The paraphyly controversy

Highlighting the articles written by Ward et al. and  Seifert et al. in Insectes Sociaux

Written by Insectes Sociaux Editor in Chief, Michael Breed

Should we change a commonly used and widely understood name when bringing classification into alignment with phylogeny? This is not a new question; it has plagued generations of systematists (e.g., Michener 1964). The Linnean system of classifying and naming organisms is a foundational tool in biological sciences. Yet not all scientists agree about how and when information about phylogeny should be embedded into classification. A particularly sticky point comes when well-accepted names for organisms conflict with newly understood phylogenetic information. The second of a pair of commentaries dealing with phylogeny, classification and scientific names appears in this month’s issue of Insectes Sociaux (Ward et al. 2016; Seifert et al. 2016).

Particularly vexsome in the social insect world have been classifications involving social parasites. Are there clear synapomorphies-traits that distinguish the social parasite from its hosts? Alternatively, is each socially parasitic species within a clade an evolutionary offshoot from shared ancestors with its host species? Or is it a distinct clade within a clade? Are parasitic characteristics, which often involve loss of foraging morphology, arrived at by convergence? The answers to these questions often affect a species’ name, in particular its placement in a genus.

The two commentaries address the question of whether, when new data reveals that genera are paraphyletic, should their naming reflect the current phylogeny. In ants, there are several examples of groups of parasitic species for which new data has caused placement within a host genus, when previously the parasites had generic status. The renaming of species has been controversial and lies at the core of the disagreement between the two groups of authors.

The much-studied relationship between bumblebees, Bombus, and their social parasites, Psithyrus, exemplifies the fascinating difficulties posed by social parasites. Psithyrus differ substantially in morphology and behavior from their hosts so much that the early literature (e.g., Plath 1922) considers Psithyrus as a distinct genus. At points in the past, Psithyrus species have also been argued to have evolved pairwise from their hosts. The contemporary understanding (Cameron et al. 2007) is that Psithyrus comprises a single clade within Bombus, and it is recognized as a subgenus. Thus, in this case the name reflects the currently supported phylogenetic hypothesis, and most bee biologists have come to accept Psithyrus as a part of Bombus.

The readers of Insectes Sociaux are very much affected by controversies over naming social parasites, as many non- taxonomic studies focus on the behavior, ecology and evolution of these insects. It is possible that a student, new to the field, might find that changed names make it difficult to connect older literature with newer work. Importantly, though, to the extent that the naming system is a hypothesis about evolutionary relationships, names that do not embody current phylogenetic knowledge could also mislead a student about evolution. Knowing that new hypotheses about phylogeny will appear in the future and, perhaps affect names, adds tension to the situation.

Name changes make things inconvenient and awkward at times; in my own work on halictid bees I have seen that the shifts between Dialictus and Lasioglossum as the genus name for the same bee¹ have been confusing to biologists outside the inner circle of halictid biologists. On the other hand, the removal of the ant genus Paraponera from the tribe Ectatommini as well as from the subfamily Ponerinae greatly informed my thinking about the evolution of Paraponera behavior, a definite positive outcome of the current reworking of ant systematics.

Why should a journal publish commentaries on an issue like this? The role of scientific journals is to document an extended conversation about theory, hypotheses and facts. My view is that it is far better for scientists to have open, and one would hope civil exchanges about controversial topics. This ensures scientific progress and helps workers outside the immediate field understand seeming contradictions. My personal agreement, or disagreement, with the point of view of either Ward et al. (2016) or Seifert et al. (2016) is much less important than the discussion of the issue within the scientific community. Additionally, one constant concern of journal editors is that the peer review system can be used to suppress work in ways that are difficult to ferret out in the editorial process. Having the conversation in a transparent way should help to advance science and maintain the integrity of editorial processes.


¹ Thus, Lasioglussum zephyrum = Dialictus zephyrus, because the Latin ending of the species epithet changes to match the ending of the genus name. I think this name mismatch inhibits non-specialists from accessing the comparative value of the literature on halictid bee behavior, but the present usage, L. zephyrum, is well justified on systematic grounds. This is a different naming issue than the question about social parasites, but it has many of the same potential outcomes and conflicts in terms of utility.



Cameron SA, Hines HM, Williams PH (2007) A comprehensive phylogeny of the bumble bees (Bombus). Biol J Linn Soc 91:161–188

Michener CD (1964) The possible use of uninominal nomenclature to increase the stability of names in biology. Syst Zool 13:182–190 Plath OE (1922) Notes on Psithyrus, with records of two new American hosts. Biol Bull 43:23–44

Seifert B, Buschinger A, Aldawood A, Antonova V, Bharti H, Borowiec L, Dekoninck W, Dubovikoff D, Espadaler X, Flegr J, Georgiadis C, Heinze J, Neumeyer R, Ødegaard F, Oettler J, Radchenko A, Schultz R, Sharaf M, Trager J, Vesnic A, Wiezik M, Zettel H (2016) Banning paraphylies and executing Linnaean taxonomy is discordant and reduces the evolutionary and semantic information content of biological nomenclature. Insect Soc 63:237–242. doi:10.1007/s00040-016-0467-1

Ward PS, Brady SG, Fisher BL, Schultz TR (2016) Phylogenetic classifications are informative, stable, and pragmatic: the case for monophyletic taxa. Insect Soc doi:10.1007/s00040-016-0516-9


Can ants get your pizza delivery faster?

Editor’s note: This is our first guest blogger post for the Insectes Sociaux blog where our blogger chooses an IS article to write about. Previously, bloggers have written about their own research. I hope you enjoy it.

A blog post highlighting the article written by A.A. Yates and P. Nonacs in Insectes Sociaux

Written by Ravindra Palavalli Nettimi

You are hungry. So you order a pizza. *Yummy, yummy, yummy*

But the delivery person is new to the city. What if he or she could use software to find the shortest path with the fewest turns to get you your yummy pizza as fast as possible? Ants could come to your rescue here!


