increase the expression of OforOrco. Furthermore, interfering with OforOrco could markedly reduce the allogrooming behavior among workers under SM1 stress, and increase the mortality of worker under SM1 stress. This study demonstrated the significant role of OforOrco in the social immunity of O. formosanus, which offers a theoretical foundation for the advancement of research on termite RNA biopesticides, and the integration of RNA interference (RNAi) with pathogens. This study is valuable for elucidating the social immune behavior and interaction network of termites.

Termite females prefer hybrid mating to parthenogenesis when conspecific males are absent

Abstract Parthenogenesis and interspecific mating are alternative reproductive strategies that species have evolved alternative strategies when the opportunity of sexual reproduction is hindered by the natural environment. However, it is still unclear how to make a choice between parthenogenesis and interspecific mating when females are unable to reproduce sexually with males of the same species. This study mainly investigated the reproductive behaviour of female termites in the absence of male termites of the same species. Here, reproductive behaviours of females Reticulitermes aculabialis (Ra♀) were observed when they encountered a male R. flaviceps (Rf♂) with the absence of male R. aculabialis(Ra♂). The results showed that Ra♀‐Rf♂ tandems were more prevalent and stable compared to Ra♀‐Ra♀ tandems. When a tandem inadvertently broke apart, the female R. aculabialis displayed longer pauses while waiting for male R. flaviceps, but shorter pauses when another female R. aculabialis was involved. Furthermore, there were mating behaviour between female R. aculabialis and males R. flaviceps, resulting in offspring produced in paired Ra♀‐Rf♂ colonies were hybridization between female R. aculabialis and male R. flaviceps. These results suggest that interspecific mating and recombination is an important aspect for the choice of reproduction strategy, which will provide new insights into the trade‐off between gene contribution rate and genetic diversity.

Accessory gland protein regulates pairing process and oviposition in the subterranean termite Reticulitermes chinensis after swarming.

Swarming and pairing behaviors are significant to population dispersal of termites. Tandem running is a key process in pairing behavior of dealates to find a mate. Succinylation can lead to significant changes in protein structure and function, which is widely involved in metabolism and behavior regulation in many organisms. However, whether succinylation modification regulates termites' tandem running is currently unknown. In this research, we performed quantitative modified proteomics of the subterranean termite Reticulitermes chinensis Snyder before and after alate swarming. The succinylation levels of accessory gland protein (ACP) were significantly altered after alate swarming. We found that ACP is enriched in male accessory gland and female oocytes of termites. The acetylation and succinylation sites of ACP affected tandem running of dealates. The transcriptome and metabolome analyses of alates injected with ACP and its mutant proteins showed that β-alanine metabolism pathway was the major downstream pathway of ACP. Silencing the significantly differentially expressed genes in the β-alanine metabolic pathway (acyl-CoA dehydrogenase, enoyl-CoA hydratase, 3-hydroxyisobutyrate dehydrogenase, methylmalonate-semialdehyde dehydrogenase) suppressed tandem running and altered oviposition of paired dealates. These findings demonstrate that protein translation modification is an important regulator of tandem running behavior of termites, which implies that the succinylation and acetylation modification sites of ACP could be potential targets for insecticide action. Our research offers a potential approach for developing novel dispersal inhibitors against social insect pests.

Extinct and extant termites reveal the fidelity of behavior fossilization in amber

Fossils encompassing multiple individuals provide rare direct evidence of behavioral interactions among extinct organisms. However, the fossilization process can alter the spatial relationship between individuals and hinder behavioral reconstruction. Here, we report a Baltic amber inclusion preserving a female-male pair of the extinct termite species Electrotermes affinis. The head-to-abdomen contact in the fossilized pair resembles the tandem courtship behavior of extant termites, although their parallel body alignment differs from the linear alignment typical of tandem runs. To solve this inconsistency, we simulated the first stage of amber formation, the immobilization of captured organisms, by exposing living termite tandems to sticky surfaces. We found that the posture of the fossilized pair matches trapped tandems and differs from untrapped tandems. Thus, the fossilized pair likely is a tandem running pair, representing the direct evidence of the mating behavior of extinct termites. Furthermore, by comparing the postures of partners on a sticky surface and in the amber inclusion, we estimated that the male likely performed the leader role in the fossilized tandem. Our results demonstrate that past behavioral interactions can be reconstructed despite the spatial distortion of body poses during fossilization. Our taphonomic approach demonstrates how certain behaviors can be inferred from fossil occurrences.

