Evolution and Biodiversity

Category: foraging

Desert ant builds landmark

Nest hill helps ants to return home in barren salt flats

desert ant Cataglyphis fortis has outstanding navigation skills

Often nothing is visible around a nest of the desert ant Cataglyphis fortis that could help foraging workers to return to the nest. In that case, the ants make a landmark themselves, Marilia Freire and colleagues show.

A foraging trip is a survival trip for the desert ant Cataglyphis fortis, which lives in salt pans in Tunisia; salt pans are vast bare plains where there was once water, but now only a salt crust remains. Ant workers venture out individually to search that barren plain for insects and other small critters that have succumbed to the relentless desert heat. After founding something, they must return to the nest with the loot between their jaws as quickly as possible, otherwise they will succumb themselves.

But the entrance to the underground nest is barely visible. That is why the ants build a landmark, when necessary, Marilia Freire and colleagues discovered.


Food is scarce, so foraging desert ants often must move far from the nest to find something. They venture up to 350 meters away. Because they have excellent navigation skills, they usually return safely.

When going out foraging, a worker constantly uses an internal sun compass to keep track of the direction in which she is walking and with a kind of pedometer she measures the distance she covers in that direction. When she finds food, she has usually followed a tortuous path, but thanks to this so-called path integration she can walk back to the nest in a straight line, i.e., via the shortest possible route. At least: she closely approaches the nest.

When within a few meters, she needs visible clues to find the exact place of the nest entrance, because path integration doesn’t work perfectly. The farther an ant has gotten from the nest, the more uncertainty creeps into the route back and thus the greater the chance is that she has to search for too long and succumbs. For the very last bit of the trip homewards, she relies on nest smell.

But in the middle of a salt pan, there is nothing to be seen at all. What to do in this case?


desert ant builds nest hill when no other visual landmarks are aroundFreire and the other researchers had noticed that the desert ants often build a hill at their nest, and that nest mounds in the middle of a salt pan are higher than on the edge, where some shrubs grow. A nest mound in the middle of a salt pan is 12 centimeters high on average (the highest they found was 30 centimeters), a nest mound at the edge only 5. So, they wondered if the mounds might serve as visual landmarks for workers returning from a foraging trip.

To find the answer, they captured ants at the nest and placed them at a distance of a few meters. Since the ants had not walked themselves, they could not use path integration. But they were placed at distances where they normally must be guided by landmarks to find the nest entrance anyway. The researchers had removed the mounds at some of the nests to see if that made a difference.

That turned out to be the case, especially for nests in the middle of a salt pan. Without a mound, ants were not able to walk directly to such nest and more often failed to find it at all. The hills therefore serve as landmarks. Next question: do ants build them specifically for that purpose? It is possible that the mounds have another main function, such as regulating the nest temperature.

Only when needed

But the researchers show that the desert ant does build its mounds as landmarks by conducting experiments in which they removed the mounds at sixteen nests in the middle of a salt pan. At eight of those nests, they placed artificial landmarks, namely two black cylinders. Three days later, the ants were found to have built a new mound at some of those sixteen nests, especially at nests without artificial landmarks, and at those nests, the mounds were taller.

Conclusion: desert ants build mounds near their nest as landmarks for foraging workers. But they only make the effort if there are no other landmarks visible, such as bushes or, in the trials, black cylinders.

Willy van Strien

Photos: ©Markus Knaden
Large: Cataglyphis fortis
Small: nest mound in the centre of a salt pan

Freire, M., A. Bollig & M. Knaden, 2023. Absence of visual cues motivates desert ants to build their own landmarks. Current Biology 33: 1-4 (31 May online). Doi: 10.1016/j.cub.2023.05.019
Steck, K., B.S. Hansson & M. Knaden, 2009. Smells like home: desert ants, Cataglyphis fortis, use olfactory landmarks to pinpoint the nest. Frontiers in Zoology 6: 5. Doi: 10.1186/1742-9994-6-5
Wittlinger, M., R. Wehner & H. Wolf, 2007. The desert ant odometer: a stride integrator that accounts for stride length and walking speed. The Journal of Experimental Biology 210: 198-207. Doi: 10.1242/jeb.02657
Wehner, R., 2003. Desert ant navigation: how miniature brains solve complex tasks. Journal of Comparative Physiology A 189: 579-588. Doi: 10.1007/s00359-003-0431-1

Garden or nameplate?

