Do honeybees make life or death decisions?

A blog post highlighting the article by Justin Schmidt in Insectes Sociaux.

By Justin Schmidt

Throughout much of human history, the ability to think and make decisions was considered a special property of our own species.  Other animals were generally considered to be acting on the basis of instinct.  Decision-making was a part of intelligence, a property most developed and humans, with other animals, including apes, dolphins, whales, and even (my emphasis) some birds like ravens and parrots having some lesser degree of intelligence.  Insects were often described as living robots that simply followed their genetic programming.  We now know that this is a grossly egocentric view and that many species have abilities to learn, as exemplified by honeybee learning in response to nectar rewards.

The ability to make decisions, that is, to tailor the individual’s behavioral response after evaluating input information, appears to be correlated with intelligence.  If so, then is decision-making mainly a property of vertebrates and lacking, or weak, in organisms such as insects?  I suspect that insects and other invertebrates have considerably greater abilities than credited for making optimal decisions based on their evaluation of presently available information.

To test this hypothesis, honeybees were challenged with attacks by a potential predator.  The attacks were a constant that was identical in all situations.  In one situation, the colony nest contained no larvae or pupae and little honey or pollen resources.  The colonies in this situation could also readily abandon their nest and successfully continue by establishing a new nest if necessary.  In the other situation, the colony nest contained a large number of larvae and pupae and substantial honey and pollen resources.  These latter colonies contained only older adult bees and a limited alternative of abandoning the nest and successfully continuing in a new nest.  The little-to-lose colonies were reproductive swarms that had been established in their artificial nest cavity for only three to four days.  The much-to-lose colonies were reproductive swarms that had been established in their artificial nest cavities for 19 to 22 days.


            The predation threat consisted of the “predator” (me) exhaling three times directly into the main nest entrance and then stepping back 5 m and collecting with short-handled insect nets all bees that were attacking my face.  This procedure was repeated as many times as necessary until no more bees were attacking.  Upon finishing collecting all attackers, the colony was collected, frozen, and all attacking workers, non-attacking workers, males, and the queen were hand-counted and weighed.  Colony resources were also removed and weighed.

The results were that the colonies with much-to-lose sent forth 2.9 times as much of their worker force as attackers than the colonies with little-to-lose.  Many other comparative measurements of the two categories of colonies showed the same trend that the colonies with much-to-lose defended more vigorously than the little-to-lose colonies.


            If a worker honeybee attacks and stings a potential predator, she loses her stinger and dies shortly thereafter.  Thus, her decision to attack is not a minor decision; it is a life-or-death decision that has serious consequences for both the individual worker and her colony.  This study revealed that individual workers can evaluate the situation and base their decisions to attack, or not, on the information present at that time.  The study demonstrates that workers can evaluate different situations and decide accordingly how to act in the best interest of the colony.  It leaves many more questions unanswered.  What are the factors the worker uses for making her decision?  Is the decision made by each individual acting alone, or is she acting in response to the actions of other individuals in her colony?  Does the worker evaluate the quantity of brood, honey/nectar, pollen, wax in the combs, or the number of other adult bees, and if so, how?  Is the presence of a queen or queen cells important?  Is the reproductive status, that is, nearness to issuing reproductive swarms, important?  And, to be fair, are the numbers of males present in the colony a factor?  What about pheromones?  As with all studies in science, the results raise more questions than they provide answers.  This is good!  I hope I have cracked the door open a little wider towards an understanding of the abilities of honeybees.

The Insectes Sociaux social media team

Hello social insect fans,

We have had some turnover in our social media editing duo for Insectes Sociaux, so we wanted to take this opportunity to introduce ourselves. Our most recent social media editor, Bernadette Wittwer, had moved on, and we will miss her. I (Madison) am staying on, and it’s my pleasure to welcome Daniela Roemer to the team. I’m excited to work with her to bring you news about new and exciting social insect research! Here’s a little bit about us:

Daniela Roemer

I am a behavioral ecologist interested in self-organized pattern formation in social insects and my study organisms are leaf-cutting ants. After receiving my Ph.D. from the Julius-Maximilians University of Wuerzburg, Germany, I spent two years as a postdoc at the Universidad de la República in Montevideo, Uruguay. Currently, I am a postdoc in the Roces Lab at the University of Wuerzburg, where I study foraging patterns and nest architecture of leafcutters, either in the lab at my university or in the field in Northern Argentina.

Aside from work, I have a passion for travel, books, and my pets (cats, fish, and stick insects). In recent years I have also become interested in science communication and sharing the hard work of my fellow scientists with a broader audience to raise awareness of the importance of scientific research and funding. As the newest member of the social media team, I am very excited to bring you the newest publications and stories from Insectes Sociaux and its blog across the journal’s social media channels.

Madison Sankoviz

I am an entomology Ph.D. candidate in the Purcell Lab at the University of California Riverside. My research focuses on nesting patterns and landscape-level genomic adaptations of Formica ants. I received a B.A. in ecology and evolutionary biology from the University of Colorado Boulder, where I studied the effects of Formica podzolica ant colonies on soil moisture, nitrogen, and plant communities. I am passionate about science communication and am always thinking about ways to bring out the stories behind scientific experiments and findings. The research of other social insect scientists continually inspires me, and I admire these entomologists’ unique enthusiasm for examining social aspects of the natural world. It’s been a blast to highlight publications and help tell the stories behind them as a social media editor for Insectes Sociaux, and I’m excited to continue in this role!