Exploring the role of loop tunnels in enhancing termite food transport efficiency: A simulation approach

Subterranean termites construct a network of tunnels beneath the ground, comprising a main tunnel and branch tunnels. While termites do not construct tunnels that turn back in a “U” shape, the intersection of main and branch tunnels often forms closed curved structures (a loop). The loop structure can have either a positive or negative effect on the transport efficiency of termites in food transportation. However, little research has been conducted on the impact of loop tunnels on transport efficiency (E) due to the technical difficulties associated with direct observation. In this study, we used an individual-based model to simulate termite behavior during food transportation within a tunnel composed of a main tunnel and a loop tunnel. The model incorporates four control variables: the number of introduced simulated termites (k1), the probability of a simulated termite using a loop tunnel when walking towards a food or nest site (k2 or k3), and the length of the loop tunnel (k4). The simulation results reveal that the E value is high for (k2, k3) = (high, high), (low, low), (high, low), and (low, high) when the length of the loop tunnel is relatively short. However, when the length of the loop tunnel is relatively long, E is high only for (k2, k3) = (high, low) and (low, high). We found that these results are primarily influenced by the frequency of traffic jams. Therefore, termites would benefit from adopting strategies that reduce the occurrence of traffic jams during food transportation. In the Discussion section, we briefly touch on the strategy from an ecological perspective.

Oleic acid triggers burial behavior in a termite population through an odorant binding protein

Social insects maintain hygienic conditions through their social immunity behaviors. Among these behaviors, burial behavior of termites is central for protecting healthy individuals from corpses. Many factors trigger burial behavior, and it is generally believed that chemicals released by corpses, such as oleic acid, are the most important cues for triggering burial behavior in termites. However, the contribution of the olfactory system to this behavior remains unclear. Here we report an odorant binding protein (OBP) that transports oleic acid and triggers burial behavior in Coptotermes formosanus Shiraki. We demonstrated that CforOBP7 is highly expressed in the antennae of workers. Fluorescent competition binding experiments exhibited that CforOBP7 has a strong affinity for oleic acid. Furthermore, the antennal response to oleic acid was significantly reduced, and oleic acid-triggered burial behavior was also inhibited in CforOBP7-silenced termites. We conclude that CforOBP7 governs the burial behavior of C. formosanus triggered by oleic acid.

Neuroregulation of foraging behavior mediated by the olfactory co-receptor Orco in termites

Olfaction is critical for survival because it allows animals to look for food and detect pheromonal cues. Neuropeptides modulate olfaction and behaviors in insects. While how the neuroregulation of olfactory recognition affects foraging behavior in termites is still unclear. Here, we analyzed the change after silencing the olfactory co-receptor gene (Orco) and the neuropeptide Y gene (NPY), and then investigated the impact of olfactory recognition on foraging behavior in Odontotermes formosanus under different predation pressures. The knockdown of Orco resulted in the reduced Orco protein expression in antennae and the decreased EAG response to trail pheromones. In addition, NPY silencing led to the damaged ability of olfactory response through downregulating Orco expression. Both dsOrco- and dsNPY-injected worker termites showed significantly reduced walking activity and foraging success. Additionally, we found that 0.1 pg/cm trail pheromone and nestmate soldiers could provide social buffering to relieve the adverse effect of predator ants on foraging behavior in worker termites with the normal ability of olfactory recognition. Our orthogonal experiments further verified that Orco/NPY genes are essential in manipulating termite olfactory recognition during foraging under different predation pressures, suggesting that the neuroregulation of olfactory recognition plays a crucial role in regulating termite foraging behavior.