The pizza delivery problem: how to get to the destination in the most efficient way possible? Illustration: R. Palavalli Nettimi

Many ants face similar maze-like challenge when foraging for honeydew secreted by scale insects in trees. In a recent study published in Insectes Sociaux, A. A. Yates and P. Nonacs from the University of California discovered that ants in a maze take straight line paths with the fewest turns.

To test whether ants can collectively find the shortest paths with the fewest turns, they attached a colony of Linepithema humile ants to a maze consisting of plastic cups connected by plastic tubes and kept some food (mmm… cheese!) in one of the cups as shown below.


A representation of the experimental array used in the study. Illustration: R. Palavalli Nettimi

Initially, the ants explored all the routes equally and laid pheromone trails as they went. The shortest paths to the food ended up getting more pheromone trails since more ants were likely to have found food sooner than the ants exploring longer paths. (In this case, there are three possible shortest paths, two of them are shown in colour). Each ant laid pheromones and also followed the pheromones trails laid by other ants, creating a positive feedback system leading to the shortest paths getting the most pheromone trails.

This phenomenon is known as the travelling salesman problem. The simple rules that the ants use to find the paths have been coded in software used by companies to find out the optimal paths (shortest distances) to deliver milk across many cities or suburbs.

But often finding the shortest path is not enough. The shortest path could involve more turns and thus a higher chance of getting lost. Or the shortest path could have a traffic jam and lead to reduced speed.

Can the ants come to our rescue again?

The researchers showed that the ants prefer the shortest paths (with fewest turns) when exploring to find the cheese. In the figure above, the green path has two turns, while the orange path has one turn to reach the food. The ants were more likely to follow the orange path than the green one. A path with fewer turns can decrease the chance of foragers getting lost. More turns in the path can make it difficult to learn the path and increase the chances of getting lost and wasting foraging time.

It is likely that the preference for the fewest turns could be a consequence of the ‘wall-following’ tendencies of the ants.

Perhaps all of the rules used by ants could be incorporated into the software to not just find the shortest path, but the most efficient path with fewest turns, or highest speed.

Has your pizza been delivered yet?


About the author:
Ravindra Palavalli Nettimi is a PhD student at Macquarie University in Sydney. He writes a blog ( and hosts a podcast called Just-questions ( ). Learn more from his website:

Interview with a social insect scientist: Lori Lach


Lori Lach

IS: Who are you and what do you do?

LL:  I am a mother and a wife and a Senior Lecturer (which is in between an Assistant Professor and an Associate Professor in the North American system) in the College of Science and Engineering at James Cook University in Cairns, Australia. I primarily research invasive social insects. In the past few years I’ve also been researching an emerging disease of honey bees and how it affects foraging behavior. I’ve lived in Australia for nearly 11 years and became a dual national (Australian-US) a couple years ago.

IS: How did you end up researching social insects?

LL: I’d been intrigued by ants during an ecology field course as an undergrad, but never really pursued it because at the time I had no idea how that would lead to a job of any kind. By the time I’d started my PhD years later I had become really interested in the consequences of biological invasions. I had the opportunity to do a summer project in Hawai’i while I was still figuring out what I would research, and while I was there I asked every scientist I met which invaders were the most overlooked and likely doing the most damage. Nine out of ten said ants, and the tenth said rats, so ants it was!


Yellow crazy ants up close. Photo: Dave Wilson

IS: What is your favourite social insect and why?

LL: I’ve been fascinated by yellow crazy ants (Anoplolepis gracilipes) since I first encountered them in Hawai’i at the start of my PhD. They’re just such a conundrum—seemingly so flighty, timid, and disorganized, and yet capable of taking down organisms much larger than they are. Attract a few hundred to a lure, take it away, and it is just mass pandemonium, not the ho-hum retreat of Argentine or big-headed ants. I thought I would get a chance to study them more during my post-doc in Mauritius, but they were really kept in check by Technomyrmex albipes (who would’ve guessed?). But now around Cairns they are a big conservation issue, so I’m in the right place at the right time to work on cracking their secrets.


A mass of yellow crazy ants next to a rainforest creek. Photo: Frank Teodo

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

LL: This is the hardest question. So far, I think discovery-wise it would have to be finding out the dramatic difference just a few invasive ants in flowers could make to the diversity and behavior of floral visitors, and I was able to show that in three different floral systems. When I started this work, most of the literature had been focused on consequences of the extraordinary abundance achieved by invasive ants and their interactions with ground fauna, so I felt like I was breaking new ground. I’ve had a couple people approach me at conferences and tell me they have been inspired by this work in deciding on their own research path. It is the best feeling to know that my discoveries are leading to others.

The best moment so far is right now. I’ve got some really great students working on a variety of really interesting projects, all involving different species of social insects. I also love that my knowledge of ant ecology, and yellow crazy ants in particular, is of direct use in efforts to protect the World Heritage rainforest from this invader. It’s a privilege to work with a really engaged community that supports science, and it’s exciting to have excellent collaborators with diverse sets of complementary skills.

IS: If teaching is part of your work, what courses do you teach? Has your work on social insects helped to shape your teaching?

LL: I had a research fellowship when I first started at James Cook University, so teaching has only recently become a substantial part of my work. I currently teach second year Ecology and a module of Field Ecology, and the occasional guest lectures in Tropical Entomology and Invertebrate Biology. Social insects figure prominently in the examples I use because they can be used to illustrate so many concepts, and really, they’re just so cool. Moreover, knowledge of social insect biology is really useful here in the tropics and can be an asset for graduates seeking employment.

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

LL: “Dark Places” by Kate Grenville. I love reading, and I’m opportunistically working my way through Miles Franklin nominated authors. I’d recommend it for the writing, which is exquisite, but not so much for the story. It was an apt title.

IS: Did any one book have a major influence in shaping your career? What was the book and how did it affect you?

LL: “Ishmael”, by Daniel Quinn is one of several books I read while I was still considering what kind of career path I should follow. “Silent Spring” by Rachel Carson, was another. These books made me take a step back and question what I wanted my priorities to be. My initial plans were to go into medicine, but I ultimately decided that I should pursue a career in which my efforts were not meant to solely benefit humankind.

