The surprising vulnerability of a termite’s secret roomate

By Igor Eloi

Igor is a PhD student based at the UFRN campus on the brazilian coast. He is fascinated by how social insects, like termites, behave and interact with other species. In this blog, he shares key insights from a research paper exploring how fragment edges affect termite guests. His lastest research in Insectes Sociaux can be read here.

A termite nest is more than a mound of earth and wood; it’s a bustling city, a climate-controlled fortress engineered and built by tiny insects. These complex structures are not just homes for termites, a rather exquisite diversity of organisms have evolved to life within their walls.

There residents are known as”termitophiles”—organisms that live their entire lives, or at least critical parts of them, in an obligatory relationship with termite society. They are not merely guests but are deeply integrated into the colony’s day-to-day routine. These creatures have evolved alongside their hosts for millennia, developing bizarre forms and behaviors to survive and thrive inside the fortress. Which raises a rather pertinent question: If a creature is perfectly adapted to live inside a protective, self-regulating termite nest, does that make it immune to changes in the outside world? In other words, what happens to these hidden, highly specialized residents when human activity, like a simple dirt road, encroaches on their world? Our study set out to find the answer, revealing just how far the ripples of habitat disturbance can travel.

We focused on studying two Aleocharinae beetles, that live in an asymmetric “obligatory relationship” with a their host. This means that while termites live fine without the beetles, the beetles themselves cannot survive without the colony. We examined two distinct types found in the arboreal nests of the termite Constrictotermes cyphergaster (Nasutitermitinae).

Lateral (left) and dorsal (right) views of the termite Constrictotermes cyphergaster (Nasutitermitinae), host species of the studied Aleocharinae beetles.

On the image below, the first, Termitocola silvestrii, is a miniature tank. This species is equipped with a “limuloid” or drop-shaped body, featuring a large, shield-like pronotum thought to be a defensive adaptation against termite attacks. Anecdotal observations suggest it may act as part of the colony’s cleanup crew, feeding on dead termites. While these beetles possess wings, researchers speculate they may lose the ability to fly after successfully settling within a host colony. This secret society has its rules, and both species rely on momentarily leaving the nest—either for reproduction or dispersal—exposing their hidden world to the conditions of the wider forest.

Lateral and dorsal views of the rove beetle Termitocola silvestrii, a limuloid (drop-shaped) Aleocharinae species associated with the termite Constrictotermes cyphergaster. The large, shield-like pronotum is thought to provide protection against termite attacks.

The second, Corotoca fontesi, has a bizarre “physogastric” body, with a swollen, soft abdomen that gives it a strange, almost larval appearance and reduces its mobility. Its life cycle is a drama of dependence and risk. To reproduce, the female must venture outside the nest during the termites’ open-air foraging expeditions (Moreira et al. 2019). She then deposits a single, motile larva into the ground litter. The larva develops alone in the soil, and how it later finds and integrates into a new host nest remains one of the fascinating mysteries of its life cycle.

Lateral and dorsal views of the rove beetle Corotoca fontesi, a physogastric Aleocharinae species associated with the termite Constrictotermes cyphergaster. The swollen, soft abdomen gives the beetle a larval-like appearance.

The central finding of our study is that despite living inside the protective, climate-controlled environment of a termite nest, the abundance of these specialized beetles is negatively impacted by proximity to a forest edge. This finding demonstrates that the so-called “edge effect”—the ecological changes that occur where two habitats meet—penetrates the defenses of the termite fortress.

One might assume that the nest would act as a perfect buffer against external environmental stressors. However, the study’s results suggest otherwise, highlighting that even for organisms living deep within a host structure, the human-made landscape changes of the outside world matters immensely.

Finally, it is our thought that the mechanisms behind the impact of edge effect over the abundance of termitophiles might lie in one (or the combination) of these:

  1. Direct Impact: The harsher environmental conditions at the forest edge—such as different temperatures or humidity—could directly harm the beetles during the parts of their life cycle spent outside the nest. For example, the larvae of Corotoca developing in the soil could be exposed to increased predation or unsuitable microclimates (Zilberman et al. 2019).
  2. Host-Mediated Impact: The termite colonies themselves might be stressed by the edge conditions. This could make them “lower-quality hosts,” perhaps with fewer resources or a smaller workforce, rendering them unable to support large populations of their beetle symbionts.
  3. Dispersal Limitation: The altered landscape near the road could act as a barrier. This might make it more difficult for adult beetles to travel between nests, limiting their ability to find and colonize nests located near the forest edge.

