Interview with a social insect scientist: Stefano Cavallo

Stefano is a biologist specializing in animal behavior and currently works as a research fellow in behavioral ecology at the University of Florence. In this interview, he recalls moment he realized that even ants show individual personalities. His lastest research in Insectes Sociaux can be read here.

IS: Who are you, and what do you do?

I’m Stefano Cavallo, a passionate biologist specialized in animal behaviour. I’m living in Pisa and currently work at the University of Florence as a research fellow in behavioural ecology. My interests range from communication and cognitive aspects of animal behaviour in invertebrates and beyond. At the moment, my project focuses on exploring phenotypic plasticity—particularly behavioural plasticity—in marine decapods.

IS: How did you develop an interest in your research?

Since I was a child, I’ve always been passionate about animals. Although I grew up in a city, I had the chance to keep and observe a variety of species—fish, amphibians, reptiles, birds, mammals, and of course, insects. Among them, social insects, and especially ants, have always fascinated me. Their remarkable social organization combined with apparent simplicity sparked both curiosity and deep biological admiration in me. As my studies in biology progressed, I developed a strong interest in behavioral biology. What I find most stimulating is the possibility of identifying similar behavioral patterns in evolutionarily distant species, both human and non-human.

IS: What is your favorite social insect, and why?

It’s hard to choose just one. I’m fascinated by social insects for very different reasons: for instance, the interspecific relationships of Atta ants, the communicative flight and cognitive abilities of Apis mellifera, and the complex social structure of Polistes dominula all capture my interest. What I find most stimulating is not a single species, but rather those organisms capable of challenging the “dogmas” of biology. For example, the recent discovery by Juvé et al. (2025) on Messor ibericus which destroy species definitions.

IS: What is the best moment/discovery in your research so far? What made it so memorable?

One of the best moments in my research was when I first realized that even ants—creatures we often think of as identical and mechanical—show individual personalities. That realization was unforgettable: it felt like discovering a hidden layer of complexity within a familiar world. From that moment on, I stopped seeing colonies as uniform units and started seeing them as societies of individuals.

IS: Do you teach or do outreach/science communication? How do you incorporate your research into these areas?

No, at the moment I don’t deal with these aspects but in the future I hope it can become part of my job as a scientist. I think it is important to disseminate scientific advances to a wide audience and shorten the distances between laboratories, research and the general public.

IS: What do you think are some of the important current questions in social insect research, and what is essential for future research?

As we know, the environment today is subject to strong anthropogenic pressures and global warming is shaping habitats very quickly. The effects on social insects are still poorly understood. I believe it is essential to focus on these aspects and understand how changing conditions act on the biology and behavior of social insects.

IS: Outside of science, what are your favorite activities, hobbies, or sports?

I love being in nature, trekking in the mountains, climbing, swimming and snorkelling

IS: What is the last book you read? Would you recommend it? Why or why not?

The last book I read was Entangled Life: How Fungi Make Our Worlds, Change Our Minds and Shape Our Futures by Merlin Sheldrake. I would definitely recommend it—it’s a fascinating and beautifully written synthesis of what we know about fungi. These organisms are extraordinary in the way they challenge traditional paradigms of biology and reveal how deeply interconnected life really is.

IS: How do you keep going when things get tough?

I practice tai-chi and mindfulness techniques to stay in the present moment and focus on beautiful things.

IS: If you were to go live on an uninhabited island and could only bring three things, what would you bring? Why?

I would bring a knife, a tinderbox and a book on edible plants. These three things would help me get food, be able to cook and warm up and not die of intoxication haha!

IS: Who do you think has had the most considerable influence on your science career?

I believe that the most important role was played by two high school teachers. My chemistry professor and biology professor taught me scientific rigor and wonder at the living world

IS: What advice would you give to someone hoping to be a social insect researcher in the future?

If I had to advise someone to hop to be a social insect researcher, I would tell them to follow the thirst for knowledge and not stop at appearances. I would ask him to always look with a critical eye at those who claim to have absolute certainties in biology.

IS: Has learning from a mistake ever led you to success?

I couldn’t point to a specific mistake, but I believe that in private life and at work we often learn by falling and making mistakes. Trying by trial and error: this is generally just how we manage to grow.

IS: What is your favorite place science has taken you?

My favourite place where science took me is Paris, in the experimental and comparative ethology laboratory of the Sorbonne University in northern Paris. I was lucky enough to work in the group led by Professor Patrizia d’Ettore who with dedicated passion dedicates herself to research in the myrmecological field.

Pesticides damage bee parenting — and their larvae pay the price

By Leeah Richardson

Leeah is a graduate student at the University of Texas at Austin with a particular interest in insect behavior and anthropogenic stressors. In this blog, she explains how, although not highly lethal to adults, some pesticides can harm bumblebees in indirect ways. Her latest research on social insects can be read here.

Worldwide we use billions of pounds of pesticides each year agriculturally to control crop pests (FAO 2024) but this negatively impacts many insects that benefit crop production – for example: pollinators. Regulatory agencies do try to minimize the impact pesticides have on pollinators, but this is largely by preventing lethal effects. Pesticides don’t always have to kill bees to harm their populations and the pollination services they provide. By asking how much of a chemical is lethal to adults, we can miss subtle yet important effects on bee behavior and reproduction.