The best part of this role is working with all the contributors to the blog and our interviewees. Thank you to all of you who have participated; we look forward to our communication with many more of you.

If you are interested in blogging or interviewing, do not hesitate to contact us via Twitter (@InsSociaux), Facebook, Instagram, or via email at and

Interview with a social insect scientist: Harmen Hendriksma

Bild Harmen Hendriksma

IS: Who are you, and what do you do?

HH: My name is Harmen Hendriksma. I aim to understand and identify drivers and threats to bee vitality. I currently work on monitoring bees in agricultural landscapes throughout Germany.

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

HH: As a nature boy, I wandered endlessly through the meadows of Friesland. Ants, butterflies, and wild bees strangely enchanted me. Then one magical day, a beekeeper passed my way, and this he said to me: “the greatest thing you’ll ever learn is just to love bees and be loved in return”.

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

HH: My first love was the bumblebee Bombus lapidarius. Yet I was given a hive as a youngster, and ever since, I have been in love with the honey bee, Apis mellifera. Would you oppose having a socialist matricentric society of vegetarians, for a change?

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

HH: We found bees to counter nutritional deficiencies; not only to differentially cover their protein and carbohydrate needs, but also particular needs for essential fatty acids and amino acids. Memorable to me is that many findings come together with crippling self-doubt.

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

HH: I generally follow a biocentric perspective. Whilst teaching Animal Behavior at Iowa State University, the 145 students felt that I sincerely cared for them, and very much care about all other animals too. I never shoehorn bees into lectures. In Germany, Israel, California, and Iowa, I gave many extension talks to beekeepers. I felt those folks all enjoyed hearing stories from a passionate bee scientist who sees beekeepers as facilitators to let bees shine in the spotlight.

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

HH: Many wild bee populations are in decline, and many honey bee colonies dwindle and die. I thusly think that drivers of bee demise need elucidation. Regarding colony structure, e.g., it would be helpful to know the pathogeneses of the many different viral diseases that plague colonies. Future research would benefit from insight into interactions (and maybe synergies) between different stressors, such as nutritional stress and disease.

IS: What research questions generate the biggest debate in social insect research at the moment?

HH: I’m biased toward the nutrition field. There is debate on if generalist bees actively balance deficient colony nutrition, and, how.

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

HH: Spread the word; Sapiens, by Yuval Noah Harari. To me, the book eloquently illustrates a misnomer; Homo sapiens, since pestis would have been a better fit.

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

HH: I have haphazard homey hobbies. I recently inoculated wetted pine wood shavings with mycelium – to grow mushrooms in my kitchen. I hung a birdhouse and a bee hotel outside my home. I glued a pigeon skeleton together that I found in my chimney. I made myself curtains on a sowing machine. And, I stream lots of movies and series – soon also to select new films for an international film festival here in Braunschweig. Once or twice per week, I go out for drinks and bites, targeting unknown cozy places.

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

HH: I tend to just keep pumping in hours and simply keep rocking. I ask for support when needing a boost.

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

HH: I would not bring anything. I’d go out adventuring like a Minecraft survival world. I’d punch down a tree with my fist to make some wooden tools, eat kelp and mobs, etc.

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

HH: During my Ph.D., Stephan Härtel showed me how to practice science. His approach, trust, and patience were most formative.

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

HH: Our social insect researcher society has super kind people that provide astounding support. Take the opportunity and learn from us, work with us, and simply become a part of our field.

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

HH: Dangling above a pool with dolphins in Eilat, Israel, to spot an invasive Apis florea bee colony. I ❤ Israel.

Interview with a social insect scientist: Tae Tanaami Fernandes


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

TTF: My name is Tae Tanaami Fernandes. I finished my Ph.D. in 2018 at the University of Mogi das Cruzes, in the Myrmecology Laboratory of Alto Tietê, under the guidance of Drª Maria Santina de Castro Morini, where we studied ant ecology. The purpose of my thesis was to observe whether abiotic factors of twig structure interfere with ant colonization. We’ve also evaluated the overlap and co-occurrence of other invertebrates in twigs colonized by ants. I am currently developing a postdoctoral project about ants on twigs at the edges of vegetable growing areas in the Alto Tietê region.

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

TTF: During my bachelor’s, I met my current advisor in a course that she taught at the university on urban ants. From that point, I became interested in these insects and started searching for an internship in myrmecology. When I started in the Myrmecology Laboratory of Alto Tietê, there was already an ongoing project that included the study of ants on twigs. Since then, I’ve never wanted to change topics.

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

TTF: I am passionate about the Thaumatomyrmex genus. Although I have never met it in the field, its biology fascinates me. The mandibles are like pitchforks; each mandible is composed of three long tines joined at the base. They are specialized predators of millipedes and are covered with detachable barbed setae that entangle potential predators. The Thaumatomyrmex workers use their long, specialized mandibles to capture millipedes and subsequently strip them of their setae.

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

TTF: I’ve had many memorable moments during the project. One of them was when I did an academic period in Mexico, where I managed to get the real dimension of my work. Another moment was when I started my master’s, and many people were working on the same project with the same energy to work and understand the process of colonization of ants on twigs.

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

TTF: Recently, I began teaching students from 14 to 18 years old in high school. Whenever I can, I talk about what it’s like to be a scientist, what we do in the laboratory and the field, and I talk about the importance of science, especially the importance of insects, which are seen as undesirable animals. I always explain it to my students so that they can spread it to their community.