α-Terpineol affects social immunity, increasing the pathogenicity of entomopathogenic nematodes to subterranean termites (Isoptera)

Biocontrol of subterranean termites is largely impeded by their social immune responses. Studies on biocontrol agents combined with natural insecticides and their possible effects on the immune defense mechanisms of termites are limited. In this study, we investigated the effects of a combined biocontrol strategy using a plant-derived insect ATPase inhibitor, α-terpineol, with the entomopathogenic nematodes (EPNs) Steinernema carpocapsae against the subterranean termite Coptotermes formosanus Shiraki. Survival assays showed that even a low lethal concentration of α-terpineol significantly increased the EPNs-induced virulence in C. formosanus. α-terpineol treatment majorly inhibited the activity of Na+- K+- ATPase, which disturbed the EPNs-induced enhancement of locomotor activity and grooming behavior in termites treated with the combined strategy. Furthermore, the combination treatment had a synergistic inhibitory effect on innate immune responses in C. formosanus, which were measured as changes in the expression of immune-related genes and activities of immune system enzymes. In conclusion, α-terpineol can weaken the immune defense of termites against EPNs at low lethal concentrations, and is a suitable non-synthetic insecticide to prove the biocontrol efficiency of EPNs on C. formosanus. This study provides a theoretical basis and technical reference for a novel biocontrol strategy that promises to overcome the problems of host immune defense in termites.

Effects of soil moisture on tunneling, survivorship, and food consumption of the Formosan and eastern subterranean termites (Blattodea: Rhinotermitidae).

Soil moisture is a critical environmental factor for the survival and behavior of subterranean termites (family Rhinotermitidae). The invasive Formosan subterranean termite, Coptotermes formosanus Shiraki, and the native eastern subterranean termite, Reticulitermes flavipes (Kollar), co-occur in the southeastern United States, while R. flavipes is distributed in a wider geoclimatic range. Previous studies showed that subterranean termites preferred higher soil moisture levels for tunneling and feeding; however, the impacts of constant moisture remained to be characterized to understand their moisture tolerance. In this study, we hypothesized that different soil moisture regimes can alter termite foraging and survival, and that the effects differ between the two species. The tunneling activity, survivorship, and food consumption of termites were documented for 28 days with different sand moisture conditions ranging from no moisture to full saturation (0%, 1%, 5%, 15%, 25%, and 30%). We found that there were no significant differences in the responses between C. formosanus and R. flavipes. In both species, termites did not survive or tunnel with 0% moisture. Termites performed tunneling with only 1% sand moisture, although they did not survive for 28 days. A minimal of 5% sand moisture was required for survival, and there were no significant differences in survivorship, tunneling activity, or food consumption among moisture contents of 5-30%. The results suggest that subterranean termites are resilient to moisture extremes. Colonies can tolerate low moisture conditions in their foraging environment for extended times, which may allow them to tunnel and find new moisture sources for colony survival.

Trichoderma metabolites trigger aggregation behavior in Formosan subterranean termites (Coptotermes formosanus).