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

LL: I’m so lucky—I live in between two World Heritage Areas—the Great Barrier Reef and the Wet Tropics rainforest. So snorkelling, hiking, camping, and just spending time outside with my family top the list. On my to-do list for 2017 is to get back into karate. I was once a brown belt, but will now have to work my way up from white again.

W7 2013 082.JPG

Lori and her family.

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

LL: I make sure that I stick with my exercise routine and spend time with my family. Stargazing provides instant perspective. It’s a reminder that we’re all just specks in space and time.

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

LL: My collecting kit, because islands are usually great places to collect invasive ants. A guide book to the flora and fauna, because islands often have weird and wonderful biota. And my journal, with lots of blank pages to fill.

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

LL: I owe a lot to whomever it was at the Australian Research Council who decided to offer prestigious early career research awards that explicitly allowed for career interruptions (e.g., parenthood or “misadventure”). At the time I applied, I had worked part-time for five years following the birth of my son in 2007. Of course I still published during that time, but was unlikely to be competitive for jobs against others who had worked full-time. If I hadn’t been awarded one of those fellowships, it is highly unlikely that I would still be a scientist today. I recently learned that until 1966, a woman working in the public service in Australia was forced to resign if she married. So Australia has come a long way.

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

LL: Take advantage of every opportunity to learn skills in a variety of disciplines—ecology, chemistry, biogeography, genetics, genomics, proteomics, bioinformatics—to name a few, because they will probably all enable you to understand these fascinating creatures that little bit more.


Sociometry may be exhausting, but it’s important and rewarding

A blog post highlighting the article written by M.L. Smith, M.M. Otswald & T.D. Seeley in Insectes Sociaux

Written by Michael L. Smith

I think Walter Tschinkel (1991) said it best when he wrote: “The list [of sociometric data] is not exhaustive, though collecting the data could be exhausting.” My research into honey bee sociometry is a case study in how right he was.

But let’s start at the beginning: what is sociometry? Sociometry is the description and analysis of the physical and numerical attributes of social insect colonies over their lifetimes (Tschinkel 1991). Sociometric data, therefore, is just about anything that you could measure in a social insect colony throughout its life, such as: the size of the nest, the number of workers, the size of the workers, the size of the food stores, the number of sexuals, etc.

Unfortunately, sociometric data are often not collected, and if they are, they’re rarely reported. It’s probably because collecting these data (plus the analyzing and writing) is tedious work. But it’s rewarding, it’s important information that forms the foundation of future research.

My primary interest is reproductive investment in honey bee colonies. In particular, I wanted to know when workers begin to build the large cells of beeswax comb that they use for rearing reproductive males or “drones.” With this question in mind, I set out to conduct a sociometric study, but not just of drone comb, I’d track the whole colony’s growth and development from birth until death. Surprisingly, this had never been done. Many studies had looked at one or two colony parameters throughout a single season, but only a couple had tracked multiple parameters in concert (Lee & Winston 1985; Pratt 1999). The study that tracked the most parameters simultaneously only did so for the first year (Rangel & Seeley 2012), and so missed out on the production of sexuals that occurs in the second year. It seemed like it was time to conduct a broad sociometric study on honeybee colonies throughout their entire life cycles.

To do this, I built and set up four large observation hives, each one about 1m x 1m. These are larger than standard observation hives, and I chose them because I needed sufficient volume (ca. 40L) for the colony to grow to its full size (Seeley & Morse 1976). I then installed into each observation hive an artificial swarm, and monitored the colonies weekly until they died.

Through the glass of the observation hives, I could observe the colonies without disturbing them. I could monitor the number of inhabitants, the growth of the comb, and the contents of the comb, all traced upon a sheet of plastic placed atop the glass of the observation hive. Together with a keen undergraduate, Maddie Ostwald, we tracked honey bee colonies from birth until death while recording worker population, drone population, comb area, comb use (cells holding brood, pollen, honey, or nothing), swarming and secondary swarming events, and time of death. This began in July 2012 and continued until January 2014, and that doesn’t count the time it took to transcribe the comb areas from the plastic sheets!

What do we get at the end of it all? Well, first and foremost, I think it’s a great way to get extremely familiar with your study organism. I grew fond of my colonies, each one with its own personality. One hive was in my office, so I’d hear them buzzing along throughout the day- the perfect office mate. Second, I’m now able to frame my experimental work within the context of these observational descriptions. For example, I now know that although all four colonies built drone comb in their first year, none of them used the drone comb for rearing drones until the second year. Despite having only four colonies, we observed a diversity of life-history strategies, including one colony that attempted to reproduce by producing queen-laid drones in worker cells (the drones were two-thirds smaller than those produced by the other colonies). We also found that drones tend to stay at home when a swarm departs, presumably because they have higher reproductive success at home, but the workers will quickly cull the drones if food stores are low. These highlights, of course, are biased by my interest in drones, so please check out the paper if you’d like to know more (Smith et al. 2016). Lastly, sociometric data are a valuable resource for all social insect biologists, and we cannot conduct comparative analyses without good descriptions of the natural growth and development of many social insect colonies.

I encourage you to think of your favorite social insect species. Is there a paper out there that describes, in painstaking detail, everything that you could possibly count, measure, and describe, from colony founding to colony death? If not, then maybe this is your chance to make it happen!



Lee, P.C. & Winston, M.L., 1985. The effect of swarm size and date of issue on comb construction in newly founded colonies of honeybees (Apis mellifera L.). Canadian Journal of Zoology, 63(3), pp.524–527.

Pratt, S.C., 1999. Optimal timing of comb construction by honeybee (Apis mellifera) colonies: a dynamic programming model and experimental tests. Behavioral Ecology and Sociobiology, 46(1), pp.30–42.

Rangel, J. & Seeley, T.D., 2012. Colony fissioning in honey bees: size and significance of the swarm fraction. Insectes Sociaux, 59(4), pp.453–462.