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

TTF: The identification of ant morphospecies. There are many more species of insects than are catalogued in the literature, and we still don’t know the biology of these species. It would be fascinating to have more people in that area.

IS: What research questions generate the biggest debate in social insect research at the moment?

TTF: Global environmental issues are the main trend. Green areas are continuously shrinking, and this directly interferes with all organisms, especially social insects. The indiscriminate use of pesticides is also a matter of considerable debate since, for many farmers, it is necessary. But with the correct knowledge, the application of pesticides could be reduced.

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

TTF: The book I am reading is Diversity of Life by Edward Wilson. I highly recommend it firstly because the author is a very renowned myrmecologist. Secondly, the text is captivating and rich in detail. In this book, he talks about field experiences in the Amazon rainforest, focusing on its environmental degradation.

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

TTF: I enjoy running, reading non-academic books, and watching TV series. In my hometown, I like watching basketball games with my mother and friends.

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

TTF: I take a few hours off to do other things that are totally different from what I was doing. Then I come back and focus on the problem, and if necessary, I discuss it with my colleagues. Everything works out in the end! This is my slogan.

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

TTF: I would take a bottle to fill with water, a raincoat (which can keep me warm in the cold too), and a knife, which can always be useful.

WhatsApp Image 2020-02-04 at 18.11.29

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

TTF: Certainly my advisor. She was always by my side with a lot of patience and persistence. She has always offered opportunities, suggestions for different courses, and lectures, which were essential for my career and for forming the professional I am today.

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

TTF: Don’t give up at the first difficulty; science needs you! Be curious, questioning, and read a lot. Always ask your supervisors and lab colleagues questions; it may surprise you how helpful these people can be.

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

TTF: A lot of places! The first place was Mexico, where I stayed for four months at the Instituto de Ecología (INECOL), working with Dr. Wesley Dáttilo. I’ve met wonderful people who helped me a lot, especially his students, as well as beautiful landscapes that are only found there. I also got to know Argentina and many Brazilian cities during courses and congresses.

Interview with a social insect scientist: Julia Eloff

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

JE: I am Julia Eloff, and I currently live in Wellington, New Zealand. I have most recently studied the population genetic structure of the invasive German wasp in South Africa. I am very passionate about the natural world and like to take part in other studies involving insects. I love fieldwork and collecting insects, as well as the lab work and molecular sides of studies.


Nature walking in Whangerei, New Zealand. At Whangerei falls.

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

JE: I’ve always had a passion for social insects, in particular, ants. When I was younger, I always believed that they were most like humans. They can solve complex problems, are highly plastic, have a distinct hierarchy, and even build intricate and complex structures. As I got older and continued to study biology, one of my undergraduate courses required a self-directed project in which, not surprisingly, I did a project on ants and their cognitive abilities. I then came into contact with Antoine, the ‘ant guy’, who was at the time doing his Ph.D. He introduced me to his supervisor Prof. Phil Lester, who asked me whether I would be interested in working on wasps. I started to do a lot of background research to find out a bit more about them and even drawing them to learn how to identify them a bit better. One thing led to another, and here I am, researching wasps.

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

JE: I think I may have hinted at earlier, but definitely ants! In particular, Solenopsis invicta or the fire ant. I have always been intrigued by their cognitive ability as a group in which they can problem solve as a collective. This is seen in floods, where they group together, creating makeshift rafts that float on floodwaters and house all members of the colony, including the workers, queens, larvae, and eggs. Although many are scared or afraid of fire ants, I tend to appreciate their collective problem-solving abilities.

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

JE: The best moment of my research would have to be the moment my lab group and I were travelling back from a conference in the South Island, New Zealand. During this time, we were stopping regularly, hunting for wasp samples that we could catch and use for research. This collecting involved lots of running around with pottles and nets, and in some cases jumping into bushes. The people who saw us awkwardly catching wasps in random areas and giving us confused and awkward glances – I can only imagine what they were thinking at the time. The amusement from the situation and confused bystanders has definitely made its way into my memory books.

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

JE: I have been a teaching assistant for several undergraduate courses at Victoria University of Wellington. The two main courses I have taught have been genetics and animal diversity, which are both excellent platforms for sharing what I do. I have also been involved in many outreach events, including judging school science competitions or university open days. I like to encourage young minds to find a passion for science and hopefully find a field in which they are truly passionate about. I won’t lie though, I may have a slight bit of favouritism when people show a passion toward the entomological sciences.

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

JE: I think cognition in insects has been an ongoing area of importance in social insect research. There have been multiple studies looking at the brain of social insects (social brain hypothesis) and others that look at visual cognition as well as other forms of learning behaviour. In particular, in invasive social insects, increased cognitive plasticity could lead them to learn in a new environment quicker and consequently adapt to their new surroundings. Therefore, I think more research in the cognition of invasive social insects and their ability to problem solve as a collective should be undertaken. This has always been an area of social insect research that I have been quite passionate about!

IS: What research questions generate the biggest debate in social insect research at the moment?

JE: I definitely think one of the biggest questions generating debate is whether we should make use of CRISPR-Cas9 gene drives in the management of pest insects. Not only is it a controversial topic where the release of genetically modified organisms concerns the public, but there are some ecological concerns as well. One of the more commonly bought up ecological risks would be the question as to whether we would be able to contain them to prevent their spread outside of the targeted areas. Another question would be whether genetic mutations could result in a rapid removal of the gene drive. Although gene drives can be highly useful as pest management tools in some situations, they may not be as effective in others. With the correct background research and careful consideration of hazards involved, the implementation of gene drives may be a useful tool in the removal of invasive social insects.

