Category Archives: Science

The Ant Daisy Chain, Described


You may remember a recent viral video showing an undescribed predatory ant behavior. It turned out that myrmecologists Christian Peeters and Stéphane de Greef had observed these ants in the field, but they’d not assembled enough data to publish, leaving us all in the dark about how the ants coordinated the amazing millipede-hauling chains.

With a bright internet spotlight on the behavior, Christian and Stéphane returned to the project and have just published a detailed description in Insectes Sociaux. The video alone is worth a thousand words:


source: Peeters C, De Greef S. 2015. Predation on large millipedes and self-assembling chains in Leptogenys ants from Cambodia. Insectes Sociaux doi: 10.1007/s00040-015-0426-2

A Gallery of Pollinators



It probably shouldn’t have taken ten years, but I’ve finally corralled a few of my better pollination photographs into a coherent gallery. Check it out at the link:

A Gallery of Pollinators and Pollination

The astute observer will notice I’ve put flies at the top. This is no accident. Flies are important pollinators, but as Morgan Jackson recently pointed out, they are unjustifiably neglected in favor of the more popular bees.

Another Reason Why Ants Don’t Have Traffic Jams



There’s a story going around about how ants never have traffic jams. It’s a fine bit of science about how ants adjust their speed and don’t overtake each other, leading to smooth, fast speeds down the formicid freeway.

But I wouldn’t recommend humans adopt ant traffic management strategies. The physics of being small gives ants an enormous advantage.

Ant are tiny compared to us, and collisions at their size involve vanishingly small forces (remember F = [m][a]?), even at high speeds. Ants can cruise along with nothing to lose; the cost of veering into an errant stone or oncoming traffic is a slight bump, barely felt. They pick up and move along.

Cars? Not so much. You can imagine how much quicker rush hour traffic would move if drivers were immune to accidents, fearless and fast. People would drive like ants.

The Puppy Spider Is Not As Big As You Think


While we’re on spider debunkings, Piotr Naskrecki’s “puppy spider” story is making the media rounds this week, spawning the usual juvenile Nopes and Kill It With Fires! With every re-telling the spider gets bigger, of course.

Not to rain on the Goliath spider parade, but what Piotr originally reported was:

they weigh up to 170 g – about as much as a young puppy.”

170 grams is big. By arthropod standards, it’s huge. But the spider is not, say, the Shelob people are imagining. For perspective, I’ve marked 170 grams on a standard puppy growth chart.


Puppy spider is not a 3 week old Labrador. It’s the weight, at birth, of a Chihuahua. Big, sure, but you’re not going to saddle this thing up and ride it to work.

The spider looks huge in the photographs for another reason, too. Piotr used wide-angle lenses that exaggerate the size of foreground relative to background objects. This wide-angle macro effect is a technique that Piotr does especially well. And it makes big spiders look larger than life.

Anyway, if you’d like one of Piotr’s puppy spiders for your living room wall, he’s just put a print on sale for prices so low he can’t be making any money:

Photo copyright Piotr Naskrecki, all rights reserved.


An Evolutionary Transition, In Vivo


Among the most dazzling products of insect evolution are leafcutter ants, which cultivate an edible fungus on a compost of fresh vegetation. The ants’ digestive chemistry is so simplified that they can only eat the fungus that grows in the underground gardens.

The leafcutter/fungus system is complex enough to seem highly improbable, and indeed, it appears to have evolved only once in the 130 million year history of ants. How could such a complex system appear?

The system did not spring forth fully-formed, of course. I was reminded of the gradual evolutionary transition on our recent BugShot course in Belize, when I happened across Trachymyrmex intermedius.

Trachymyrmex intermedius


Contrary to appearances, T. intermedius is not a leafcutter ant.

At least, not technically. True leafcutters belong only to the genera Atta and AcromyrmexTrachymyrmex is instead the sprawling, paraphyletic genus from which the leafcutters arose. These ants also farm fungus, but they typically use dead vegetation, caterpillar frass, and other bits of detritus. Like so:


Green vegetation is not the usual fare for Trachymyrmex, but T. intermedius and several others do take it on occasion. Seeing a few of these small ants trundling off with a harvest more fit for their larger cousins was just a reminder that animal behavior is naturally variable, and that variation is what allows the evolutionary process to explore new paths.

Little ant, big thoughts.

Did LiveLeak Just Leak An Unknown Ant Behavior?


Those corners of the internet prone to viral outbreaks are abuzz today with an intriguing ant video:

Is it real? Yes.