Seeley, T.D. & Morse, R.A., 1976. The nest of the honey bee (Apis mellifera L.). Insectes Sociaux, 23(4), pp.495–512.

Smith, M.L., Ostwald, M.M. & Seeley, T.D. 2016. Honey bee sociometry: tracking honey bee colonies and their nest contents from colony founding until death. Insectes Sociaux.

Tschinkel, W.R., 1991. Insect sociometry, a field in search of data. Insectes Sociaux, 38(1), pp.77–82.

The winner takes it all, the loser standing small!

A blog post highlighting the article written by Bang and Gadagkar in Insectes Sociaux

Written by Alok Bang

Remember the song by ABBA, ‘The winner takes it all’? In a nutshell, the fate of the dejected lover ABBA portrays can be extended to any conflict. Winners keep winning, and monopolise resources and opportunities. Losers keep losing and forego everything. Or, do they really?

But before coming to that, let’s discuss conflict in animal societies. Why is conflict of utmost interest and importance? For the simple reason, that it is omnipresent. Think of societies most harmonious and in unison, such as those of paper wasps, honey bees, ants and termites – where tens to hundreds and sometimes millions of individuals live and work together – are strewn with conflict. Individuals in these seemingly cooperative societies fight with each other often, over food, mates, territories and other opportunities. Who wins and who loses, thus, has a direct impact on an individual’s survival and reproduction, thereby affecting its evolutionary fitness.

Classically, researchers have focussed on role of individual characteristics such as age, size, weight, hormones and genes, in making winners and losers. While this approach has been important, it has excluded the role an individual’s social environment might play. Environment may influence fighting behaviour, fighting abilities, strength, and finally self-assessment of one’s strength, but this has been largely ignored.

Take the case of self-assessment of one’s fighting ability. When individuals fight, are they winning or losing merely based on their strengths, or does self-assessment of strength influence the outcome of a contest? Human history is laden with examples of an underdog, who is physically average or even weak, defeating a stronger opponent, because of a heightened perception of her strength. Similarly, a strong individual is known to lose a contest if she has a diminished self-assessment of her strength. Self-assessment can thus be influential – as much if not more – than the actual strength, in deciding the outcome of a fight.

In the past two decades, the phenomenon of winner-loser effects have come to the forefront of such enquiries into external determinants of fighting abilities and contest outcome. Simply put, they refer to an increased probability of winning or losing a contest based on prior experience of winning or losing, respectively, even if everything else is randomised. A prior experience of winning may enhance and a prior experience of losing may diminish an individual’s perception of its own fighting ability; thereby, affecting the contest outcome. Such studies have been mostly performed in vertebrates and research on the role the environment plays in conflict outcome in invertebrates is severely lacking.


A typical Ropalidia marginata colony. R. marginata is found abundantly in peninsular India. Photo credit: Thresiamma Varghese.

In the first study of its kind that investigated the role of prior experience on current contest outcome in a eusocial species, we chose the primitively eusocial Indian paper wasp, Ropalidia marginata as the model system. To control for a wasp’s environmental experience, the focal individuals to be included in the experiments had to be devoid of any prior fighting related experience. This was achieved by bringing adult-less colonies of R. marginata into a controlled environment, keeping thorough census records of all individuals being born on a colony, and isolating these individuals as soon as they were born.

The other important step was the method of choosing focal individuals for winner and loser effect experiments. We achieved random-selection by giving pre-decided contest outcomes to random focal individuals in the first contest, independent of their intrinsic strengths. We chose this method because these experiments aim to investigate the effect of experience, and not strength, on the contest outcome.


A lone R. marginata female. R. marginata females, like in many other paper wasp societies, can found new colonies individually as well as in a group. Photo credit: Alok Bang

In experiments performed to investigate winner effects, a pre-decided winning experience was given to a random focal individual by pairing it with an extremely weak individual (termed habitual loser) of the population. This ensured that the pool of focal individuals used for testing winner effects did not include only strong individuals, but included individuals with a wide range of intrinsic strengths. Similarly, in experiments to investigate loser effects, a pre-decided losing experience was given to a random focal individual by pairing it with an extremely strong individual (termed habitual winner) of the population. This, in turn, ensured that the pool of focal individuals used for testing loser effects did not have only weak individuals, but included individuals with a wide range of intrinsic strengths. The focal individuals with such pre-decided contests were then paired with a random naive individual in the second contest.


R. marginata females engaged in a dominance-subordinate interaction. The behaviour displayed here is called ‘sit over’, where the dominant individual sits over the subordinate individual and renders her immobile. Fights such as these are quite common in R. marginata. This helps to establish a dominance hierarchy between individuals, and has important implication on survival and reproduction of individuals, and work regulation in the colony. Photo credit: Alok Bang.

Each experiment thus consisted of a first contest between a focal individual and a habitual loser/winner, giving it a pre-decided contest that occurred for one hour, followed by a 45-minute gap, which then was followed by a second contest of one-hour between the focal individual with a random naïve opponent. In such an experimental set-up, a second successive win (or loss), in significantly more than half the cases, would indicate that the individuals’ self-perception was impacted due to their prior experience. During all these contests, dominance-subordinate interactions between individuals were observed, and winners and losers of the contests were declared. All experiments were carried out  blind.

We indeed found that there was a significantly high number of pairs in which a win was followed by a second win, and a significantly high number of pairs in which a loss was followed by a second loss, indicating that both winner and loser effects are present in the Indian paper wasp, R. marginata.

Winner effects may evolve due to advantages associated with winning, but why would a species evolve loser effects? Moreover, how do two such apparently opposing phenomena concurrently exist in a species? Winner and loser effects are most likely independent or even interdependent effects. If self-assessment of winners and losers are independently advantageous, these effects would exert independent feedback loops on individuals and co-exist. For example, winning a contest may allow winners a higher access to resources and mates, thus developing and reinforcing winner effects. Losing a contest on the other hand, though seemingly disadvantageous, may allow individuals to forego costs associated with fighting such as injuries, exhaustion and death. If the benefits of avoiding these costs are much higher than the benefits acquired from winning, loser effects will simultaneously develop in the population.