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

JE: The last book I read was ‘All Creatures, Great and Small’ by James Herriot. I absolutely loved the book and reading about all his funny tales as a veterinarian and the heart-warming moments. It’s definitely a book I would recommend, especially to all my fellow animal lovers. It’s a book based on true-life stories filled with humour and compassion and multiple animal tales.

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

JE: I’ve always been a nature walk enthusiast. I love to experience nature and to look at the plants and creatures. I also love to draw and paint. From portraits of people to drawing my favourite insect creatures, it’s a passion I’ve had since childhood. More recently, I’ve been trying to get in touch with my sporty side and taking part in some martial arts classes.

Some of Julia’s insect sketches and drawings. @artyjewlz

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

JE: One thing that definitely helps when things get tough is taking advantage of my creative outlet. I like to pick up a pencil and put my emotions on paper. Sometimes just remembering those little moments that made me smile in the day helps too.

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

JE: First, my sketching gear to get me through those tough moments, and document all the nature I see. Secondly, some seedlings to garden for entertainment and maybe food. Lastly, my guitar, music definitely can calm the soul.

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

JE: I mentioned Antoine Felden before, but he has definitely been there for me. Without him, I would not have been where I am now. He was there for me during the roughest of times and let me feed his ants. He excited my science career with his passion and helped me stay in social insect research. Secondly, Mariana Bulgarella, she taught me everything I know, from lab work to analyses. She is one of the most amazing teachers I have ever had who kept my passion for science alive.

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

JE: Keep talking to people about your passions; you never know when you may speak with the right person (like ‘Antoine the ant guy’), who will introduce you to the right people to start your science career.

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

JE: Recently, I took part in a moth survey at Zealandia Ecosanctuary, New Zealand. This survey took place at night. Between the combination of the night sky and the sanctuary, there were thousands of glow worms surrounding us. It made me feel like I was in a scene from Avatar. Not only that, but seeing kiwi in the wild, as well native gecko species and tuatara. It was definitely the right place at the right time.

Insectes Sociaux Best Paper 2019

It is a pleasure to announce that the winner of the 2019 Best Paper Award is “Vitellogenin and vitellogenin-like gene expression patterns in relation to caste and task in the ant Formica fusca” by Claire Morandin, Anna Hietala and Heikki Helanterä at the University of Helsinki, Finland. Here is the journey of how this project came to be, in Claire’s words.

When I started my Ph.D. in early 2011, Vitellogenin (Vg) was the “hit” gene that everyone was interested in. Vg is one of the most studied genes involved in the division of labour across social insects. My first Ph.D. project was to look at Vg expression patterns across multiple Formica ant species, but upon constructing the first Formica transcriptome (Dhaygude et al., 2017), we surprisingly came across not only one Vg gene, but four. To understand these new Vg-like-genes (as we named them), we performed gene expression analyses, protein modelling, and evolutionary analyses, and found that these three new homologues were the result of ancient duplications. They partly differ in their conserved protein domains and have undergone rapid evolution after duplications. Furthermore, their expression patterns and thus their likely roles in social regulation were not consistent across the seven Formica species we looked at, providing important new insights into the complexity of insect social behaviour and gene expression variation amongst even closely related species.

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Understandably, we got really excited about these Vg-like genes and knew we needed to look deeper into their roles. In species with multiple conventional Vg (such as Solenopsis invicta), the multiple copies show sub-caste and task-related expression patterns, potentially linked to the loss of reproductive constraints and evolution of new functions for the duplicated copies (Wurm et al., 2011). We then questioned if similar caste- and/or task-related expression differences would likewise have emerged during the ancient duplication events between/among the conventional Vg and the three Vg-like genes. Hence, to begin to comprehend the role of Vg homologs and their potential involvement in division of labour, we designed a study that would allow us to investigate the relationships of expression patterns not only between castes (queens vs. workers), but also between different tasks (nurses vs. foragers), between colonies with and without a queen, and between several points in time.

At the same time, Anna started her master thesis with us. It was a challenging and stimulating project that involved a lot of field and lab work. Once the snow melted, we went out around Tvärminne Zoological Station in the south of Finland and collected dozens of F. fusca colonies. The time frame was tight, as we needed to find them after they came out of hibernation, but before they got too active, in order to catch the entire colony (or most of it), the queens included. As we needed colonies with several queens, we could not afford to miss queens while collecting. Back in the lab, the colonies were carefully sorted to find and count all of the queens (that’s a tedious job that involves going through a bucketful or two of nest material and soil for each nest, and hundreds or even a few thousand workers that do not appreciate your efforts). Experimental nests were established in plastic trays with a feeding platform so we could differentiate nurses and foragers. The experiment lasted for 20 days; every five days we collected nurses, queens, and foragers from each nest for gene expression analyses and checked whether queens or workers had been laying eggs. After that, we brought back the samples to the University of Helsinki for gene expression analysis. Anna extracted RNA from more than 500 individuals and performed qPCR analysis (needless to say it was a challenging task for someone who has never done any lab work after one or two basic courses, but she managed brilliantly). Apart from the wet lab work, Anna also dissected the ovaries of a few hundred workers to see whether queenlessness incites ovary development in workers.