The quality isn’t great, but the clip appears to show an Asian Leptogenys daisy-chaining their bodies in parallel lines to haul away a large millipede. I have spent the morning searching the technical literature for mention of this unusual behavior, and am coming up empty. Some Leptogenys species, including L. diminuta, L. nitida, and L. processionalis, are known to forage in groups and transport prey “cooperatively” (source, source). What is meant by “cooperative” is often vague. (For more, see this excellent recent review of cooperative transport by Helen McCreery).  Yet I didn’t find any explicit description of workers linking up, mandible to abdomen, to pull together.

Is ponerine daisy-chaining an unknown behavior? Possibly. It is also possible my search skills aren’t up to the task. If you know of a description of it, please drop a note in the comments. I am not the only one interested, either:


I did, however, happen across a higher quality video from a Cambodian beekeeper:

I presume the swelling music helps motivate the ants to pull harder. But, I digress.

Steve Shattuck took a photograph recently in Borneo capturing a variation on this behavior, with workers forming a chain by biting the legs of a preceding ant.


Again, I don’t think the behavior has been formally described beyond this smattering of visual media.

Regardless of documentation, daisy chaining raises some definitely unanswered questions and will make a fine Ph.D. thesis for some lucky student. How do ants organize themselves in chains? What cues do they use? How do they know to let go? Is chaining employed only for particular sizes or species of prey? How does the behavior effect overall foraging efficiency? What are the evolutionary precursors to chaining? And, do these ants have any other tricks up their coxae?

 ***Update 8/30/2014 – 

In the comments, Roberto Keller suggested that the eminent ponerophile Christian Peeters might know something. And indeed, Christian emails in with the following:

I observed this fascinating behaviour in Cambodia 4 years ago. Stéphane De Greef was with me and some of his photos are attached.

The behaviour was very stereotyped: mandibles grab preceding ant’s gaster (between first and second segment).

Seiki Yamane identified it as Leptogenys sp. 47, closely related to L. chalybaea described from Borneo by Emery (but stronger sculpture especially on gastral tergites).

The millipedes were 130mm long, identified as order Spirostreptida (Diplopoda). Ant is 16mm long.

Back then I reviewed the literature and found no other record of chain behaviour in Ponerinae. No record of millipede predation in Leptogenys.  Specialized hunting on millipedes is restricted to Thaumatomyrmex, Probolomyrmex and Gnamptogenys, but these are solitary hunters on a very different kind of millipedes (polyxenids).

I started writing a ms on this behaviour (formation of chains in ants through a self-assembling behaviour) but sadly I have not been able to get further observations. It seems to happen at certain times of the year only.

By an amazing coincidence, two days ago I finished fieldwork in northern Thailand and came across the same Leptogenys species. There were cleaned out ring segments of big millipedes outside entrances. Unfortunately I did not observe any raids.

Image by Stéphane De Greef, used with permission.

Image by Stéphane De Greef, used with permission.

postscript: The virality of the video also illustrates both the good and the bad about the internet. The good, of course, is that this fascinating ant behavior found its way in front of scientists who otherwise might not have seen it. On the other hand, the viral nature of the video means that actual person who filmed it is drowned out among the hundreds of uncredited, unsourced copies. Securing the information about where and when the video was taken, and verifying the species, is going to be difficult. This is one reason why crediting sources online is important. Lose the credit, lose the data.

Ant Research Roundup: Parasites Edition


One measure of the importance of ants is the number of parasites that have evolved to exploit their abundant resources. This week has seen a cluster of new ant parasite studies. Among them:


Formica subintegra, photographed while raiding in upstate New York.

1. Socially parasitic Formica move nests during raiding season to richer hunting grounds. From the abstract of Apple et al (2014):

Five summers of monitoring the raiding behavior of 11–14 colonies of the slavemakers Formica subintegra and Formica pergandei revealed relatively frequent nest relocations: of 14 colonies that have been tracked for at least three of 5 years, all but one moved at least once by invading existing host nests. Movements tended to occur in the middle of the raiding season and were typically followed by continued raiding of nearby host colonies. Spatial patterns of movements suggest that their purpose is to gain access to more host colonies to raid.

source: Apple, J.L., Lewandowski, S.L., Levine, J.L. 2014. Nest relocation in the slavemaking ants Formica subintegra and Formica pergandei: a response to host nest availability that increases raiding success. Insectes Sociaux, doi: 10.1007/s00040-014-0359-1


Myrmica queen in Arizona covered in mites.