Here we show that wasp behaviour is not only governed by their own internal constitution, but to a considerable extent by their surroundings. The role of external and social determinants of behaviour balances the hitherto unduly skewed importance given to individual characteristics.

Finally, is the Indian paper wasp R. marginata a unique and only eusocial species that displays winner-loser effects? It is definitely the first eusocial species, but we believe it will not be the last. The uniqueness of R. marginata in this regard may have less to do with ecology of the species, and more due to lack of such investigations in other social insects. This study should steer efforts towards finding the presence, extent and longevity of winner-loser effects in other social species. A comparative approach to studying proximate and ultimate factors governing winner and loser effects in social species will be key to understanding sociobiology of group living animals.

Facultative slave-making ants tolerate alien slaves but not their masters

A blog post highlighting the article written by Włodarczyk in Insectes Sociaux

Written by Tomasz Włodarczyk

Many ant species in nature are closely associated with other ant species. The closest form of such an association is called a mixed colony where ants of both species inhabit common nest, share food and raise their brood side by side. Mixed colonies arise as a result of social parasitism when one species exploits the labor of the other, such as in slave-making ant species. Slave-maker ants raid the nests of the host species, steal the pupae and bring them back to their home nests. Newly-emerged individuals integrate into the parasite’s society and perform all domestic duties.

In the lab, we can also create mixed colonies using species that would never form such an association in nature. The species don’t even have to co-exist geographically. This is because ants learn (imprint) colony odor after eclosion from pupae and use it as a template for subsequent nestmate recognition. Thus, by putting together callow ants of different species we can create a mixed colony of individuals that have imprinted on the odor of each other.

As a part of my PhD project I investigated the recognition behavior of ants using a colony of the facultative slave-making ant species, Formica sanguinea. By supplying them with pupae of Formica polyctena or F. rufa -which soon emerged- I formed mixed colonies (Włodarczyk 2012, Włodarczyk and Szczepaniak 2014). These experiments were inspired by the studies conducted by Wojciech Czechowski (1994) whose results suggested that F. sanguinea ants acquire their recognition signature form their slaves, as in the obligate slave-making species Polyergus samurai (Yamaoka 1990).


Formica sanguinea is a facultative slave-making species enslaving ants from the subgenus Serviformica. Here, a colony with black F. fusca slaves has been excavated.

Later I became curious about how things look in the case of F. sanguinea colonies containing the most frequently used slave species, F. fusca. Results of chemical studies revealed that odor of F. sanguinea ants is quite different from that of its host species (Martin et al. 2008, Włodarczyk and Szczepaniak in prep.). Moreover, we found that enslaved F. fusca ants develop a chemical recognition signature which is intermediate between that of their parasite and ants from free-living colonies (Włodarczyk and Szczepaniak in prep.). This raised the question about how recognition cue diversity in F. sanguinea colonies affect the recognition abilities of ants. Even more interesting was whether there are differences in the recognition abilities between F. sanguinea and F. fusca ants given that the parasite is the only party to be under selective pressure to live in such a condition.

I collected eight queenright F. sanguinea colonies containing F. fusca slaves and maintained in the the laboratory. The slave-making F. sanguinea ants and their slaves were exposed on a Petri dish to anesthetized ants from alien colonies. I measured the number of aggressive behaviors in various encounter combinations. I showed that F. sanguinea ants are able to discriminate other individuals from the same species from alien colonies towards which they exhibit aggressive behavior. However, slaves from alien colonies were generally tolerated. This result supports the hypothesis that F. sanguinea ants are intrinsically tolerant to individuals whose odor indicates that they are slaves. Otherwise slave-making ants might accidentally attack their own slaves, which possess a recognition signature that deviates from that of the other slaves. This situation would arise when slaves from new source colony appear in the slave-maker’s society.

The other result was that slaves (F. fusca) are poor at discriminating slave-making ants and slaves from alien colonies and do not exhibit an overt aggression toward them. This could be explained by the high within-colony recognition cue diversity that hampers formation of an accurate template during colony’s odor learning phase. This is intuitive explanation since it might be hard to recognize an object of a given class when this class is relatively heterogeneous. Thus, there is no recognition barrier for F. sanguinea ants to take over slaves from alien colonies. However, such a phenomenon has not been recorded for F. sanguinea ants. Therefore we can hypothesize that intraspecific raids play at best very limited role as a way of slave gaining.


Formica fusca slaves showing aggressive behavior towards an anesthetized conspecific ant.

Moreover, I conducted an experiment in which slaves and slave-makers were reared separately. After about 2-month period, ex-slaves elicited aggression in ants from stock colonies (both in slave-makers and in slaves). Conversely, slave-makers separated from slaves were still treated as nestmates. This result suggests that F. sanguinea exert a strong impact on the odor of F. fusca ants, possibly by the transfer of recognition cues during food exchange.

The results of my study highlight that selective pressures associated with different life histories can lead to differences in recognition systems between social insect species.



Czechowski W (1994) Impact of atypical slaves on intraspecific relations in Formica sanguinea Latr. (Hymenoptera, Formicidae). Bull Pol Acad Sci 42(4):345–350

Martin SJ, Helantera H, Drijfhout FP (2008) Evolution of species-specific cuticular hydrocarbon patterns in Formica ants. Biol J Linn Soc 95:131–140

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Interview with a social insect scientist: Mark Brown

IS: Who are you and what do you do?

MB: Mark Brown. I’m a Professor at Royal Holloway University of London, where I lead a research group that seeks to understand host-pathogen interactions in bumblebees, as well as aiding in the conservation of bumblebees. We also enjoy investigating other aspects of social insect biology.

IS: How did you end up researching social insects?

MB: In my 2nd year at university, I was lucky to have Deborah Gordon – ant biologist – as one of my tutors. After teaching me for a term, she asked me if I’d like to come and work for her as a field assistant in the desert in Arizona for the summer. I said yes, to a large degree because I thought it was an opportunity to combine biology with travel. But then I met the ants and fell in love (with the ants, I should add!), and since then it’s been social insects all the way!