Why vicuñas create communal dung piles

Vicuñas use permanent latrines to defecate and urinate

Vicuñas live in arid, cold and barren areas, high in the Andes. They set up permanent places to defecate and urinate and use those latrines for decades. There is disagreement about why.

High in the South American Andes, where the soil is arid, rocky, and barren, some places stand out because they are green, overgrown with plants. The greens islands developed because vicuñas come there repeatedly to defecate and urinate. Why do they use such latrines? To create gardens with plants that they can feed on, Kelsey Reider and Steven Schmidt suggest. No, the dung piles are kind of nameplates that mark their territory, William Franklin thinks.

Unpalatable bunch grass

Vicuñas are one of the few animal species that live in the Andes at altitudes of more than 4000 meters, right up to the edge of snow. They mainly live in groups that roam over a territory of almost 20 square kilometers. Climate change is also noticeable here; glaciers dwindle and retreat to the mountain tops. Where they melt, a bare bottom appears which is poor in plant nutrients, so that it takes decades before a noteworthy vegetation is formed. Vicuñas are the first to enter the newly exposed soil at the edge of the glaciers.

With their droppings, they enrich the soil with nutrients. They defecate and urinate only on permanent latrines or dung piles which persist for decades. Consequently, fertilized places are created where vegetation can develop more quickly.

First, a vegetation appears that is dominated by the tough and little nutritious Peruvian feather grass, Stipa pichu. It is not until hundreds of years later that a grassier vegetation develops, with the grass Calamagrostis vicunarum, other grasses and herbs.


In those grassy places vicuñas forage preferentially. Because the places are still used as latrines also, the animals run the risk of picking up gastrointestinal parasites. But places with tasty vegetation are so scarce that it is worth the risk.

That is why Reider and Kelsey believe that the vicuñas maintain latrines in order to concentrate their dung and accelerate the development of nutritious vegetation locally. In other words, latrines are gardens where they grow food.

Franklin thinks otherwise, however. Vicuñas that use a young latrine at the edge of a glacier or start a new one will not be able to enjoy a tasty yield themselves, because generations will have passed before there will grow anything edible. When it comes to food breeding, it would be better for an animal to choose an older latrine where plant growth is already substantial.

Instead, he thinks that the dung piles mark the territory of a group. This is important because if an animal enters another group’s territory accidently, it will be violently attacked and chased away and is at risk of serious injury. By marking the territory at fixed places with the characteristic group scent, especially at the borders, a group manages to keep its members within their own safe territory. So, at a border, two groups may be seen peacefully grazing side by side, each in its own area.


Every group member contributes to these scent markings, and whoever contributes benefits from the fact that the nameplate is maintained.

As a result, vegetation develops on bare ground, gradually becoming more attractive. Which is a nice side-effect for future generations and other mammals that visit the grassy places: mountain viscacha (Lagidium viscacia) and Andean fox (Lycalopex culpaeus).

Willy van Strien

Photo: Dick Culbert (Wikimedia Commons, Creative Commons CC BY 2.0)

Franklin, W., 2021. Vicuña dung gardens at the edge of the cryosphere: Comment. Ecology 102: e03522. Doi: 10.1002/ecy.3522
Reider, K.E. & S.K. Schmidt, 2021. Vicuña dung gardens at the edge of the cryosphere. Ecology 102: e03228. Doi: 10.1002/ecy.3228

Leaf cutters prevent traffic jams

Take no heavy load when traffic flow is high

leaf cutter ants carry small leaf fragments on crowded trails

When the number of workers on foraging trails is high, leaf cutters maintain the flow by carrying only small pieces of leaf with them, Mariana Pereyra and Alejandro G. Farji-Brener show. Otherwise traffic jams would arise.