For many animals, including most bee species, offspring receive care from adults in order to survive. For example, in a bumblebee colony, worker bees chew through the wax covering their larval sisters to feed them pollen and nectar, regulate the temperature of the colony, and perform hygienic behaviors, so that these larvae can develop into adult bees. These caretaking behaviors are critical, but can be vulnerable to environmental stressors, like exposure to insecticides.

Bumblebee (Bombus impatiens) workers displaying caretaking behaviors on a brood mass.

The insecticide Flupyradifurone (FPF) is especially interesting to study in the context of how it may influence bee caretaking behaviors FPF is not likely to outright kill adult honeybees or bumblebees at the concentrations present agriculturally – so it can be sprayed on flowering crops, but recent studies have shown that it has negative effects on bumblebee larvae (Fischer et al. 2023, Richardson et al. 2024). This raises the question: are bee larvae themselves sensitive to FPF, or is the problem that exposed adults provide poorer care to developing larvae?

We conducted two experiments to determine whether FPF is directly toxic to larvae through ingestion or if FPF has indirect effects by impairing caretaking behaviors (below).

FPF could influence larvae directly (due to ingestion) or indirectly (by impairing caretaking behaviors provided by the adults).

We first did an experiment where we fed larvae by hand so that parental care was completely standardized for all of the larvae. To do this, we took larvae from an existing colony and kept each larva in an individual well of a 24-well plate, provisioning them with a sugar water/pollen mixture four times per day for three days. This mixture was either untreated (control) or contained FPF at one of four concentrations. If FPF was directly toxic, we expected to see higher mortality or delayed molting with the treated larvae. Instead, we found no differences between our untreated control groups and the treated larvae.

Process of removing all larvae from an existing colony under red light, then sorting them into size categories (instars), then placing them into individual wells in 24-well plates to be hand fed a sugar water/pollen mixture.

We then conducted a cross-fostering experiment to test for both direct and indirect effects to larvae. We created small “microcolonies” with four worker bees each that begin laying eggs and rearing offspring once separated from their queen. Half of these microcolonies were given FPF treated sugar water for two weeks, while the other half got untreated (control) sugar water.

Bumblebee microcolony with three worker bees on the brood mass they produced. Larvae are kept under the wax covering, so at the end of the experiment we peel this back to count and weigh them.

After two weeks, we swapped the adults to new microcolonies so that we now had groups of larvae that had never been exposed to FPF being cared for by FPF-treated adults, and groups of larvae that had been exposed to FPF being cared for by untreated (control) adults for three days (see figure below). This design allowed us to see whether larval outcomes depended on what the larvae themselves had previously ingested or on the status of their caretakers.

Design of our cross-fostering experiment to decouple adult and larvae exposure during the second phase and we found that whether the adults had been treated during phase 1 most influenced the size of the larvae we recovered from the microcolonies (adapted from Richardson et al. 2025).

We expected that if FPF directly impaired the larvae then whether or not the larvae themselves had been exposed to FPF for the two weeks prior to cross-fostering would strongly influence larval outcomes, but if indirect effects due to impaired parental care was most important then whether or not the adult caretakers had been exposed to FPF would instead be most influential. We found that it was the adult exposure to FPF that had the largest impact on larval outcomes (particularly the size of the larvae). Larvae tended by FPF-exposed adults were consistently smaller than those cared for by untreated adults, regardless of whether the larvae themselves had ingested FPF previously.

Both our hand-feeding and cross-fostering experiments showed that the larvae were surprisingly tolerant to direct FPF exposure via ingestion, but they were highly sensitive to impaired care. Together, these findings suggest that FPF’s harm to bumblebee larvae is driven mainly by changes in adult behavior, not by direct toxicity to the young.

Bee declines are complex, driven by habitat loss, climate change, disease, and pesticides (Goulson et al. 2015). Our study highlights the importance of testing not just whether pesticides kill adults, but also whether they disrupt the social and parental behaviors that larvae depend on. Future work should extend these kinds of experiments across more bee species and under field conditions, where multiple stressors interact.

References:

Fischer, L. R., Ramesh, D., & Weidenmüller, A. (2023). Sub-lethal but potentially devastating—The novel insecticide flupyradifurone impairs collective brood care in bumblebees. Science of The Total Environment, 903, 166097. https://doi.org/10.1016/j.scitotenv.2023.166097


Goulson, D., Nicholls, E., Botías, C., & Rotheray, E. L. (2015). Bee declines driven by combined stress from parasites, pesticides, and lack of flowers. Science, 347(6229), 1255957. https://doi.org/10.1126/science.1255957


Pesticides use and trade. 1990–2022. (2024). Food and Agricultural Organiation of the United Nations. https://www.fao.org/statistics/highlights-archive/highlights-detail/pesticides-use-and-trade-1990-2022/en


Richardson, L. I., DeVore, J., Siviter, H., Jha, S., & Muth, F. (2025). Bumblebees exposed to a novel ‘bee-safe’ insecticide have impaired alloparental care and reproductive output. Insectes Sociaux. https://doi.org/10.1007/s00040-025-01054-w


Richardson, L. I., Siviter, H., Jha, S., & Muth, F. (2024). Field‐realistic exposure to the novel insecticide flupyradifurone reduces reproductive output in a bumblebee (Bombus impatiens). Journal of Applied Ecology, 61(8), 1932–1943. https://doi.org/10.1111/1365-2664.14706

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

by Paige Caine

Paige Caine is a PhD student in Dr. Michael Goodisman’s lab at Georgia Tech. She study fire ants and yellowjackets 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

At what point does a male social wasp leave his natal nest to reproduce?