Our results showed that each of these genes had a unique caste-specific expression pattern in F. fusca. Expectedly, we found a significant caste and worker task-related increase for the conventional Vg. We found that task (nurses vs. foragers) was the only factor that explained expression variation among workers in any of the studied genes and that removing the queens did affect expression, despite the fact that the proportion of fertile nurses increased significantly. As in previous studies (Kohlmeier et al., 2018; Salmela et al., 2016), our results are consistent with the idea that Vg-like-A may be involved in worker behaviour, and Vg-like-B in stress resistance in ants, while Vg-like-C displayed a consistent forager-biased expression pattern (just like in other social insects (Harrison et al., 2015), suggesting that Vg-like-C might have sub functionalized to a completely different role.

With this project, we aimed to get a clearer picture of the roles of these newly found Vg-like genes for caste differentiation. We still do not know their precise roles, and for example, tissue-specific expression analyses would be an important next step, but at least now we know that their expression patterns are consistent with roles in the division of labour separate from the conventional Vg. We hope this study will spark further interests in these really interesting Vg homologues, and hopefully, at some point we will find out exactly what these genes are doing at a molecular level.

As many projects do, this one took its time as well, and there were challenges along the way. For example, extra effort was needed from Anna who had to write the thesis in English and not in her native Finnish as Claire as her supervisor needed to understand it as well. It was a long journey from the beginning to Anna’s MSc thesis, and then publication in Insectes Sociaux. After all the work for this study, the award means a lot to all of us!


Dhaygude, K., Trontti, K., Paviala, J., Morandin, C., Wheat, C., Sundström, L., & Helanterä, H. (2017). Transcriptome sequencing reveals high isoform diversity in the ant Formica exsecta. PeerJ, 2017(11), 1–31.

Harrison, M. C., Hammond, R. L., & Mallon, E. B. (2015). Reproductive workers show queenlike gene expression in an intermediately eusocial insect, the buff-tailed bumble bee Bombus terrestris. Molecular Ecology, 24(24), 3043–3063.

Kohlmeier, P., Feldmeyer, B., & Foitzik, S. (2018). Vitellogenin-like A–associated shifts in social cue responsiveness regulate behavioral task specialization in an ant. PLoS Biology, 16(6), 1–26.

Salmela, H., Stark, T., Stucki, D., Fuchs, S., Freitak, D., Dey, A., … Sundström, L. (2016). Ancient duplications have led to functional divergence of vitellogenin-like genes potentially involved in inflammation and oxidative stress in honey bees. Genome Biology and Evolution, 8(3), 495–506.

Wurm, Y., Wang, J., Riba-Grognuz, O., Corona, M., Nygaard, S., Hunt, B. G., … Keller, L. (2011). The genome of the fire ant Solenopsis invicta. Proceedings of the National Academy of Sciences of the United States of America, 108(14), 5679–5684.

Interview with a social insect scientist: Robert J. Warren II


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

RW: My interest is in the impact of global change (species invasion, climate change, and habitat fragmentation) on species interactions, which quite often steers me toward social insects.

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

RW: Really, at its core, my interest in research stems from walks in the woods. My favorite research projects tackle natural history observations with ecological theory.

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

RW: The Aphaenogaster rudis complex (woodland ants in eastern U.S. deciduous forests) certainly are my favorite social insects because I spent much of my life hiking and playing in woodlands, and I never knew that a single species was so dominant (both in abundance and impact). Now, I see them everywhere.

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

RW: I spent my doctoral research trying to explain the contrasting distribution of two myrmecochorous plants in the Southeastern U.S. based on the niche requirements of the plants. One day, when loading equipment into the back of my truck, I thought, ‘what if it is the ants?’ My research had solely focused on plants up until that time, and incorporating ants opened up a whole new world for me (including new insights into global change ecology).

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

RW: I teach ecology and biostatistics in a biology department (SUNY Buffalo State). I share my research and field experience with students – most of whom want to go into the medical field and do not think ecology is relevant to their careers – to connect with the students and help them see a different aspect of biology.

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

RW: I am fascinated with how often social insects get cheated. We know of many great benefits that come with sociality/eusociality, but one of the main benefits seems to be that a colony can amortize the cost of being cheated across many individuals and hence tolerate it more easily than a solitary organism.

IS: What research questions generate the biggest debate in social insect research at the moment?

RW: In my little area, ant-mediated seed dispersal has long been described as a mutualism despite little evidence supporting palpable benefits for the ants. I have received surprisingly strong and emotional negative responses, particularly from plant-oriented folks, when questioning it as a mutualism.

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

RW: Last Train to Memphis, by Peter Guralnick. I would recommend it. Guralnick does an amazing job detailing the rise of Elvis Presley, including showing that Presley had a deep understanding and passionate love for black music, and you realize that a lot of the resistance to Presley in his early years was racist resistance to his integrating black musical styles into what became rock and roll.

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

RW: I love hiking and all that stuff, and I spend a lot of time doing home improvement on our 19th Century Victorian home. I also love watching Indiana University basketball and University of Georgia football.

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

RW: I have made a few great decisions in my life, and my wife of 27 years is one. She is my foundation.

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

RW: These days that question is a little open given that we can bring a Kindle and mp3 player with thousands of options, but I will try and answer in the traditional sense. I would bring Alice in Chains “Grind” because I can listen to it again and again, Hermann Hesse “Siddhartha” because I seem to find something new every time I read it, and a large supply of Tabasco sauce because I like it on my seafood.