IS: What is your favourite social insect and why?

MB: Can I ask for two? The first – Messor andrei – was the subject of my PhD research. They’re beautiful black ants, who make a habit of carrying seeds like parasols back to their nest. The second – Bombus lapidarius – I have yet to work on, but they’re a beautifully smart bumblebee that make the most elegant nests.


Bombus lapidarius, one of Mark’s favourite species. Photo credit: Jürgen Mangelsdorf / flickr

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

MB: That’s a tough one. Discovering that the parasite Crithidia bombi had a major impact on bumblebee fitness has to be up there, as before then it was seemingly a parasite without virulence. However, I think that work with Matthias Fürst, Dino McMahon, Juliet Osborne, and Robert Paxton, where we showed that honey bee pathogens spill over into wild bumblebee populations, is at the top. Understanding the dynamics of viral diseases in the field has important practical implications, as well as being exciting from a pure research perspective, and so our finding has had a real impact on the field.

IS: If teaching is part of your work, what courses do you teach? Has your work on social insects helped to shape your teaching?

MB: I teach courses in invertebrate biology, conservation biology, and a field course in ecology and conservation on the island of Samos, Greece. It’s surprising how often my examples involve social insects of one kind or another. 😉

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

MB: “A Place of Greater Safety” by Hilary Mantel. A character-focused history of the firestorm that was the French Revolution, this is definitely worth reading! Mantel writes incredibly incisively about people and their motivations, and how this shapes history. For anyone who wants to understand the politics of science, and how this can impact careers and the trajectory of science itself, this is a great primer.

IS: Did any one book have a major influence in shaping your career? What was the book and how did it affect you?

MB: I read “The Trouble with Lichen” by John Wyndham when I was a teenager. This inspired me to become a research scientist (in particular, a biochemist, which lasted only until I realised that the ‘bio’ aspect was rather limited), and also to recognise that gender has a significant impact on recognition and career advancement in science (this was long before I’d heard of Rosalind Franklin). We still have a long way to go to make science a level-playing field for all genders and orientations, but it’s a goal we have to reach.

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

MB: Reading fantasy novels, and spending time with my nieces. It would be walking safaris through the Zambian bush, but I can’t afford to do it often enough to call it a hobby!

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

MB: I remember that I’m a very lucky man – I have a family, friends, and a job I love – and try to focus on the day-to-day until I get my perspective back.

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

MB: My husband, good Swiss chocolate, and an endless supply of paperback books (none of these need explanation!).

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

MB: My PhD supervisor, Deborah Gordon, taught me how to look at ants, and how to think and write like a scientist. Paul Schmid-Hempel, my post-doc boss, introduced me to the intriguing world of host-parasite interactions, and also taught me how to play the scientific game. I owe them both a huge debt.

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

MB: Get outside and watch the animals. If you can spend hours watching ants or bees (apologies to my termite and wasp colleagues!), and still come away fascinated, then you’ve got a good foundation to build on. If you decide it’s not for you, get another job, earn loads of money, and set up a charitable foundation to fund the research you’d like to see done.


Messor andrei, Mark’s first ant love. Photo credit: photographer (unknown) and / Wikimedia commons


Two views on invasive ants

Highlighting the articles written by Calcaterra et al. and Santos in Insectes Sociaux

Written by Insectes Sociaux Editor in Chief, Michael Breed

Species ranges are often defined by natural barriers such as lakes, rivers oceans or mountains. Human actions can break down natural barriers or transport species across those barriers- creating biological invasions with serious consequences. Social insects play a large role in the overall picture of biological invasions, with many species of ant, including fire ants, Solenopsis invicta, and Argentine ants, Linipithema humile, wreaking havoc on ecosystems. Wasps, such as Polistes dominula and Vespa velutina, have gained serious footholds in non-native habitats. The Formosan termite, Coptotermes formosanus, is equally a scourge. Even the honey bee, Apis mellifera, whose introduction into the New World is often seen as a benign event and whose current presence is valued for its role as a pollinator, has had dramatic effects on native bees and possibly on plants that rely on native bees for pollination.

In this issue two papers highlight different aspects of social insect invasion biology. Calcaterra and colleagues (2016) take on the original community roles of ants that become invaders by comparing ecological dominance among ants in both native and introduced communities. They take advantage of the natural occurrence in Argentinian ecosystems of several prominent invasive ant species, including two of particular note, Solenopsis richteri and L. humile. These two species are ecologically co-dominant in the Otamendi Natural Reserve, near Buenos Aires, Argentina, with S. richteri being numerically the most abundant ant and L. humile being the best at recruiting colony mates to new food sources. However, these ants co-exist in this reserve with 47 other species of ant, including several that are also invasive, suggesting that ecological resistance to the qualities that favor invasive success have evolved among ants in these communities. Competition may not be the primary factor in structuring these communities, reducing the impact of species with high competitive abilities. These two invasive species were abundant across several habitat types, suggesting that they are ecologically flexible. Three non-invasive species were also dominant in the habitats, meaning that the characteristics that lead to invasiveness are not the sole qualities that lead to dominance in a native habitat, although the invasive potential of these other species is unknown.

Santos (2016) reviews the roles of ants in urban ecosystems. In a meta-analysis based on over 100 papers published about ants found in urban environments, Santos found that many known invasive ants, including S. invicta and L. linipethema are also prominent in the literature on urban ants. Other invasive ants occurring in urban environments include the pharaoh’s ant, Monomorium pharonis, and Tapinoma sessile. The majority of the studies of urban ants came from the United States (with a bias towards southern and northeastern states) and Brazil (with a bias to southeastern Brazil). While urban ants are more often aesthetic pests rather than vectors of disease or destroyers of food supplies, some species present public health problems by stinging or being incidental purveyors of bacteria and fungi. Their presence in urban environments is thus a concern due to public demand for control and in some cases public health concerns.