The fungus that leaf cutter ants grow in their gardens needs fresh plant material continuously to grow on. And so ant workers walk up and down trails that are cleared and maintained free of debris. They leave the nest to cut leaf fragments from plants and return with a piece in their jaws.

Sometimes ants carry extra-large leaf fragments, causing them to move slowly. That is cumbersome when the trail is crowded, because then a slow ant may hinder the flow. Accordingly, when many ants are walking on the path, they only take small loads with them, Mariana Pereyra and Alejandro Farji-Brener write.

Truck-driver effect

In earlier research, Farji-Brener and colleagues had shown that workers of the leaf cutter Atta cephalotes sometimes carry a strikingly large piece of leaf, up to twice the normal size, to deliver a large gain at the nest. But such extra large burden also has disadvantages; a heavily loaded ant runs slower and hinders the ants that come behind her carrying a normal load. Their walking speed may be reduced by up to 50 per cent. So, a traffic congestion may form behind a heavily loaded worker; the researchers call it the truck-driver effect. It slows down the entire column.

A slow ant on the trail is especially obstructive when it is busy, because in that case, ants walk close together and cannot overtake a slow colleague. At high ant flows, the biologists observed relatively few ants with a heavy load. Is that because the ants are so ‘wise’ not to enter a busy path with a heavy load?

Steady flow

Pereyra and Farji-Brener now answered that question in another species, Acromyrmex crassispinus. They offered workers pieces of ‘leaf’: filter paper soaked in orange juice. They presented pieces of normal size and of extra large size and observed what choice the ants made when different numbers of ants were walking on the trail. And indeed: only at low ant flows, workers selected extra large pieces of paper; when many ants were running, they only picked up the smaller pieces.

Various reasons are thinkable for avoiding large pieces; they make it more difficult to manoeuvre in case of obstacles, the chance of collisions is greater and a heavily loaded ant is more vulnerable to predators. But the fact that ants tend to ignore the large parts at high ant flows suggests that they also do so in order not to obstruct traffic. In this way, leaf cutters optimize colony performance. All going at the same speed: on a busy path, that is the best way to keep a steady flow.

Just like on highway.

Willy van Strien

Photo: Atta cephalotes ©Alejandro Farji-Brener

Pereyra, M. & A.G. Farji-Brener, 2019. Traffic restrictions for heavy vehicles: Leaf-cutting ants avoid extra-large loads when the foraging flow is high. Behavioural Processes, online November 25. Doi: 10.1016/j.beproc.2019.104014
Farji-Brener, A.G., F.A. Chinchilla, S. Rifkin, A.M. Sánchez Cuervo, E. Triana, V. Quiroga & P. Giraldo, 2011. The ‘truck-driver’ effect in leaf-cutting ants: how individual load influences the walking speed of nest-mates. Physiological Entomology 36: 128-134. Doi: 10.1111/j.1365-3032.2010.00771.x

Pair bonds in bats

Female Egyptian fruit bat selects male that shared its food

In Egyptian fruit bat, a fruit-eating mammal, males take the initiative to mate, but females determine whether mating occurs. They strongly prefer a friend that often offered them food, Lee Harten and colleagues write.

Bats are social animals, and so is the Egyptian fruit bat (Rousettus aegyptiacus), which occurs in Africa and the Middle East. The fruit-eating mammals live in large colonies of up to thousands of specimens. Individuals within a group maintain friendship bonds with a few others, meaning that they share food.

Lee Harten and colleagues previously reported that the animals have two ways to obtain food. A risky way is to get fruit from a tree on their own. When a bat lands in a tree to collect food, it runs the risk of being caught by a predator, such as a snake or a cat. Therefore, the bats forage high in the trees. And when a fruit tree has thin foliage, they fly with their catch to a safe place to consume it.

There is also a funky method that the bats often use. If a colony mate holds a fruit in its mouth, they approach it and try to steal it. The bat that has obtained the fruit may respond aggressively, but sometimes it will have its catch scrounged.