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

RW: Retired University of Georgia Professor Ron Pulliam. I entered his lab as a former newspaper journalist and left a scientist.

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

RW: Work with people that you like and find projects for which you have passion because it is a helluva lot of work to do for low pay and little reinforcement.

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

RW: As a literal ‘place,’ I love doing work in the Southern Appalachian Mountains where, even though there are thorns, wasps, and steep slopes, I can work in shorts and a t-shirt and get lost in the green. As a figurative ‘place,’ I love working with students and scientists, which is endlessly fascinating and rewarding.

Interview with a social insect scientist: Tim DeLory


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

TD: My name is Tim DeLory. I am currently a Ph.D. student at Utah State University in Dr. Karen Kapheim’s lab, where we study the evolution of sociality in bees. The objective of my dissertation is to study genome evolution in bees across a range of bee social life histories using a bioinformatic approach. I am particularly interested in the potential impact that these complex social phenotypes can have on the evolution of genomes.

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

TD: I first learned about social insects in my undergraduate entomology class. My professor, Kevin Alexander, explained the haplodiploid mating system of hymenopterans to us and showed us some of the different species of bees with variable sociality. This piqued my interest in social insects. The following summer, I had the opportunity to attend a Research Experience for Undergraduates (REU) at UNC Greensboro under the guidance of Olav Rueppell, where we studied recombination rates in honeybees. It was after this REU that I knew I wanted to make studying social insects my career path and use bioinformatics approaches to do this.

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

TD: I would have to say Meliponini. Maybe it’s cheating to say a whole tribe is my favorite, but it’s their diversity of behavior and life histories that fascinates me. They have a wide variety of castes and nesting strategies. I have also always loved reading about interspecific interactions between eusocial species, and the ​Lestrimelitta limaois a fascinating example of this. They have a “robber” caste instead of a foraging caste which robs the reserves of other stingless beehives. The pollen-collecting morphology of ​Lestrimelitta is reduced and vestigial. It is interesting to me that the forager caste is not inextricably linked to pollen-collecting in bees. On a personal note, I think they are beautiful (especially ​Tetragonisca angustula) and have an incredible variety of kinds of honey.

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

TD: I would say the most memorable moment in my research was when I was able to see the results of a simulation that I made when finishing a fire ecology project I had been working on before starting my Ph.D. It relied on some field data we had collected as well as GIS data that other students had gathered. The results were interesting, but the moment was memorable because I liked the collaborative effort that went into it. I was able to really integrate my math background, as well. It gave me a strong sense of affirmation that research in the life sciences was the right career path for me.

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

TD: I am a teaching assistant for some introductory biology labs at my university. I try to discuss my research whenever appropriate in these classes to demonstrate that people do apply the skills they are learning. When teaching, I also try to expose my students to other career opportunities beyond academia or medical school, such as land management agencies or non-profit work. I have some friends in these areas that I can refer them to if they have questions. I think this helps students feel less siloed by a biology degree, which hopefully translates to retention. As far as conversations with the community at large, I really consider it a victory when I can explain that there is a lot of research beyond honeybee health going on in entomology, and why basic scientific research, in general, is valuable.

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

TD: I am relatively new to research in social insects and research in general. But I would say some of the recurrent issues I have noticed have to do with the rate of advancement of sequenced genomes, while the characterization of insect life histories has not advanced nearly as rapidly in recent years. In 06’, the honeybee genome was assembled, and there was a lot of information about its social life history to contextualize that genome. We are now in a position where we can afford to sequence bees whose more basic qualities are still a mystery. It will be interesting to see how new assessments of bee life histories can help to direct sequencing efforts. Or perhaps as sequencing becomes more and more affordable, we will sequence results to direct which bees we select to do behavioral studies in the field.

IS: What research questions generate the biggest debate in social insect research at the moment?

TD: I know that the value of kin selection theory as a framework for understanding social evolution has been repeatedly brought into question over the years, and has sparked many debates. Although I am not sure if this issue is as contentious today. One recent question that seems to generate a lot of discussion and differing opinions is the role of using insects with semi-social or facultatively eusocial lifestyles to understand eusocial evolution as a whole. I think the research community has not yet reached a consensus on what we can reasonably infer about the evolution of extant eusocial lineages using less socially complex insects as ancestral proxies.

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

TD: The last book that I read was called The Dragon Reborn. It was a fun read for me. I would recommend it if you really like reading Tolkein-like fantasy books with expansive world building and have read the first two books of the Wheel of Time series. I would not recommend it if you are interested in a book with deep or dynamic characters. The plot moves pretty slowly as well because it is part of a much larger series.

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

TD: I like different outdoor activities such as hiking, rock-climbing, and skiing. I also enjoy jiu-jitsu.

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

TD: Mostly, I just think about what got me interested in social insects in the first place. The idea of these various insect societies interacting and inhabiting the planet and acting out these amazing and intricate narratives reminds me of a science fiction novel. It’s endlessly fascinating. It’s easier to push through when I can remind myself that my job is to research something that is fundamentally cool.

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

TD: I would bring seeds. I used to have a large vegetable plot when I was younger, and I loved tending to it. So, I think gardening would be fun, and an excellent way to have a steady source of food. I would bring my collection of books. I think I could read Frank Herbert’s Dune once a year and not get sick of it. Lastly, I would bring a volleyball, because it is always good to have a friend.