For those of us whose interest lies in the beauty of social insects’ complex systems of communication and division of labor, as well as their major significance in terrestrial ecosystems rather than in applied entomology, reading these two papers is a dark reminder that introduced social insects can have significant negative impacts on both humans and natural communities. By the translocation of ants across geographic barriers that would be otherwise impassable for ants, humans have created real problems of immediate importance. Transfers of social wasps, social bees, and termites have happened in much the same ways as for ants. These movements have largely resulted from inadvertent actions or carelessness with shipment sanitation and quarantine. We cannot unreel these past actions; in fact, more than half a century of futility in trying to control fire ants and the “Africanized” version of the honey bee suggest the irreversibility of these introductions. Biologists who work on social insects should have much to bring to discussions of how to ameliorate the impacts of social insect populations as they become established in natural communities and in urban environments.


Calcaterra L, Cabrera S, Briano J (2016) Local co-occurrence of several highly invasive ants in their native range: are they all ecologically dominant species? Insect Soc 63:**-** DOI 10.1007/s00040-016-0481-3

Santos MN (2016) Research on urban ants: approaches and gaps. Insect Soc 63:**-** DOI 10.1007/s00040-016-0483-1


Editor’s note: If you are interested in Luis Calcaterra’s explanations of how invasive ants are like tango dancers, please click here for his blog on his work.

Montane ants use their bodies to make warm homes for themselves (and others)

A blog post highlighting the article written by Baudier and O’Donnell in Insectes Sociaux

Written by Kaitlin Baudier

Army ants are predatory and nomadic. To find food, the army ant Labidus praedator marches in carpet-like swarms both on the forest floor and underground, very different from most ants that you likely imagine walking in neat rows. Army ants have temporary nests that they can move around called bivouacs. Bivouacs are constructed from clustered, interlocking bodies of worker ants that are not foraging for insect prey. Bivouacs can either be free hanging aboveground or within a cavity or series of cavities belowground. In fact, other arthropod species live within the bivouacs. Army ants are important ecological keystones in the tropics, largely because of the other arthropods that rely on them for food and shelter while living within army ant bivouacs.

Bivouacs allow the ants to thermoregulate- warming and reducing temperature variation to accommodate the specific thermal needs of the colony’s maturing brood (eggs, larvae, and pupae) suspended within the bivouac. However, of the approximately 1,000 species of army ant in the world, all of our understanding of bivouac thermoregulation comes from only two species that form aboveground bivouacs. Research has shown that aboveground army ant bivouacs at low elevations can actively warm brood, but until now underground and high-elevation bivouacs were poorly understood.

Excavated soil bivouac

Figure 1: Excavated soil mounded atop an underground bivouac of Labidus praedator (Photo credit: Sean O’Donnell)

Montane bivouacs of L. praedator have a different structure than lowland bivouacs. Though lowland L. praedator have historically been observed nesting in preexisting cavities, often with other species of army ant, these high-elevation ants construct largely self-excavated bivouac chambers at the base of trees, seemingly independent of other army ant species (Figure 1).

In our study, we examined underground, lower montane L. praedator bivouacs to ask:

  1. Do below-ground army ants warm their bivouacs?
  2. How do thermal tolerances of ants and inquilines (other arthropods that reside in the bivouacs) compare to bivouac temperatures?
  3. Are high elevation bivouacs challenged by cold?

In order to find out, we placed temperature and humidity loggers at different depths in an underground bivouac of Labidus praedator and also in adjacent soil. After five days of recording we excavated the bivouac, keeping careful notes on depths at which we found brood, young adult workers, mature adult workers, and inquilines.


Next, we collected worker ants and inquiline millipedes as subjects of thermal tolerance assays to estimate the range of temperatures tolerable by each group. We exposed the ants and millipedes to either a range of increasing or decreasing temperatures. The most extreme temperature for which individuals were able to move was considered either its maximum thermal tolerance, or minimum, depending on the test each individual received. We then compared these thermal tolerances to environmental temperatures to see if any of the colony members were “thermally challenged” in these high elevation underground bivouacs.


Figure 2: Subjects compared for thermal tolerance (A-B) bivouac-dwelling millipede (Calymmodesmus sp.), (C) host adult Labidus praedator worker (mature), (D) a ‘callow’ or recently-molted young adult L. praedator worker


We found that L. praedator bivouacs were significantly warmer than reference soil. Temperatures within the bivouac were on average 6.2°C warmer than soil at similar depths, and more than 8°C warmer than average nighttime surface lows. Both reference soil temperature and bivouac temperature varied little at depths of 40cm below the forest floor. Together these two findings suggest that army ants may passively use soil to buffer against diel fluctuations in temperature, while actively warming the nest with their collective metabolic heat. Below-ground bivouacking species of army ant are able to survive at both higher latitudes and elevations than their highly-studied aboveground relatives. The thermal buffering effects of soil are likely key to creating a homeostatic and warmed brood environment despite seasonal and daily low temperatures.

Thermal tolerances of bivouac-resident millipedes and ants varied, and this variation corresponded to placement within the bivouac. More cold-sensitive individuals were located deeper in the bivouac where they experienced hotter and less variable temperatures. Callow and mature L. praedator workers had the same heat tolerance, but callow workers were more susceptible to immobility at low temperatures. The most cold-sensitive, however, were the millipedes, many of which had minimum tolerable temperatures at or above temperatures occurring on the bivouac surface. This suggests that warmth within army ant bivouacs may be an added benefit for arthropods that live in bivouacs at high elevations, enabling some bivouac-dwelling species to occupy geographic ranges that they would not be able inhabit without the ants and their bivouacs.

Editor’s note: You can find the video above and many other ant videos on Kaitlin Baudier’s YouTube channel AntGirl

Interview with a social insect scientist: Miriam Richards

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Moonshine, Miriam and Blueberry. Photo credit: Michele du Moulin

IS: Who are you and what do you do?

MR: Miriam Richards – I am a professor at Brock University, in the Niagara Region of southern Ontario, Canada. I teach courses in Animal Behaviour, Ecology, and Evolution. I do research on the social behaviour, ecology and evolution of carpenter and sweat bees, and on restoration of local bee communities. I also spend a lot of time gardening and communing with farm animals – this fits right in as applied ecology and animal behaviour.