By Daniela Torres Garcia

In this blog, Daniela Torres Garcia, from the University of São Paulo, describe how she discovered that the number of females in a Mischocyttarus cerberus wasp nest influences the departure of males for mating. This latest research on social insects can be read here.

In social hymenopterans, male reproductive success depends entirely on the timing of reproduction, as males play no role in maintaining the colony—at least in the widely studied species. Males of many species, including social wasps, undergo post-pupal sexual maturation within the natal nest before dispersing to mate, during which time they rely on their nestmates for protection and food.

This leads us to a key question: do all males leave the nest at the same time, or is there something that makes some of them stay longer or shorter in their natal nest?

To answer this, we observed a population in southeastern Brazil of the Neotropical species Mischocyttarus cerberus. We conducted a rigorous monitoring of the nests of this species over several weeks to track male dispersal, and we found that the time a male spends in the nest before leaving varies. Some males leave the day after emerging, while others remain for almost a week.

Nest of M. cerberus with females and males. Photo by Andres Rodrigues De Souza.

Given this variability, we asked what factors might be influencing male dispersal timing. Does the social context affect this variability? That is, does the number of adult females in the nest influence how long the males stay? Do males stay longer when more adult females are present?

We addressed these questions using two approaches: on the one hand, observationally, by monitoring 36 natural nests; on the other hand, experimentally, by manipulating the number of females in 22 nests to see whether this caused a change in male dispersal behavior. And what did we find? Males in nests with more females stayed longer, thereby delaying their dispersal.

On average, males left after 3 days, but some took up to 8 days. We found that in nests with three females, males stayed for about 2.8 days, whereas in nests with only one female, they left after just 1.7 days. This suggests that females modulate male dispersal, which can last up to 8 days—similar to another social wasp, Polistes lanio (up to 7 days) (Southon et al., 2020). Why? Probably because staying in the nest is safer and more comfortable. More females mean better defense against predators and more food available. It is worth remembering that the sting—the primary defense mechanism of this group—is associated with the female reproductive system and thus is absent in males.

Male M. cerberus resting on the underside of a leaf within the study area. Photo by Andres Rodrigues De Souza.

Therefore, it is not surprising that males from nests with more females delay their dispersal to complete their sexual maturation in a safer and more comfortable environment, thereby increasing their survival and future reproductive competitiveness (i.e., by accumulating energy reserves). The accumulation of these reserves could help them avoid having to expose themselves on flowers to obtain food once dispersed.

Taken together, these results highlight the role of social context in shaping male reproductive strategies and suggest that pre-dispersal social life may be an underestimated factor in the physical fitness of males in social insects.

The reproductive biology of male social insects has often been studied at mating sites, such as leks and swarms (Beani et al., 1992; Beani et al., 2014). However, less attention has been given to male behavior prior to reaching these sites (e.g., Southon et al., 2020), despite its potential to influence male competitive ability. Therefore, pre-dispersal social life may be an overlooked aspect of male paper wasps’ reproductive strategies.

Left: Researcher tagging M. cerberus males for tracking, under an air conditioning unit. Right: M. cerberus nest under study, with several workers visible on the cells

References

Beani L, Dessì-Fulgheri F, Cappa F, Toth A (2014) The trap of sex in social insects: from the female to the male perspective. Neurosci Biobehav Rev 46:519–533. https://doi.org/10.1016/j.neubiorev.2014.09.014

Beani L, Cervo R, Lorenzi CM, Turillazzi S (1992) Landmark-based mating systems in four Polistes species (Hymenoptera: Vespidae). J Kansas Entomol Soc 8:211–217 https://www.jstor.org/stable/25085358

Garcia, D. T., Santos, E. F., Santos, S. A., do Nascimento, F. S., Krams, I., Rantala, M. J., & de Souza, A. R. (2025). Social context predicts male dispersal in nests of a paper wasp. Insectes Sociaux, 1-4. https://doi.org/10.1007/s00040-025-01050-0

Southon, R. J., Radford, A. N., & Sumner, S. (2020). Hormone-mediated dispersal and sexual maturation in males of the social paper wasp Polistes lanioJournal of Experimental Biology223(23), jeb226472.

What makes a queen successful?

By Luisa M. Jaimes-Nino

Luisa is a researcher at Johannes Gutenberg University Mainz. She specializes in studying the life-history traits of ants, their senescence, and the genetic and non-genetic mechanisms influencing queen fitness. Read her latest article in Insectes Sociaux here.

Queens, and kings in termites, are highly fertile and long lived insects. But certainly there is variation to which extent they are fertile. We were intrigued to understand what causes variation in fitness traits?

A Cardiocondyla obscurior queen (right), and brood and workers (left). ©Laure-Anne Poissonier.

We wondered if very fertile queens were also more successful by producing very fertile daughters. Is this caused by a genetic factor or perhaps the maternal status, such as maternal age? Old mothers might produce less fit queens and workers, and this can have a detrimental effect for the future of the whole colony. The negative effect of parental age, known as Lansing effect, has been documented across a wide range of taxa but not yet investigated in social insects.

We used Cardiocondyla obscurior ants as model given their high variability in fertility and  longevity. We investigated how fertility varies by selecting mothers that produced a low and high number of eggs after 15 weeks. We profited from a study in which we monitored a batch of queens in controlled conditions for their entire life (Jaimes-Nino et al., 2022), and monitored their daugthers too, to test if they presented a similar fertility and longevity.