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

TD: I had a fantastic group of professors during my undergrad at Western Colorado University (WCU) who all encouraged me to do research and taught me valuable skills to do my research. My research advisor at WCU, Dr. Jonathan Coop, really comes to mind. He gave me a lot of creative control over designing a simulation for a project of his that I was working on. I felt like a colleague that was making a real contribution. Beyond that, he has always respected my personal decisions, including leaving school to work when I needed a break.

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

TD: I would say that reading a healthy amount of papers is helpful for this in the beginning. The literature will give you a sense of what intrigues you about social insects. You can check out the lab sites of the authors of the papers that you especially liked, and contact those authors. That is how I met my Ph.D. advisor. It has been my experience that everyone I have contacted in the social insect community is affable. They will recommend other labs, too, if they think those labs could be a good fit for you.

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

TD: I had the opportunity to go to an old wildfire site from the 2002 Rodeo-Chedeski burn in Arizona to collect data for a fire ecology research project. I was on top of a mesa looking around, and it really made me realize how expansive the burn was. That vista definitely made a strong emotional impression. I could also see how the landscape had such a dynamic recovery response from one drainage to the next during the time I spent there.

Colonies on the march move maternal alleles across tropical landscapes

A blog post highlighting the article by Soare et al. in Insectes Sociaux.

By Sean O’Donnell

Army ants are strange beasts. New World army ants (Ecitoninae) share a suite of unusual characteristics, including mass foraging raid behavior, that distinguish them among ants and indeed among all social insects- the army ant syndrome. Their strangeness may breed success: recent evolutionary analyses suggest the army ant syndrome is ancient (perhaps 80 million years old), and army ants are among the most important ecological players in the tropical forests where they thrive.

One component of the army ant syndrome relates to their mode of reproduction. Like some other social insects, notably honeybees (Apis) and some paper wasps (Epiponini), army ant colonies reproduce by swarming. Swarming involves groups of workers and one or more reproductives (queens, in Hymenoptera) moving away from the natal colony to establish a new daughter colony. Army ants are unusual because their impressively large queens are wingless. During periodic bouts of reproduction that occur every few years, an army ant colony produces a single surviving daughter queen who mates with a number of visiting winged males that arrive from distant colonies. After mating, the young queen inherits about half of the worker force and walks away from her natal colony to start a new society.

Differences in mobility between queens and males occur in some ant species and are usually associated with significant sex differences in dispersal distance. In these species of dependent-founding ants, homebody queens settle in or near their natal nests; males fly longer distances to seek mates. This sex difference can be important for population structure: the mobile longer-flying males move genes (alleles) greater distances. It was long assumed that wingless army ant queens were relatively mobility-challenged, and that queens were therefore less important than winged army ant males for maintaining gene flow in army ant populations.

However, another unusual key feature of the army ant syndrome is colony nomadism. Army ants do not dig and occupy permanent nests. Rather, they regularly move or emigrate among a series of temporary shelters where they bivouac, assembling a temporary nest from the interlinked bodies of the workers. In the well-studied species Eciton burchellii, colonies are on a five-week cycle, of which two weeks are spent emigrating among a series of nesting sites. Single emigrations can traverse 100 m linear distance. Successive emigrations tend toward directionality: a right turn on one night’s emigration path is likely to be followed by a left turn the following night. Do such colony movements, summed over the three years between reproductive bouts, contribute to maternal (female) gene flow?

Video of an an Eciton burchellii colony starting to emigrate. Source: S. O’Donnell.

We hypothesized that colony emigrations would contribute to maternal gene flow and reduce or eliminate sex biases in gene dispersal. We tested for sex biases in gene flow by measuring the genetic relatedness among males and females (queens) in a population of the army ant Eciton burchellii parvispinum in the mountains of Costa Rica. We collected samples of workers from a total of 40 colonies in a roughly 10 km X 10 km area. We sampled colonies in the same geographic area in 2006 (25 colonies), and again roughly three years later in 2009 (15 colonies). Three years represents the typical generation time (time between reproductive bouts) for E. burchellii colonies.

We genotyped workers by using PCR primers to amplify seven highly variable microsatellite DNA regions. We then reconstructed the maternal (queen) genotype for each colony as well as the genotypes of the males that had fathered the workers we sampled. We asked whether relatedness among the queens and their mates decreased with distance among the colonies with years, as well as testing for spatial genetic structure between the 2006 and 2009 samples.

We found no significant difference in spatial genetic structure between the sexes, either within or between the 2006 and 2009 samples. In fact, there was some evidence against genetic philopatry for queens: the queens sampled in 2009 were significantly unrelated to queens sampled in 2006 that were collected nearby (within 0.5 km). These patterns suggest maternal dispersal via emigrations contributed to gene flow in these army ants, reducing or eliminating male biases in dispersal. Because colony emigrations summed over the lifetime of army ant queens queen may contribute to gene flow across the landscape. The ecological consequences mean that habitat connectivity is essential to permit colony emigrations and support genetic diversity in populations of this keystone species.


Interview with a social insect scientist: Antoine Felden

IS: Who are you and what do you do?

AF: Hi, I’m Antoine Felden, and I live in Wellington, New Zealand. Currently, I study the Argentine ant immune system in response to pathogens, mostly using transcriptomics and functional genetics (RNA interference). I also have other ongoing projects, too, like the study that just published in Insectes Sociaux about how a mite might be associated with wasp pathogens. I’m getting more and more experience in computational biology, and that is what drives me. I love fieldwork (and bench work to some extent, too) but I thrive working with computers and digging deep into datasets.