Our previous studies have shown that some Trichoderma fungi trigger aggregation behavior in Formosan subterranean termites, Coptotermes formosanus Shiraki. However, the mechanisms underlying the induction of termite aggregation by Trichoderma fungi remain unclear. Here, we found that the aqueous or acetone extract of Trichoderma asperellum Samuels, Lieckfeldt & Nirenberg and Trichoderma virens Pers. ex Fries isolated from the gut or cuticle of C.formosanus elicited significant termite aggregation in 2-choice tests. We then screened 9 Trichoderma metabolites (3-acetoxy-2-butanone, phenol, 3-ethoxypropionic acid, ethyl 2,4-dioxovalerate, diglycolic acid, d-valine, styrene, 3-aminopyridine, and hexyl acetoacetate) that triggered termite aggregation. Among them, phenol (1000μg/mL), 3-ethoxypropionic acid (10μg/mL), ethyl 2,4-dioxovalerate (1000μg/mL), and diglycolic acid (1000μg/mL) showed the strongest activities, triggering termite aggregation throughout the 24-h period. As T.asperellum and T.virens produce different metabolites that trigger aggregation behavior in termites, the mechanisms underlying the interaction between subterranean termites and Trichoderma fungi likely vary. Future studies are needed to test whether these chemicals can attract termites and increase bait consumption.

The root stubble loss due to foraging by the workers of Odontotermes horni on different crops under dryland agroecosystem.

The study was mainly focused on the foraging behavior of worker termites of O. horni on various crops grown under dryland agroecosystems and how they cause loss to root stubbles. The results showed that soybean had the highest loss to root stubbles at 88.62%, followed by maize at 73.72% and finger millet at 70.16%. In terms of partial root stubble loss, maize had the highest percentage at 26.54%, followed by finger millet at 21.82% and soybean at 3.01%. On the other hand, complete root stubbles eaten by the workers were more in soybean at 86.61%, followed by maize at 57.84% and finger millet at 55.85%.

Trophic Path of Marked Exuviae Within Colonies ofCoptotermes gestroi(Blattodea: Rhinotermitidae)

Nitrogen, a limiting growth factor in wood-feeding insects, was hypothesized to play a role in the recently discovered behavior of subterranean termites returning to the nest to molt. Coptotermes gestroi (Wasmann) exuviae is approximately 11% N by dry weight, and therefore a potentially rich source of recyclable nitrogen. Exuviae from a C. gestroi colony were marked with immunoglobulin G (IgG) and were fed to two-year-old C. gestroi colonies. IgG-marked exuviae were detected with an enzyme-linked immunosorbent assay. The IgG marker was later detected in every caste and life stage except first-instar larvae (L1). The proportion of individuals positive for the marker varied by caste, with the queens always being positive for the marker. The queens and second-or-higher-instar workers (W2+) had significantly higher concentrations of the marker than the eggs and L1. The trophic path of exuviae includes individuals that directly fed on marked exuviae (workers and possibly second-instar larvae) and individuals that secondarily received marked exuviae through trophallaxis (queens, kings, and soldiers). This study described the trophic path of consumed exuviae and demonstrated its role in the recycling of nitrogen in a subterranean termite. Molting at the central nest may be an efficient means to transfer nitrogen from shed exuviae to recipients and may be a nitrogen recycling behavior conserved from a termite ancestor.

Visibility graphs of animal foraging trajectories

The study of self-propelled particles is a fast growing research topic where biological inspired movement is increasingly becoming of much interest. A relevant example is the collective motion of social insects, whose variety and complexity offer fertile grounds for theoretical abstractions. It has been demonstrated that the collective motion involved in the searching behaviour of termites is consistent with self-similarity, anomalous diffusion and Lévy walks. In this work we use visibility graphs—a method that maps time series into graphs and quantifies the signal complexity via graph topological metrics—in the context of social insects foraging trajectories extracted from experiments. Our analysis indicates that the patterns observed for isolated termites change qualitatively when the termite density is increased, and such change cannot be explained by jamming effects only, pointing to collective effects emerging due to non-trivial foraging interactions between insects as the cause. Moreover, we find that such onset of complexity is maximised for intermediate termite densities.

Efficacy of leaf extracts of apple of sodom (Calotropis procera) for the control of subterranean termite Odontotermes Obesus

Despite serious safety concerns, Calotropis procera is used for gastrointestinal disorders. The subterranean termites Odontotermes obesus is extremely invading wood pests that cause direct economic losses worth billions of dollars throughout the world particularly in humid and subtropic regions. Leaf extract of Calotropis procera in two solvent hexane and acetone, is used against workers of subterranean termites, mortality. LC50 and tunneling behavior of termites was observed at different concentration of 3%, 7% and 10% respectively and distilled water applied a control. Two solvent hexane and acetone is used for extraction. Data have been analyzed using software Minitab (version 16).