In this experimental study, each daughter queen was mated to her wingless brother, kept in a single-queen colony on a plaster nest, and monitored monthly for productivity (i.e., number of eggs and pupae produced). ©LM Jaimes-Nino.

Our model species is polygynous, meaning that a large number of queens cohabit within a single colony. However, it is known that those queens that live longer, produce also more eggs (Kramer et al., 2015) and more queen daugthers (Jaimes-Nino et al., 2022). Previous studies have shown that queens remain heathly for a long portion of their lives because their mortality rate and gene expression pattern remain stable until old age (Harrison et al., 2021; Jaimes-Nino et al., 2022). Therefore, it is important to test whether old queens produce daugthers that are equally fit compared to those of younger queens, given that they produce the majority of them within the colony!

A) Wingless male and queen, B) three chambered plexiglas insert covered by plastic foil served as nest, and C) queen (right) and (worker) with brood inside the artificial nest chamber. ©Photo LM Jaimes-Nino.

Our results showed that mothers and daugthers do not “look” alike —  they do not have a similar fertility or longevity. This could indicate that their genetic background does not account for the observed variation. This can be expected from C. obscurior, as it exhibits extreme inbreeding.

Furthermore, contrary to the Lansing effect reported in other taxa, we found that daugthers produced by old mothers were just as fit as those produced by young mothers. This aligns with the hypothesis that, since the majority of sexuals are produced later in life, there must be mechanisms in place to maintain the health of these queens as they age! We believe that selection against aging remains strong in older queens. The specific mechanisms by which C. obscurior queens are able to produce equally fit daugthers at such advanced ages awaits further investigation.

Wingless (ergatoid) males with long mandibles mate in the nest with closely related gynes. ©Laure-Anne Poissonier.

Strikingly, the maternal lines differed in productivity suggesting background variation influenced by the maternal environment or male quality. In this species, ergatoid males (worker-like males) figth against rivals to monopolize queen access. Our study offers new avenues of research, to disentangle the effect of mother, father and developmental environment, on the final reproductive success of queens.

References

Harrison, M.C., Jaimes Niño, L.M., Rodrigues, M.A., Ryll, J., Flatt, T., Oettler, J., et al. 2021. Gene Coexpression Network reveals highly conserved, well-regulated anti-ageing mechanisms in old ant queens. Genome Biology and Evolution 13: 1–13. https://doi.org/10.1093/gbe/evab093

Jaimes-Nino, L.M., Heinze, J. & Oettler, J. 2022. Late-life fitness gains and reproductive death in Cardiocondyla obscurior ants. eLife 11: 1–17. https://doi.org/10.7554/eLife.74695

Kramer, B.H., Schrempf, A., Scheuerlein, A. & Heinze, J. 2015. Ant colonies do not trade-off reproduction against maintenance. PLoS ONE 10: 1–13. https://doi.org/10.1371/journal.pone.0137969

How does a caterpillar use its tentacles to get the attention of ants?

By Amalia Ceballos-González

In this blog, Amalia from the University of São Paulo tells the story of how she and her colleagues studied a strange functional behaviour in a myrmecophilous riodinid caterpillar. Read her latest article in Insectes Sociaux here.

Caterpillars that establish close interactions with ants have developed various adaptations to maintain the ants’ attention. These adaptations involves specialized organs that produce nutritional rewards or chemical signals to attract ants. The butterfly families Lycaenidae and Riodinidae provide many examples of myrmecophilous caterpillars, including species with these organs. In our recent study, published in Insectes Sociaux, we explored the impact of these specialized organs on ants by focusing on a species from the less-studied family Riodinidae, Synargis calyce, which interacts with various ant species. In our study area, the most frequent interaction involved the ant species Camponotus crassus.

Caterpillars of Synargis calyce interacting with different ant species. (a) With Camponotus crassus, (b) with Camponotus renggeri, (c) with Wasmannia auropunctata, and (d) with Paratrechina longicornis. ©Amalia V. Ceballos-González.

Caterpillars of this species possess two pairs of tentacular organs. The first pair, known as ATOs (Anterior Tentacle Organs), likely release volatiles that influence ant behavior, although there is insufficient evidence to confirm this. The second pair, known as TNOs (Tentacle Nectary Organs), secrete a nutritive substance (primarily composed by sugars and amino acids) that ants consume. Whether these organs work synergically or if one is more relevant than the other was still unclear for our study species and it is also the case for many other species of the family Riodinidae.

Illustration showing the position of the tentacular organs (TNOs and ATOs)  in a caterpillar of the Riodinidae family. Below, a photograph of Synargis calyce indicates the two pairs of tentacular organs with arrows (Yellow = ATOs, Blue = TNOs). Drawing adapted from DeVries (1991). ©Amalia V. Ceballos-González.

To uncover those aspects, we aimed to explore this pair of tentacular organs by checking ants’ reaction. Our research was conducted at the University of São Paulo, Ribeirão Preto campus. Our first objective was to create an ethogram documenting the behavioral interactions between caterpillars and ants. During these observations, we identified a striking behavior. The ethogram revealed that after the eversion of ATOs, ants exhibited stereotyped “jumping” behavior. This behavior involved ants rapidly lifting their legs and jumping towards the caterpillar’s head.

 Synargis calyce caterpillar interacting with a Camponotus ant.