Probably talking about Toulouse and bees at the Biology & Genomics of Social Insects conference © Cold Spring Harbor

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

AF: I have had a passion for ants as far back as I remember. That is what got me into biology in the first place. I worked on a lot of different topics over the years, from behaviour to pathogens and immune response, locomotion mechanics, learning, and nutrition, all using ants as study models, and I’ve always been fascinated by what I was doing. So I suppose the short answer to that question is: ants.

IS: What is your favorite social insect and why?

AF: I suppose I don’t need to say it twice! I think ants tap into the fascination I have for miniature worlds. I was a pretty weird kid staging up small sceneries and playing with little things until an unusually old age, and ants perfectly fit this kind of aesthetic. Then, growing up and studying, I learned about the complexity of their biology, and a more scientific fascination about them emerged. I am amazed by all the different characteristics that have evolved in different species, from kidnapper ants to fungus-growing ants, army ants, aphid-tending ants, unicolonial ants, and so one. There is so much diversity in the ant world that one can never stop being amazed.

More anecdotally, there was a colony of Formica selysi living in the sandy backyard of the family holiday house on the Atlantic coast of France. After looking forward to seeing them again every year in summer, I have to say they have a special place in my heart; they looked stunning with their iridescent silver cuticle, running in the sun.

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

AF: One highlight discovery was when I was to study appetitive conditioning in Argentine ants to decipher nestmate recognition mechanisms, back in Berkeley with Ellen van Wilgenburg and Neil Tsutsui. I was using something similar to the bee “proboscis extension reflex” in ants to figure out if there were able to learn some olfactory cues better than others. Watching them under the stereomicroscope sticking their tongue out as they remembered that a given smell was followed by a sweet treat was a successful experiment like few happen. That was a while ago; now, my academic path is taking me to fuzzier transcriptomic studies where discoveries often bring more questions than answers.

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

AF: Questions that revolve around caste differentiation and its molecular basis have the potential to have significant implications for our understanding of developmental biology as well as the evolution of sociality. For our society’s benefit, there might also be a lot to learn in studying aging in ants where queens can have such an extended lifespan compared to the average worker. What is essential for further research in most fields is to develop more models for functional genetics, like in the CRISPR-ant studies published in the last couple of years. It will be paramount to investigate these processes better, and I’m really excited about this!

I also think that unicoloniality that some ant species display is an odd phenomenon and that its ultimate mechanisms are still poorly understood. What caused the formation of these large supercolonies of ants? What are the behavioural consequences of such an upscale colony size? Is unicolonial social structure viable over evolutionary time? I think these questions still need more attention.

IS: What research questions generate the biggest debate in social insect research at the moment?

AF: I see that there is still a lot of room for debate in themes revolving around the evolution of sociality. The recent papers by Boomsma & Gawne (2018) on superorganismality and Linksvayer & Johnson (2019) about misconceptions on social complexity were pretty thought-provoking and highlight how much there is still to understand.

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

AF: First, the series (especially the first volume) called Les fourmis by Bernard Werber is a must-read for science fiction enthusiasts and naturalists alike. It is a very well-read novel, and there are still some accurate science facts in there (if one lets scientific rigour loose a little). I also love most of Virginie Despentes’ work. She writes about lives going on destructive train wrecks. Her characters are often objectively dark and despicable people, but following their struggles from their own perspective makes you uncomfortably sympathetic with their twisted journeys. I definitely recommend checking them out; there are English translations available!

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

AF: Since I moved to New Zealand, I took up surfing more seriously. I’m lucky to live in Wellington; there is a nice beach easily-accessible by bus (I’m one of these university people who don’t have a driving licence… I know I’m not the only one out there!). It’s often a mix of chance and negotiation skills to hop on the bus with a surfboard, but it’s always worth a try. And thanks to the Argentine ants that like it hot and sunny, fieldwork in the warmer northern tip of New Zealand is conveniently located around beautiful surf beaches. Beyond that, I really like hanging out with my friends.


Collecting Argentine ants living inside beehives in the Far North of New Zealand, with Jana Dobelmann © Phil Lester

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

AF: Highs come with lows, and I try to remember that problems eventually drift away just like clouds in the sky. When issues arise, I try to not lose sight of their resolutions that come with trying a bit harder.

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

AF: I’d bring a sound system, a laptop, and solar panels – provided that supplies are already on the island. Then I’m all set to take science notes, write a book, then celebrate the outcomes with a dance.

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

AF: My first internship mentor – Tim Brütsch, in Lausanne, who was doing his Ph.D. at the time. He taught me the necessary dedication for experimental work, how the academic world works, and sparked my interest in an evolutionary perspective on biology. I’d also like to mention my dear colleague Raphael Aguillon (not a social insect researcher – but no one is perfect) who asks the big questions and also showed me what passion for science looks like.

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

AF: Stay around good people and trust that with time, luck always strikes. I have both of those at Victoria University working with Phil Lester: a pleasant working environment with good funding to answer our questions, as well as having been at the right place and the right time.

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

AF: I went to beautiful Tokelau to deal with a Yellow Crazy ant invasion as part of Monica Gruber’s Pacific Biosecurity program. Tokelau is a tiny atoll nation north of Samoa. It’s pretty hard to get to, as it is only accessible by boat every fortnight from Apia, and it is one of the most incredible places I’ve seen, full of lovely people. To this day, I can still reminisce how constantly stoked I was to be there.


Telling Tokelau kids about invasive ants © Evan Brenton-Rule