The effect of ergosterol on the allogrooming behavior of termites in response to the entomopathogenic fungus Metarhizium anisopliae.

Termites have physiological and behavioral immunities that make them highly resistant to pathogen infections, which complicates biocontrol efforts. However, the stimuli that trigger the pathogen-avoidance behaviors of termites are still unclear. Our study shows that workers of Coptotermes formosanus exposed to the conidia of Metarhizium anisopliae exhibited a significantly higher frequency and longer duration of allogrooming behaviors compared with untreated termites. Volatile compounds in the cuticle of control termites and termites previously exposed to a suspension of M. anisopliae conidia were analyzed and compared using a gas chromatography-mass spectrometer (GC-MS). Our results showed that the amount of ergosterol differed between the fungus-exposed and control termites. Choice tests showed that termites significantly preferred to stay on filter paper treated with ergosterol (0.05, 0.1, or 1.0mg/mL) compared with control filter paper. In addition, termites exposed to ergosterol followed by M. anisopliae conidia were allogroomed at a significantly higher frequency and for a longer duration than termites exposed to alcohol (the solvent used with the ergosterol in the ergosterol trials) alone followed by M. anisopliae conidia. These results showed that ergosterol may enhance the allogrooming behavior of termites in the presence of entomopathogenic fungi.

A High Soldier Proportion Encouraged the Greater Antifungal Immunity in a Subterranean Termite.

Termites possess a mighty social immune system, serving as one of the key obstacles to controlling them biologically. However, the dynamic mechanism coordinating the social immunologic defense and caste distribution of the termites remains elusive. This study used the Coptotermes formosanus Shiraki and an entomopathogenic fungus as a host–pathogen system and experimentally manipulated a series of groups with different caste compositions of workers and soldiers. Then, the impact of demography on the behavior and innate immunity of termites was explored by analyzing the fungus susceptibility of the respective caste, efficiencies, and caste preferences of sanitary care, as well as the expression of the immune genes and phenoloxidase activity. Overall, to ensure the general health and survival of a group, the infected workers were found to sacrifice their survivorship for maintaining the soldier proportion of the group. If soldier proportion was limited within a threshold, both the survivorship of the workers and soldiers were not significantly affected by the infection. Correspondingly, the infected group with a higher proportion of soldiers stimulated the higher efficiency of a non-caste-biased sanitary care of the workers to the nestmate workers and soldiers. Moreover, the innate immunities of the infected workers were found to be more intensely upregulated in the group with higher soldier proportions. This suggested that the adjustable non-caste-biased sanitary care and innate immunity of the workers would contribute to the flexibility of the worker–soldier caste ratio in C. formosanus. This study, therefore, enhanced our understanding of the functional adaptation mechanism between pathogen-driven social immunity and the demography of the termites.