Next, we conducted experiments in which we experimentally manipulated – by allowing or preventing them to evert – the two types of caterpillar organs (TNOs and ATOs), to determine their role in maintaining ant attendance. Our findings demonstrated that TNOs are more effective in maintaining the attention of attendant ants, likely due to the rewards these organs provide. However, we also found that caterpillars with only functional ATOs received more attention compared to those with neither organ functioning. This indicates that TNOs play a central role in sustaining ant-caterpillar interactions, while ATOs serve a complementary function.

Three caterpillars (possibly third instar) interacting with Camponotus ants.

In conclusion, the interactions between S. calyce caterpillars and attendant ants are primarily driven by the rewards produced by TNOs, with ATOs playing a smaller, supportive role. These findings are consistent with observations in Lycaenidae species, which exhibit similar mutually beneficial relationships with ants. The evolution of these organs may represent a case of convergent adaptation to environmental pressures experienced by caterpillars in both families.

Viruses of eusocial insects: high and underestimated diversity

By Anna Zueva

In this blog, Anna Zueva, researcher at the A.N. Severtsov Institute of Ecology and Evolution (Moscow), reveals the hidden world of viruses in eusocial insects. Read her latest article in Insectes Sociaux here.

Eusocial insects are a kind of human society in miniature. Each part of the community has its own functions and features. The system of social insects’ family works as a coherent and well-coordinated mechanism, and the life of each individual strongly depends on the life of the whole complex.

The functioning of even a fine-tuned system can be disrupted. For example, in the history of mankind there are known episodes when whole civilizations suffered from local epidemics and even pandemics caused by microscopic entities – viruses. But is that the case for social insects?

Our work started from our interest in viruses of invertebrates. Tropics are especially promising for us, as this region is known for its great biodiversity and has lots of still unknown biological species, including of course microorganisms and viruses.

We started our investigation with the research on viruses of termites of Cát Tiên National Park – a part of the Đồng Nai Biosphere Reserve (Fig. 1). We are grateful to the South branch of the Joint Russian-Vietnamese Tropical Research and Technological Center for invaluable help in our studies.

Figure 1. A gap in the tropical monsoon forest of Cát Tiên National Park (Vietnam). ©Anna Zueva

We took samples of termites of three different species feeding on different substrates – lichens and fungi – as we expected that food resource can affect the composition of viruses associated with insects (Fig. 2, 3).

Figure 2. One of the studied species of termites, Hospitalitermes bicolor (Haviland) feeding on lichens. ©Alexei Tiunov
Figure 3. A small termitarium. ©Andrey Zuev

Thought we did not observed any visible symptoms of infection, we detected four new viruses related to viruses previously discovered in termites. We also found the evidence of presence of virus probably belonging to termites’ food substrate (Litov et al. 2022), which partly support our suggestion about the effect of feeding type on the insect virome.

Potentially, eusocial insects can be a model for studying the spread of viruses via social interactions. In the recently published review (Zueva et al 2024), we aimed to actualize the information on the diversity of viruses associated with termites and ants, which are among the most functionally important soil invertebrates. In our review we analyzed 93 articles dedicated to viral findings in both groups of insects. To date, viruses were detected in 54 ant species and in 28 species of termites. We have pointed out 270 viruses and viral genetic variants detected in soil-dwelling social insects, and less than one third of them were associated with termites (Fig. 4). It is obvious that the virome of termites is still mostly undescribed. In addition, both for ants and termites, the information on symptoms or on replication of viruses in their insect hosts remains strongly limited. More studies of the virome of soil-dwelling eusocial organisms with more attention to viral replication and infection symptoms are needed (Zueva et al. 2024).

The most amazing is, that despite the presence of numerous potentially harmful viruses and intense interactions between individuals within a colony, evidences of massive viral epidemics in termites are virtually unknown. We found the information only about one possible virus-caused termite family extinction (Chouvenc et al. 2013). This is especially surprising since termites are important pests of human structures and agriculture, and the search for viruses that infect them has been ongoing for a long time.

Figure 4. Visualization of the presence of viruses in ants and termites (based on Zueva et al. 2024).

There are numerous other unsolved questions on the virome of social insects. How many more viruses of eusocial insects we still don’t know about? Are they able to cause acute dangerous infections or are they just present in the insect tissues and do not manifest themselves until the immune system of host is critically compromised?

We are planning to continue our work in tropics, both on social insects and beyond them. We are sure that this region is a great source of new virological investigations, both on social and solitary invertebrates. By the way, a recent research of our team revealed at least eight new viruses in millipedes collected in the Cat Tien National Park (Litov et al. 2024).

References:

Chouvenc T, Mullins AJ, Efstathion CA, Su NY (2013) Virus-like symptoms in a termite (Isoptera: Kalotermitidae) field colony. Florida Entomologist 96(4):1612–1614. https://doi.org/10.1653/024.096.0450

Litov A.G., Semenyuk I.I., Belova O.A., Polienko A.E., Thinh N.V., Karganova G.G., Tiunov A.V. (2024) Extensive diversity of viruses in millipedes collected in the Dong Nai Biosphere Reserve (Vietnam). Viruses, 16: 1486. https://doi.org/10.3390/v16091486.