2. Myrmica just can’t catch a break. A literature survey by Witek at al reports an array of about 40 parasites on these common holarctic ants, including butterfly larvae, socially parasitic ants, fly larvae, fungi, and others.

My sense is that Myrmica isn’t unusual in its parasite load; rather, Myrmica geography has lent itself to observation by natural-history obsessed northern Europeans. As myrmecology advances elsewhere, plenty of other ants will turn out to have lives just as miserable as Myrmica‘s.

source: Witek, M., Barbero, F., Markó, B. 2014. Myrmica ants host highly diverse parasitic communities: from social parasites to microbes. Insectes Sociaux, doi: 10.1007/s00040-014-0362-6


Ophiocordyceps sp. on a Camponotus in Belize.

3. The famous ant-killing Ophiocordyceps fungus, when injected into a non-host species, fails to induce the stereotyped death-bite behavior. From the abstract of a paper by de Bekker et al:

…brain manipulation is species-specific seemingly because the fungus produces a specific array of compounds as a reaction to the presence of the host brain it has evolved to manipulate.

The real news here is the development of a technique to infect the fungus across species and monitor the results. This method will be powerful going forward, especially since the test of specificity in this study is weaker than the title would suggest. The researchers only injected a single non-host Camponotus, and they’ll need more for a proper assay of host range. I presume such work is coming, as the Hughes lab is generally thorough and highly productive.

source: de Bekker, C., Quevillon, L., Smith, P.B., Fleming, K., Ghosh, D., Patterson, A.D., Hughes, D.P. 2014. Species-specific ant brain manipulation by a specialized fungal parasite. BMC Evolutionary Biology 2014, 14:166  doi:10.1186/s12862-014-0166-3


Mycocepurus castrator (photo by Christian Rabeling)

4. A parasite of fungus-growing ants provides convincing evidence for sympatric speciation. Older generations of biologists have been slow to accept that new species can arise while in physical proximity to their parent species, as mating between forms could erase any nascent differences. But Christian Rabeling et al have genetic data in Mycocepurus goeldii and its parasite M. castrator (ouch!) that make any other scenario highly unlikely. From the abstract:

Based on differing patterns of relationship in mitochondrial and individual nuclear genes, we conclude that host and parasite occupy a temporal window in which lineage sorting has taken place in the mitochondrial genes but not yet in the nuclear alleles. We infer that the host originated first and that the parasite originated subsequently from a subset of the host species’ populations, providing empirical support for the hypothesis that inquiline parasites can evolve reproductive isolation while living sympatrically with their hosts.

source: Rabeling, C. Schultz, T.R., Pierce, N.E., Bacci, M. 2014. A Social Parasite Evolved Reproductive Isolation from Its Fungus-Growing Ant Host in Sympatry. Current Biology. doi:

What do Tatuidris armadillo ants eat?


Check this out:

It’s the first video of live Tatuidris, among the rarest and least understood of all ants. Until recently, no one had even seen one alive. The video is from a new paper in Insect Science, where a team of Belgian myrmecologists report their observations on a recent collection of live specimens. Here’s the abstract:

Ants of the genus Tatuidris Brown and Kempf (Formicidae: Agroecomyrmecinae) generally occur at low abundances in forests of Central and South America. Their morphological peculiarities, such as mandibular brushes, are presumably linked with specialized predatory habits. Our aims were to (1) assess the Tatuidris abundance in an evergreen premontane forest of Ecuador; (2) detail morphological characteristics and feeding behavior of Tatuidris; and (3) define the position of Tatuidris in the food web. A total of 465 litter samples were collected. For the first time, liveTatuidris individuals were observed. Various potential food sources were offered to them. A nitrogen stable isotope ratio analysis (15N/14N) was conducted on Tatuidris tatusia, other ants, and common organisms from the leaf-litter mesofauna. We found a relatively high abundance of T. tatusia in the site. Live individuals did not feed on any of the food sources offered, as usually observed with diet specialist ants. The isotope analysis revealed that T. tatusia is one of the top predators of the leaf-litter food web.

So Tatuidris is a top micro-predator. But of what?

sources:  Jacquemin J, Delsinne T, Maraun M, Leponce M. 2014. Trophic ecology of the armadillo ant, Tatuidris tatusia, assessed by stable isotopes and behavioral observations. Journal of Insect Science 14(108). Available online:

Greibenow, Z. 2014. Glimpsing Armadillo Ants. Gentle Centipede blog:

A Myrmicine Phylogeny Shakes Things Up

Monomorium kiliani

Monomorium kiliani, an Australian myrmicine. The narrow, two-segmented waist is characteristic of this subfamily.