References:

Moreira IE, Pires-Silva CM, Ribeiro KG, et al (2019) Run to the nest: A parody on the Iron Maiden song by Corotoca spp.(Coleoptera, Staphylinidae). Papéis Avulsos De Zoologia 59:e20195918–e20195918.

Siqueira-Rocha, L., Eloi, I., A Luna-Filho, V. et al. Aleocharinae termitophiles are affected by habitat fragmentation in deciduous dry forests. Insect. Soc. (2026). https://doi.org/10.1007/s00040-025-01076-4

Zilberman B, Pires-Silva CM, Moreira IE, et al (2019) State of knowledge of viviparity in Staphylinidae and the evolutionary significance of this phenomenon in Corotoca Schiødte, 1853. Papéis Avulsos De Zoologia 59:e20195919–e20195919. https://doi.org/10/gng3q8

Insect architects: How termites, ants, and bees build without blueprints

by Paige Caine

Paige Caine is a PhD student working in Dr. Michael Goodisman’s lab at Georgia Tech. She studies fire ants and yellowjacket wasps. In this blog, she explains how social insects, such as termites, ants, or bees, collectively manage to build complex nests. Her latest research on social insects can be read here.

A builder stands at the foot of her construction, a massive skyscraper towering thousands of times her height. The imposing architectural feat stretches stories underground as well, and is home to thousands of individuals. She can’t see the results of all her hard work though; she’s blind. In fact, the entire team of builders responsible for the structural triumph is blind, and they didn’t have a chief architect or any blueprints to guide them. How did they do it?

To answer this question, let’s meet the construction crew: Cathedral Termites. Native to Australia, this species of termites has blind workers measuring only about 3-4.5 millimeters long, yet they build massive nests to house their queens, kings, and young. 

A Cathedral Termite (Nasutitermes triodiae) mound.

But this feat isn’t unique to Cathedral Termites—most social insects construct some form of nest. These structural marvels range in size and shape, from Cathedral Termite mounds to charismatic honeybee hives to tiny ant homes contained within acorns.  In the absence of realtors, social insects often use collective decision-making to choose a nest location that optimizes temperature, sunlight, precipitation level, predation risk, and proximity to resources (Jeanne and Morgan 1992; London and Jeanne 2000; Suzuki et al. 2007). These strategies typically involve sending a few scouts to locate potential nesting sites. The scouts then recruit colony-mates to “vote” on sites by physically going to that site and contributing to the recruitment effort. Eventually, a quorum is reached, and the losing party packs up from their rejected sites and heads to the winning location (Pratt 2005).

Once the site has been chosen, a range of different construction methods are used to build the nest. Termites and ants tend to excavate their homes, while social bees and wasps tend to build their homes from manipulated biological material—chewed up wood pulp in the case of social wasps or wax in the case of some bees.

A social wasp nest from the yellowjacket Vespula squamosa. While these structures are built underground, this nest has been excavated (left), and then separated into the individual layers of comb (right).

A common problem during collective construction—and one most human commuters are accustomed to—is crowding. To excavate a massive structure composed of tunnels and chambers, ants and termites must navigate narrow spaces containing hundreds or even thousands of individuals. One way termites solve this problem is through something referred to as a “bucket brigade;” like humans passing water towards a fire via a series of buckets, some termite species form a queue and pass excavated material along from individual to individual until it reaches the deposition site (Bardunias and Su 2010). Some ants, on the other hand, utilize “laziness” to avoid crowding, by having certain individuals sit still while a minority actually contributes significantly to construction (Aguilar et al. 2018).

But, if there’s no blueprint and no architect in charge of doling out specific tasks, how are all these individual construction behaviors coordinated?

One common means of coordination is stigmergy, which means communicating across time via the environment. Each time an individual interacts with the incipient construction, they leave behind traces of their behavior, either by shaping the material or leaving behind chemicals. These cues tell individuals who later approach the construction what has been done, and what’s left to do.

A diagram displaying stigmergy at work in honeybee nest construction, based on (Nazzi 2016). Different colored bees indicate distinct individuals A) Bee #1 interacts with cells constructed by her nestmates. B) Cells act as cues for bee #1 to extend the floor of the nest. C) Floor acts as a cue for bee #2 to begin constructing stubs of a wall. D) Third bee detects these wall stubs and responds by adding to the stubs to encircle a cell. E) Fourth bee notices cells constructed by her nestmates and responds by extending the floor further. F) After additional building activity by several bees, the comb of cells hits the edge of the cavity.