Litov AG, Zueva AI, Tiunov AV, Van Thinh N, Belyaeva NV, Karganova GG (2022) Virome of Three Termite Species from Southern Vietnam. Viruses 14(5):860. https://doi.org/10.3390/v14050860

Zueva AI, Zuev AG, Litov AG, Karganova GG, Tiunov AV (2024). Viruses of ants and termites: a review. Insectes Sociaux, 1-12. https://doi.org/10.1007/s00040-024-01008-8

Interview with a social insect scientist: Tom Ratz

Tom Ratz is a researcher at the University of Zurich, studying social interactions in arthropods like Drosophila and beetles. One of his most surprising discoveries came during his PhD while observing burying beetle mothers. Check out his latest work in Insectes Sociaux here!

IS: Who are you, and what do you do?

I am an SNSF Ambizione Fellow based at the Department of Evolutionary biology and Environmental Studies, University of Zurich, Switzerland. My research broadly explores social interactions in arthropods and their role in evolution. My current focus is on agonistic interactions in the highly aggressive species of fruit fly Drosophila prolongata. My group uses a combination of behavioural experiments, quantitative genetic tools, and experimental evolution to test how the competitive environment shapes the evolution of social and non-social traits. 

IS: How did you develop an interest in your research?

My fascination with the insect world began early, around the age of eight, when I started collecting beetles in the backyard of my house. I was captivated by the hidden, bustling world of insects happening all around us, often unnoticed. I wanted to pursue a career in entomology and enrolled in a biology degree. During my studies, I found myself particularly drawn to ethology and behavioural ecology. Applying these fields to insects felt like an exciting way to maintain a connection to entomology while exploring broader scientific questions about behaviour and ecology.

Aggressive encounter between two male Drosophila prolongata. In the first image (left), one male chases the other, leading to an escalation into a fight involving leg fencing (right).

IS: What is your favorite social insect, and why?

Burying beetles are my favourite social insects. I studied them during my PhD and still find their social behaviour incredibly enigmatic–most of which takes place on or inside the decaying carcass of a vertebrate! In a sense, what is a crypt to some is a cradle for burying beetles. Aside from their important ecological role as efficient buriers of small rodent and bird corpses, the complexity of their social interactions within family is, to me, unparalleled in the arthropod world. These behaviours include larvae begging for food and parents regurgitating a “soup” of pre-digested carcass flesh to feed them. Conveniently, burying beetles are mostly undisturbed by experimental conditions, making their behaviour relatively easy to observe and study in the lab.

IS: What is the best moment/discovery in your research so far? What made it so memorable?

A memorable discovery was analysing the data of my first PhD experiment and finding that they absolutely defied our initial predictions. Contrary to expectations, burying beetle mothers didn’t reduce care when experimentally handicapped by a led weight attached to them –instead, they provided more care. At first, this result was puzzling to me, but it became a revelation about the importance of understanding a species’ natural history. It makes sense for a parent to increase investment towards the current brood when prospects for future reproduction are low, which is the case with handicapping, even if the cost of care is higher. This insight highlighted a crucial lesson: while theoretical predictions are valuable, they must be contextualised within the specific biology of the study system.

IS: Do you teach or do outreach/science communication? How do you incorporate your research into these areas?

In both teaching and science communication I try to incorporate my own research as case studies to illustrate key concepts and bring scientific research to life. I find that people are more engaged when they can interact directly with the researchers behind the studies.

Lab stock and experimental populations of the fruit fly Drosophila prolongata

IS: What do you think are some of the important current questions in social insect research, and what is essential for future research?

In my opinion, some critical questions in the field include clarifying the role of social behaviour in shaping population dynamics and evolutionary responses. It is increasingly clear that social interactions within a group can drastically influence the population growth, survival, and how animals respond and adapt to environmental changes. However, what remains less understood is when and to what extend behavioural dynamics taking place among interacting individuals can impact group fitness and drive long-term phenotypic evolution.

IS: Outside of science, what are your favorite activities, hobbies, or sports?

One of my top hobbies is spending time outdoors. I’ve been fortunate to live near beautiful natural landscapes and mountains, which has allowed me to enjoy hiking throughout the year. It’s a great way to clear the mind and recharge. I’m also a regular at the bouldering gym. And of course, entomology remains an important hobby of mine.

IS: What is the last book you read? Would you recommend it? Why or why not?

The book “In search of us: adventures in anthropology” by Lucy Moore, and I highly recommend it. It’s a fascinating account of the origins of anthropology, told through the stories of people who helped found the discipline. It’s rich in field work and historical anecdotes. The author does a nice job of highlighting the complexities of the influential figures in the field–acknowledging both their biases as Westerners and their progressive ideas ahead of their time.

Burying beetles parents feeding their larvae.

IS: How do you keep going when things get tough?

Talking things through with colleagues, friend, or family often helps. While it may not directly solve the problem, verbalising it can normalise the issue and make it feel less dramatic (which it often is). Sometimes, simply going for a walk works wonders–a change of scenery can help put things into perspective.

IS: If you were to go live on an uninhabited island and could only bring three things, what would you bring? Why?

If it can count as one item, I’d bring my partner with me. She’s incredibly resourceful and crafty, and would surely be a great survival companion (as she is in life!). I’d make sure she brings her Swiss army knife, so that’s item number two covered. And obviously a tube to collect beetles as my third item.

Tom, collecting beetles in Greece

IS: Who do you think has had the most considerable influence on your science career?