 IS: How did you end up researching social insects?

 MR: I’ve always been interested in animals and as far back as I can remember thinking about what I would be when I grew up, it was related to animals. By grade 8 I wanted to study animal behaviour and ecology, but I was completely focussed on vertebrates. I first thought about comparative social behaviour in bees in my third year Animal Behaviour course (the same one I teach now!) when I wrote an essay on the evolution of sociality in bees. That was about 1982, and I must have read Michener’s work, but I don’t have the essay any more to check it out. And besides, I was fully occupied studying common tern behaviour (my undergrad thesis and first publication) and then snow goose demography (my MSc thesis). And I really wanted to be a zookeeper and breed endangered species.

My MSc experience was traumatic and nearly the end of my career in science. We decided to have a baby during grad school, because in the mid-1980s, being a pregnant post-doc or new faculty member was clearly a stressful thing. Unfortunately, being a pregnant grad student also turned out to be stressful – such a thing was practically unheard of then. When my supervisor learned I was pregnant, he told the lab technician that this proved I wasn’t serious about research. A member of my committee informed me (and several other members of the lab) that women with small children really couldn’t do science. I managed to finish my MSc, but decided to abandon research as a career.

Over the next couple years, I worked at a pet store, as a secretary for my husband’s PhD supervisor at York University in Toronto, then as a zookeeper at the Metro Zoo in Toronto, thought about various options, like medical school and environmental law, and realized they were not for me. While I was a secretary, I heard about a new professor who had just been hired at York, who studied bees and was interested in doing molecular ecology (although it was not called that yet). This was Laurence Packer, and after I realized that being a zookeeper meant feeding and cleaning, not saving animals from extinction, I finally made my mind up to do a PhD. I joined Laurence’s lab as his first grad student. That was a fateful decision – I got to know bees, and I adore the science of social evolution. And after a few more years of fateful twists and turns, I got really lucky and landed a job at Brock. And here I am!

IS: What is your favourite social insect and why?

MR: My favourite social insect is Xylocopa virginica. Talk about charismatic microfauna! It has everything going for it: it’s big enough to see and readily identifiable. It lives in social groups that are organized in ways that are very different from eusocial bees, even from the primitively eusocial bees that we still love to study. It is becoming very common in southern Ontario, so it is easy to find. The males also have interesting social behaviour. And the males’ antics as they guard territories are hilarious.


Xylocopa virginica at her nest entrance. Photo credit: Lyndon Duff

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

MR: That’s a hard question, because we have lots of great moments, and I don’t look back on great moments and compare them to now. I really like when we find some new behaviour, especially something that is unusual or contradicts a major theoretical paradigm, but I am not sure what I’ve done that is more important. I think the best moments are when students confidently and authoritatively describe their work, or when they stand up to my arguments and bring me over to their point of view. Then I see that they have become scientists, and I feel like I’ve done my job.

 IS: If teaching is part of your work, what courses do you teach? Has your work on social insects helped to shape your teaching?

 MR: I’m in a very small department, so I teach all sorts of general topics, from introductory ecology and evolution, to animal behaviour, molecular ecology, behaviour genetics, etc. I use social insect examples in my teaching all the time, including analysing data sets that we’ve already published (or not). More importantly, my teaching has shaped my research. For instance, there are major differences in the way scientists think about insect versus animal sociality, and sometimes I think that insect sociobiologists need to broaden their definitions a bit. Our entire line of research monitoring bee communities in restored landfill sites was largely inspired by my teaching community and ecosystem ecology, fields way outside my research comfort zone.

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

MR: I mostly read murder mysteries, for their escapist value. Every so often I read a science book, but only in the summer when I can focus. I am about to read Clutton-Brock’s new book on mammal societies.

 IS: Did any one book have a major influence in shaping your career? What was the book and how did it affect you?

MR: Michener’s 1974 book is one of my favourites – I still use it when I need a quick synopsis of the social behaviour of some group that I don’t really know much about. In the last few years, I’ve begun to realize that it is getting out of date, but it is 40 years old, after all. I still recommend it to new students, though.

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

MR: The absolute joy of our lives is our 2-year old granddaughter, but it’s a long distance relationship that is maintained by Skype and visits every few months. I love being outside tending my garden and my animals. Recently, I bought a mule named Moonshine. He’s so much fun to ride through the woods. I am also training one of my donkeys to become a saddle and cart donkey for my grand-daughter.

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

MR: One of my great discoveries in life is that my outlook is actually quite independent of my circumstances. For instance, I am often tired, cranky and depressed in February. It’s easy to find reasons to explain this to myself, so I try to remember that I am one of the luckiest people on the planet. I have a great job, I live in a great house on a fabulous little hobby farm, we’re all healthy and busy, my graduate students generally have been successful and done interesting work, my undergraduates sometimes tell me I inspire them, and every so often somebody is appreciative for the work I do. Of course I have low moments, but very few of them are “real”, just irritating foibles of brain chemistry, I think.

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

MR: I need to bring my husband, my camera, and a guidebook. I need the guidebook so I don’t miss anything great, the camera to record it because I look at things more carefully when I take pictures, and my husband, because it’s always more fun when he’s along, too.

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

MR: Laurence Packer, my PhD supervisor, set me on my current path. He also helped me stay on that path, and has helped me to new opportunities over the years. And my husband, Adonis Skandalis, who is a molecular biologist and my chief scientific advisor, collaborator, and sounding board, even though we work on completely different topics.

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

MR: I encourage them not to put all their eggs in the academic basket. Research jobs are hard to come by and social insect jobs are harder still. Academic positions are the most obvious path into research, but not the only path. I think the incredible growth of citizen science will present a lot of opportunities for people with scientific training to do science, even if they don’t get to call themselves scientists on their tax returns. Science is a way of knowing – and a way of life, too. Let’s go back to a time when scientists were people who made discoveries using scientific method, even if you earn a living doing something else.



Xylocopa virginica female foraging. Photo credit: Lyndon Duff