We’re only halfway through the year, but already 2014 will be remembered as pivotal for studies of ant evolution and classification. Following right on the heels of Schmidt & Shattuck’s massive ponerine revision comes an important new study from the Ant Tree of Life group. Ward, Brady, Fisher, and Schultz (2014) have reconstructed the first thorough genus-level phylogeny of the great ant subfamily Myrmicinae.

How important is this study?

Roughly half of all ants are myrmicines, both in abundance and in species diversity. Their numbers include fire ants, harvester ants, leafcutter ants, big-headed ants, acrobat ants, and so on, to the tune of some 6,000+ species.

So… Boom! Suddenly, we’ve been given a detailed picture of the evolution of half the ants. This is big. It is so big I cannot cover the paper in detail. Instead, I’ll just give a few preliminary thoughts, as follows:

1. This is a well executed study, as we’ve come to expect from the Ant Tree of Life team, applying a thorough analysis to over 250 carefully selected taxa and 11 genes. It’s also a shining example of an older generation of genetic techniques, alas, and while I am confident the stronger results will mostly endure, be aware that an incoming next-gen tide of full genomes, and the 6,000 yet-unsampled myrmicine species, may yet overturn some of the findings.

2. The deep history of Myrmicinae, starting 100 million or so years ago, mostly occurred on those continents that drifted to become the Americas. Echos of these earliest divisions are heard in six clear, genetically distinct groups that Ward et al have formally set the up as a new system of tribes, replacing an earlier, messier scheme. The six groups are listed here in their order of divergence: Myrmicini (MyrmicaManica), Pogonomyrmecini (Hylomyrma & Pogonomyrmex), Stenammini (AphaenogasterMessorStenamma, and relatives), and three sprawling groups with thousands of species: Solenopsidini, Attini, and Crematogastrini. 


The myrmicine big picture. (Sharpie on office paper, 2014, limited edition print available, unless I recycle it).

3. The news is not all good. The clarity deep in Ward et al‘s tree fades for slightly younger events. Early relationships within some of the the six tribes are discouragingly ambiguous. This study has resolved some problems, myrmicine taxonomists face a difficult road ahead. Many of the world’s greatest genera do not form natural groups and will have to be redone. These include Aphaenogaster, Pheidole, Tetramorium, and especially Monomorium, which splatters almost comically across the Solenopsidines.

What, really, is Monomorium? Modified from Figure 1 of Ward et al (2014).

Distressingly, fuzzy resolution in a data set with this many markers and taxa means achieving proper resolution, if at all, will likely be expensive. Myrmicines may have speciated so explosively that we may never be able to reconstruct what happened with confidence.

4. The authors correct a few of the more obvious instances of paraphyly. Notably, the New World “Messor“, being unrelated to their old world doppelgangers, were moved to a revived Veromessor, and several social parasites like Protomognathus and Anergates have been sunk into the host genera from whence they evolved: Temnothorax and Tetramorium, respectively. There are other changes, too; they are listed in the abstract

Most of the identified problems- such as what to do with Monomorium and Aphaenogaster were left for targeted future research.

5. Remember the dispute over Pyramica vs. Strumigenys? The argument was fundamentally over how ant mandibles evolve. Apparently, high energy trap-jaws arise easier than anyone imagined. According to Ward  et al, not only is the assemblage of trap-jaw ants formerly included in dacetini a polyphyletic splatter, even within the genus Strumigenys the trap jaw has arisen at least twice.


A phylogram of Strumigenys, modified from Figure 1 in Ward et al 2014, showing strong support for the parallel evolution of trap-jaws in the genus.

6. The rare and bizarre African myrmicine genus Ankylomyrma is not a myrmicine at all! Rather, Ward et al‘s results unambiguously tie it to the equally bizarre Tatuidris of the Neotropics, sitting on a distant branch of the ant tree. Peas in a poneromorph pod…

Ultimately, Ward et al have crafted a sobering view of how little we still know about ant evolution, and how much remains to be done.


Aphaenogaster fulva, photographed in Illinois.

source: Ward PS, Brady SG, Fisher BL, Schultz TR (2014) The evolution of myrmicine ants: phylogeny and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae). Systematic Entomology, online early. DOI: 10.1111/syen.12090

disclosure: I received my Ph.D. from Phil Ward’s lab where much of this study was completed, and I contributed a few of the samples, but I was long gone by the time the study was initiated and have had no other involvement with the research.