My PhD supervisor, Per Smiseth, has been a major influence on my scientific career. He’s a mentor with exceptional work ethics. Another key figure was Sylvain Pincebourde, who took me as an intern in his lab when I just a first-year undergraduate. That was my very first  real research experience, and was incredibly formative. I also owe a great deal to many other mentors and colleagues who have had an important role in shaping my interest and career in Science, including Joël Meunier, Pierre-Olivier Montiglio, Niels Dingemanse, Cristina Tuni, Stefan Lüpold, and Wolf Blanckenhorn.

IS: What advice would you give to someone hoping to be a social insect researcher in the future?

A general piece of advice, not just for someone interested in social insects, is to focus on work you’re passionate about and truly enjoy. With genuine interest and motivation, nearly everything seems to become achievable.

IS: Has learning from a mistake ever led you to success?

Yes! And, sadly, the most painful mistakes are often the ones that teach us the most.

IS: What is your favorite place science has taken you?

The tropical forests of Panama are a favourite of mine. I also have a soft spot for Mediterranean ecosystems. Despite their dryness, I’m always surprised by their abundance and diversity of plants and animals.

Ants and their commensals: The intriguing presence of other groups in ant nests

By Mariane Dias-Soares and Cléa S. F. Mariano

In this blog, Mariane Dias-Soares and Cléa Mariano explore the diverse organisms cohabiting ant nests in the Neotropics, from gastropods to myriapods. They explain how do these guests interact with ants, sharing resources and space within the nest environment. Discover more about these intriguing interactions in their latest work for Insectes Sociaux, here.

What attracts these other groups? What are these groups? Are there really gastropods inside ant nests? What are commensals? Do ants benefit from their presence? Why aren’t they expelled? These are some of the most frequent questions when the topic of conversation is our research and our article. Let’s now address each of these questions, the work done so far, and the next steps toward the discoveries that researching an ant nest provides us…

The ant nests provide a protected environment for the workers, the queen, and all of their immatures, as well as storing food and maintaining stable temperature and humidity. When studying these nests, the presence of other groups was observed, which, attracted by these resources, coexist with the ants. These groups may spend part of their life cycle inside the nests or even their entire existence.

Gastropod near the immatures of N. verenae. Photo: Laís Bomfim

Our research aims to identify which groups are associated with different ant species in a Neotropical region. In my master’s studies, I focused on the species Neoponera verenae, an ant from the subfamily Ponerinae that nests in various substrates such as dry cocoa pods, soil, and decomposing logs. In our study, we found a variety of groups, including Myriapoda, Isopoda, Araneae, Lepidoptera, Pseudoscorpiones, Collembola, Acari, Coleoptera, Diptera, Dermaptera, and Gastropoda, among others. This highlights the great diversity of organisms that coexist within these ant nests.

Caterpillar in a N. verenae nest near workers and immatures. Photo: Mariane Dias-Soares]
Researchers during new field collections in the Neotropical Region of Brazil. Photo: Mariane Dias-Soares

Noticing the high number of groups within the ant nests sparked in us the need not only to identify which groups inhabit them but also to understand the interactions that occur in these environments. In our article, we studied the facultative commensalism of gastropods in N. verenae nests, presenting novel records and proposing hypotheses about this type of interaction.

There are different types of interactions between ants and gastropods. In the case of facultative commensalism, the gastropods coexist peacefully with the ants, benefiting from the protection provided by the colony, the available food, and the environmental stability, while also being found outside the nests. For the ants, however, we did not observe any apparent benefit or loss. Further research will delve deeper into these issues.

Gastropods recorded inside N. verenae ant nests. (yellow arrows indicate immatures, and orange arrows indicate snails). Photos from the article by Dias-Soares et al. (2024)

Through various observations and records made in the field and laboratory, we found the presence of several gastropod species inside the ant nests. Among the gastropods found, the family Achatinidae was the most abundant. These gastropods coexisted harmoniously with the workers and the young individuals in the nest (larvae, pupae, and eggs), moving freely without being disturbed by the ants. We also observed that the gastropods produced a foam, which generated a pacifying effect that prevented their expulsion from the nests. This is one of the strategies used by these organisms to inhabit ant nests.

Our study presents novel records of the interaction between ants and gastropods, leading us to explore various unresolved questions. One of these questions is the degree of interaction between immature ants and gastropods, as we found individuals in the chambers that contained the immatures. Additionally, we are investigating the chemical nature of the mucus involved in these interactions and identifying the new species of gastropods found in the nests, in collaboration with Dr. Sthefane D’ávila. Ongoing studies focus on analyzing the chemical strategies used, the morphological adaptations and behaviors exhibited, and the existence of mimicry within these nests. There is still much to be discovered in the vast world that is an ant nest…

Some members of the research team currently conducting collections for the new phase of the Project. from left to right: Fred da Silva, Mariane Dias-Soares and Jossiane Dias
Part of the research group led by Cléa Mariano and Jacques Delabie, focusing on studies of various ant species and other groups present in ant nests

Interview with a social insect scientist: Tomer Czaczkes

Tommy is a researcher at the University of Regensburg, where he leads the ACElab since 2016. He study value perception and decision-making in invertebrates (mostly ants). His latest work in Insectes Sociaux can be found here.

IS: Who are you, and what do you do?

My name is Tomer (Tommy, please) Czaczkes, and I study the behaviour of mostly ants, sometimes bees, and very occasionally other arthropods. My current focus is on comparative psychology – understanding how animals think, learn, and make decisions. I’m trying to apply our hard-earned knowledge of behavioural ecology to controlling invasive ants. I also dabble in collective behaviour.

Tommy Czaczkes thinking about Lasius fuliginosus.