Now that we know how social insects build their remarkable nests, another natural question is why?

Social insect nests offer many advantages to residents. For one, they offer protection from weather, much like a human home. They also protect against infection, with many species actively incorporating antimicrobial bacteria or other antibiotic agents into the walls (Tranter et al. 2013; Madden et al. 2013; Chouvenc et al. 2013). Nests also enable protection against larger threats, functioning as defendable fortresses. In fact, many species employ guards at nest entrances, and often close their doors at night (Bennett and Baudier 2021).  Finally, nests help large insect societies organize their behaviors by physically contributing to division of labor, as well as by influencing the efficiency of collective tasks like foraging.

Termites nesting in paper (left), which they eat as they construct. Ants nesting in soil (right), showcasing some of the many interior tunnels and chambers.

Overall, social insect nest construction is an impressive feat, and the results are both structurally remarkable and highly functional. One day, we may be able to imitate such techniques using swarm robotics. Today, many engineers are already working on bio-inspired robot collectives capable of construction. Robotic models are even being designed to test hypotheses about collective behaviors in social insect groups, an approach recently termed “robophysics.” In the future, robophysical models may unlock some of the principles underlying social insect nest construction, strengthening our understanding of collective behavior in both engineering and biology.

Robots engaged in construction. Left photo credit: Eliza Grinnell/Harvard SEAS. Right photo credit: Daniel Soto, Joonha Hwang

References:

Aguilar J, Monaenkova D, Linevich V, et al (2018) Collective clog control: Optimizing traffic flow in confined biological and robophysical excavation. Science 361:672–677. https://doi.org/10.1126/science.aan3891

Bardunias PM, Su NY (2010) Queue Size Determines the Width of Tunnels in the Formosan Subterranean Termite (Isoptera: Rhinotermitidae). J Insect Behav 23:189–204. https://doi.org/10.1007/s10905-010-9206-z

Bennett MM, Baudier KM (2021) The Night Shift: Nest Closure and Guarding Behaviors in the Stingless Bee, Tetragonisca angustula. J Insect Behav 34:162–172. https://doi.org/10.1007/s10905-021-09779-9

Caine, P.B., Robertson, A.T., Treers, L.K. et al. Architecture of the insect society: comparative analysis of collective construction and social function of nests. Insect. Soc. (2025). https://doi.org/10.1007/s00040-025-01057-7

Chouvenc T, Efstathion CA, Elliott ML, Su N-Y (2013) Extended disease resistance emerging from the faecal nest of a subterranean termite. Proceedings of the Royal Society B: Biological Sciences 280:20131885. https://doi.org/10.1098/rspb.2013.1885

Jeanne RL, Morgan RC (1992) The influence of temperature on nest site choice and reproductive strategy in a temperate zone Polistes wasp. Ecological Entomology 17:135–141. https://doi.org/10.1111/j.1365-2311.1992.tb01170.x

London KB, Jeanne RL (2000) The interaction between mode of colony founding, nest architecture and ant defense in polistine wasps. Ethology Ecology & Evolution https://doi.org/10.1080/03949370.2000.9728440

Madden AA, Grassetti A, Soriano J-AN, Starks PT (2013) Actinomycetes with Antimicrobial Activity Isolated from Paper Wasp (Hymenoptera: Vespidae: Polistinae) Nests. Environ Entomol 42:703–710. https://doi.org/10.1603/EN12159

Nazzi F (2016) The hexagonal shape of the honeycomb cells depends on the construction behavior of bees. Sci Rep 6:28341. https://doi.org/10.1038/srep28341

Pratt SC (2005) Quorum sensing by encounter rates in the ant Temnothorax albipennis. Behav Ecol 16:488–496. https://doi.org/10.1093/beheco/ari020

Suzuki Y, Kawaguchi LG, Toquenaga Y (2007) Estimating nest locations of bumblebee Bombus ardens from flower quality and distribution. Ecol Res 22:220–227. https://doi.org/10.1007/s11284-006-0010-3

Tranter C, Graystock P, Shaw C, et al (2013) Sanitizing the fortress: protection of ant brood and nest material by worker antibiotics | Behavioral Ecology and Sociobiology. Behavioral Ecology and Sociobiology 68:499–507. https://doi.org/10.1007/s00265-013-1664-9