29/01/2022
Currently reading:https://www.jstor.org/stable/pdf/1929134.pdf
An article going quite in depth into the relationship between ant and fungus.
Just a heads-up - now we're gonna get speculative here again, and some of what I say might not be nailed to the ground as absolute truth. Maybe in the future it will be. Who knows? I am just trying to put together a picture here, so bear with me.
The common level of knowledge about ant-fungus mutualism, is, I think, mainly regarding the most specialised genera - the New World leafcutters in tribe Attini.
At the nuptial flight, the queen alate brings a pellet of fungus mass and spores with her, in a small cavity in her mouth that basically fills the function of the cheeks of a hamster. On ants this is called an 'infrabuccal cavity'. Ants can't eat solids, but they can store a little bit in this cavity.
When she founds her colony, she will nurture the fungus and grow it from this single pellet, and fertilise the substrate and feed the fungus with her own excrement and even some eggs.
When the first workers appear, they in turn begin feeding the fungus grass clippings - and the ants in turn eat the fungus. It's their only source of food and they are completely dependent on their relationship with this fungus. If it dies, the colony dies too. The fungus, in turn, is equally reliant on the ant.
This is what is usually featured in documentaries. These are the fungus growers we are familiar with from Planet Earth or similar shows.
However, the ant-fungus mutualism hole seems to go deeper. Less advanced Attini ants grow slightly different fungus, and they grow it slightly differently.
Once again - just like in the case of ant parasitism - ant species are so diverse and niched that for every step in the evolution toward a highly specialised niche - like that of the leaf cutters - there are little side tracks to genera that haven't travelled the entire way - at least not yet. And so we can try to put together an evolutionary history from the little pit-stops on the way.
Atta and Acromyrmex are the "highest" level of fungus-gardener. They feed their fungus fresh clippings from living plants. Living plants are protected by toxins, and both these "highest" genera - and their fungi - are adapted to handle this extra load of toxins, unlike their "lower" cousins.
Right underneath the leaf cutters, we have collectors of dead vegetation: genera like Trachymyrmex and Sericomyrmex. They don't cut live plants, but collect their plant matter from the ground. Fruit, flowers, seeds, leaves. They lack the specific mutation that lets them handle living plant matter.
Among these species, by the way, there is a very promising candidate for the ant-keeping hobby - Trachymyrmex septentrionalis. They hibernate, and the fungus does too - and they appear in locations as far north as New York.
These would probably be far easier to keep in the northern hemisphere than any of the tropical leaf cutters. And they are easier to feed too. The only drawback is that you don't get the polymorphism of higher Attini. Workers are pretty much all the same shape and size. The specialisation has not gone as far. Extreme polymorphism signifies an extremely niched lifestyle.
The types mentioned so far all count as "higher" genera.
Beneath these collectors of dead plant matter, there are many species that grow their fungus on excrement, invertebrate remains, and other detritus - exactly like the leaf cutter queen does right at the beginning of her colony's life.
An example of a genus with a way of life fitting this category is Cyphomyrmex - another genus in Attini. In their case, the substrate for the fungus is pretty much what you normally find on a typical ant thrash heap, and some of the species are yeast farmers. Simpler ants, in smaller colonies, farming simpler fungi.
Here we might be getting close to a possible origin story. At some point, many millions of years ago, some ants might have discovered that the stuff growing in their thrash was actually viable food, and started eating it. Eventually they started to maintain and grow it, leaving other food sources behind.
Fast forward to today and we have obligate symbiosis of ant and fungus.
This origin is speculative, of course. Just that disclaimer again. I think it makes very much sense though.
So far, all of these species belong to tribe Attini, and exist exclusively in the New World. And all of them are, compared to other ants, very, very specialised.
But there are other associations between ants and fungi, also in the Old World. These associations are more primitive, often speculative or hypothesised, and from what is known, there are no obligate fungus farmers in the entirety of Europe, Africa or Asia.
Some examples of primitive associations with fungi are very interesting, though, and might also shine a light on other possible origins of fungus farming - both in the present and in the future.
One such example of possible "early" ant-fungus association is Lasius fuliginosus. I have read that Lagerheim believed this species cultivated fungus in the walls of their nest, and as a food source. But I have also read elsewhere, later, that this has been somehow disproven.
But in the study I linked above, they had actually looked into the contents of post-nuptial flight queens' infrabuccal cavities, and found that the pellets they kept there largely consisted of spores and fungal hyphæ.
If this species had no symbiosis with a fungus, why would this be the case? How would the pellets consistently contain large amount of hyphae and spores if this was not available in the nest? And why would the queens bring it with them if it wasn't somehow beneficial or necessary?
Is this a possible pathway into obligate fungus farming? Will L. fuliginosus have a future as a fungus agriculturalist? Is the lack of this information what has caused me to fail in starting a colony of this species? Maybe they are easier to start in natural substrates, than they are in a test tube setup?
Another interesting observation can be made in genus Messor - specialised seed collectors. There have been studies where people tried to figure out how these species stopped mold from infecting their granaries, and it appears that they have built in, chemical means of handling this issue - Anti-bacterial, and anti-fungal substances - and also substances that stop seeds from germinating.
Leaf cutters in the New World also have such features, and use similar substances to remove contaminants, bacteria and competing fungus from their gardens.
If a seed collecting species forms an alliance with a fungus that can live on pre-chewed, disinfected seed mass, then that would also constitute a possible pathway into fungus gardening.
I can imagine at least these three paths into this kind of specialised lifestyle. I think the jury is still out on which one actually took place in the Attini ants.
The hypothesis I have heard talked about the most is the pathway through seed collection into fungus farming. But unless something very interesting has happened in myrmecology lately, that I have completely missed, this question is still unsettled.
Interesting stuff nonetheless, and this is no doubt a fun read!
Irving W. Bailey, Some Relations between Ants and Fungi, Ecology, Vol. 1, No. 3 (Jul., 1920), pp. 174-189