IS: How did you develop an interest in your research?

Ah, well. During my undergraduate the average grades for different modules were available, and I noted that while cell biology and microbiology had pretty low average grades, behaviour and ecology had quite high ones. I know which side of the bread is buttered, and, honestly, I never expected to stay in research. Then, during my undergraduate project, I realized that while the miserable vertebrate ecology people would have to trek for hours through the forest to sight their animal, I, as an experimental behavioural ecologist working on ants, could collect 50 datapoints in half a day, while drinking rum.

IS: What is your favorite social insect, and why?

Oooh, a tough question! I’m torn between two ant species: Lasius niger and Pheidole oxyops. L. niger is perhaps the most common ant in Europe, and as my PhD supervisor Francis Ratnieks always says “it’s the common animals that are most interesting. They’re clearly doing something right.”. L. niger are extremely smart, polite, helpful, and make excellent colleagues. P. oxyops, however, do wonderful cooperative transport – the collective carrying of loads. They have an amazing, explosive recruitment behaviour, and love cheese. They’re also extremely common, but alas, in Brazil and not in Germany, where I’m based.

Pheidole oxyops carrying a 10x10mm square of choose by the corners (published in Insectes Sociaux).

IS: What is the best moment/discovery in your research so far? What made it so memorable?

Seeing ants being visibly disappointed when they received food which was poorer than what they were expecting – poor things! It was clear from the moment I did the first pilot on that project that we would have a clear and strong effect. It was memorable because it was simply so easy to relate to: the disappointed ants would check the food, break away, try again to make sure, and circle around looking for the good stuff they were sure was there before. It was simply so cute and relatable.

IS: Do you teach or do outreach/science communication? How do you incorporate your research into these areas?

I enjoy going into schools and kindergartens, to talk to kids about ants and insects in general. It’s always fun to bring an ant colony or two, and show the “mama ant” and her babies. For the bigger kids, it’s fun to do a pheromone following assay – makes me feel like an ant whisperer, who can use my super science powers to talk to insects.

IS: What do you think are some of the important current questions in social insect research, and what is essential for future research?

This is showing my own biases here, but I think the question of insect sentience and intelligence is a huge question, and social insects are central to the experimental examination of these topics. We’ve had a slew of high impact work reporting all sorts of impressive cognitive abilities, with a big swing from behaviourism to cognition. I expect that very soon the swing will move the other way again, with people starting to push for simpler explanations, or attempting replication studies. Animal behaviour as a subject is overdue a big replication study, the likes of which shook up the worlds of experimental psychology and cancer research (amongst others) recently. I have attempted to replicate some of my own work, with some things replicating wonderfully, and others simply not there next time I looked. And yes, I publish the failed replications too.

Lasius niger worker who is very satisfied with her drop of sucrose solution.

IS: Outside of science, what are your favorite activities, hobbies, or sports?

I really enjoy hiking in the mountains, when I can get out. When not, I’m a big fan of sci-fi books and computer games. My mind is still somewhat blown by my VR set.

IS: What is the last book you read? Would you recommend it? Why or why not?

I’m almost through “Delusions of Gender” by Cordelia Fine. The book speaks against the supposed ‘evidence’ for a simplistic biological basis for gender roles. Would I recommend it? It’s convincing and helpful, but sometimes feels like being bludgeoned with an endless series of (reasonable) criticisms of studies. It’s well researched and useful, but perhaps not the page turner it could have been.

IS: How do you keep going when things get tough?

Give up! No, really. On days where I can’t focus, I simply stop working. If an experiment runs into wall after wall, I’ll drop it. But for things like rejections, failures, etc – I take the long view, and remind myself that this is normal, and this too shall pass. Oh, and moaning. Moaning helps.

IS: If you were to go live on an uninhabited island and could only bring three things, what would you bring? Why?

Assuming my basic survival needs were met? My ebook stuffed with books (for entertainment), a solar charger to charge it, and a Swiss army knife to bootstrap other tools from. I think I’ve played too much Minecraft.

IS: Who do you think has had the most considerable influence on your science career?

Certainly my Doctoral supervisor, Prof. Francis Ratnieks. He has an absolutely excellent eye for interesting biology. Moreover, I admire (and have tried to emulate) his quick, cheap, and cheerful approach to research projects – avoiding the huge, long term, ultra-high tech projects, and preferring short, fun, and simple projects which require only some ants, a few strips of paper, and some drops of sucrose. And a good idea, of course.

In this experiment, Tommy’s team was testing whether ants prefer food they have worked harder for (they do). A good example of their experimental designs. Note the Lego, paper runways, and complete lack of high tech gubbins.

IS: What advice would you give to someone hoping to be a social insect researcher in the future?

Read “The Ants” by Hölldobler and Wilson. Yes, it’s almost 35 years old, but it’s a wonderful primer to most of the major topics in social insect biology. I read it cover to cover to prepare for my PhD, and that knowledge has stood me in good stead since then.

IS: Has learning from a mistake ever led you to success?

Not nearly as much as I would have hoped. I seem doomed to making the same mistakes over and over again. However, at least by now I recognise them with absolute clarity in hindsight.

IS: What is your favorite place science has taken you?

The La Selva biological field station in Costa Rica, where I did my Bachelors project (on leaf cutter ants). Being surrounded by researchers for the first time, in a beautiful jungle, with amazing animals, was life changing. I also met my future wife there, so that was a nice bonus.