Detrimental Effects of Metabolic Gas Accumulation within Termite Bioassay Arenas

Effects of metabolic gas accumulation on termite behavior in tightly sealed laboratory bioassay arenas were compared with arenas with partial air exchange, or open ventilation. Laboratory studies with termites are often conducted in glass or plastic arenas. These studies can last from a few days up to 12-16 weeks or longer. Based on personal observation and published laboratory studies, how tightly study arenas are sealed is not consistent. In some instances arenas are tightly sealed with impermeable rubber stoppers or tight-fitting lids. In other studies, arenas are covered with semi-air-permeable Parafilm®, or loose-fitting aluminum foil or ventilated lids to allow air exchange. Generally, arenas are covered and not completely open to the air, thereby maintaining high moisture requirements needed to avoid termite dehydration. In this study, to determine metabolic gas accumulation and gas effects on termite groups collected from two field colonies, bioassay arenas were capped with either tightly sealed metal lids, semi-air-permeable Parafilm, or stainless steel mesh to allow non-restricted ventilation. Gas concentrations in these arenas and termite behavior and mortality were determined over time. During the study no visible ataxic termite behavior or significant mortality was observed within Parafilm or stainless-steel-mesh capped arenas. In contrast, tightly sealed arenas quickly accumulated significantly more metabolic gases compared with the other arena configurations, causing obvious abnormal termite behavior within 96 hours after these arenas were sealed. Within 8 to 10 days 100% moribund or dead termites occurred in tightly sealed arenas, whereas termites in semi-permeable membrane or non-restricted ventilated arenas were behaving normally. Excess gas accumulation in tightly sealed arenas first elicited negative effects between 96 and 144 hours. Our results show that tightly sealed bioassay arenas must be avoided, even for short-term studies lasting only a few days, as metabolic gases accumulate rapidly, causing negative effects on termites beginning within 4 to 6 days.

A Review of the Evolution of Termite Control: A Continuum of Alternatives to Termiticides in the United States with Emphasis on Efficacy Testing Requirements for Product Registration.

Simple SummaryTermites are structurally destructive pests that can infest a consumer’s most important investment, their home. The evolution of termite control is complex, involving people’s understanding of termite biology and behavior, building construction, the use of soil termiticides, baits, wood treatments, and physical barriers, as well as regulations from different industries (pest control and building construction). Soil termiticides have been the standard method of treatment for decades, but the concern for human and environmental health has driven the development of alternatives. This article discusses the evolution of termite control methods that were alternatives to the standard of the time and the regulatory structure that provides a level of consumer protection for some products.The global economic impact of termites is estimated to be approximately USD 40 billion annually, and subterranean termites are responsible for about 80% of the total impact. Twenty-eight species of termites have been described as invasive, and these termites are spreading, partially due to global trade, making effective control methods essential. Termite control is complex, as is the biology and behavior of this social insect group. In the U.S., termite prevention and control (with claims of structural protection) is regulated by more than one industry (pest control and building construction), and at the federal and state levels. Termite prevention has historically relied on building construction practices that do not create conducive conditions for termite infestations, but as soil termiticides developed, heavy reliance on pesticides became the standard for termite control. The concern for human and environmental health has driven the development of termite control alternatives and regulation for products claiming structural protection. Product development has also provided unprecedented opportunities to study the biology and behavior of cryptobiotic termites. Technological advances have allowed for the re-examination of questions about termite behavior. Advances in communications via social media provide unrestricted access to information, creating a conundrum for consumers and science educators alike.

Differential time allocation of foraging workers in the subterranean termite

BackgroundForaging in group living animals such as social insects, is collectively performed by individuals. However, our understanding on foraging behavior of subterranean termites is extremely limited, as the process of foraging in the field is mostly concealed. Because of this limitation, foraging behaviors of subterranean termites were indirectly investigated in the laboratory through tunnel geometry analysis and observations on tunneling behaviors. In this study, we tracked subsets of foraging workers from juvenile colonies of Coptotermes formosanus (2-yr-old) to describe general foraging behavioral sequences and to find how foraging workers allocate time between the foraging site (food acquisition or processing) and non-foraging site (food transportation).ResultsOnce workers entered into the foraging site, they spent, on average, a significantly longer time at the foraging site than the non-foraging site. Our clustering analysis revealed two different types of foraging workers in the subterranean termite based on the duration of time they spent at the foraging site and their foraging frequency. After entering the foraging site, some workers (cluster 1) immediately initiated masticating wood fragments, which they transferred as food boluses to recipient workers at the foraging site. Conversely, the recipient workers (cluster 2) moved around after entering the foraging site and received food from donating workers.ConclusionsThis study provides evidence of task specialization within foraging cohorts in subterranean termites.