Individuals differ in their strategy. Some usually pick their own fruit, while others are more likely to try to scrounge it. The scroungers are more anxious. They are afraid to land on a place with food, and if they do, they are so vigilant that most times, they will not be able to pick any fruit. For faint-hearted bats, scrounging from others is the better option.

Often scroungers don’t approach any arbitrary colony member, but they have one or two partners that they regularly approach, and that tolerate it. So, a network of affiliations exists.

Overall, Egyptian fruit bat males and females use different strategies. Males are more likely to collect fruit on their own than to scrounge, while for females it is the other way around. Only during lactation – a female produces one pup once or twice a year – they shift to collecting food on their own; they then need extra energy. Outside that period, they prefer to scrounge, each from its own set of favorite males.


Now, Harten shows that those relationships have big consequences. In his lab, he kept a colony of wild-born Egyptian fruit bats, fifteen males, ten females and the young that were born in the lab. Genetic paternity analysis of the pups showed that in most cases, the father was one of the males that the mother preferred to get food from. The transfer of food from father to mother had been most intensive in the period just before pregnancy.

It is not a direct exchange of food for sex, because not all food-sharing bonds result in a descendant. But by tolerating a few females to prig food, a male has a chance to sire offspring later. Although a male takes the initiative to mate, a female decides whether or not to accept it. If she does, the male gets something in return for its generosity. Such delayed reciprocity is probably an explanation, but maybe not the only one, that the animals share food with some others.

Each male has a number of regular scroungers and a chance to produce a young with one of them. The relationships persist during a breeding season, but when a new period starts, females select another male to sire their young.

Willy van Strien

Photo: Egyptian fruit bat with fig. Artemy Voikhansky (Wikimedia Commons, Creative Commons CC BY-SA 3.0)

Harten, L., Y. Prat, S.B. Cohen, R. Dor & Y. Yovel, 2019. Food for sex in bats revealed as producer males reproduce with scrounging females. Current Biology, online May 23. Doi: 10.1016/j.cub.2019.04.066
Harten, L., Y. Matalon, N. Galli, H. Navon, R. Dor & Y. Yovel, 2018. Persistent producer-scrounger relationships in bats. Science Advances 4: e1603293. Doi: 10.1126/sciadv.1603293

Mode of transport

Ants select best carrier material for collecting fluid food

Ant Aphaenogaster subterranea uses tools to collect fluid food

The ant Aphaenogaster subterranea uses absorbent material to carry fluid food back to its nest. It selects the most easily portable material, or it takes material that is discovered first, as Gábor Lörinczi and colleagues observed.

Ants of many species possess a greatly distensible crop and a highly modified proventriculus, in which they transport large volumes of liquid food back home for their nest mates. But other species, among which Aphaenogaster subterranea, don’t possess such internal carrier sac. Still, they are able to collect fluid food. They drop debris in such food source, and grasp the food-soaked material with their mandibles to take it with them. Their choice of carrier material is flexible, Gábor Lörinczi and colleagues write.

Aphaenogaster subterranea occurs in forests in Central and Southern Europe. It lives in colonies, mostly in a nest in the soil, under a stone. Specialised workers collect fluid food for the colony: fruit pulp and body fluids of dead arthropods.

Aphaenogaster is flexible in its choice of toolsLörinczi held a number of nests in plastic boxes the laboratory and conducted experiments in which the nests were connected to a foraging arena via a plastic tube. In the foraging arena, he offered a drop of honey diluted in water or honey enriched with sugars as a liquid food source on a plastic disc; or he offered pure water as a control. Around the disc, he placed different piles of carrier material: small soil grains (1 millimetre in diameter), large soil grains (2 millimetre in diameter), pieces of pine needles (5 millimetre in length), pieces of plant leaves (5 millimetre in length) and pieces of sponge (5 millimetre in length). He then observed the foraging behaviour of the ants.


If all piles of ‘carrier bags’ were close to the food bait (at a distance of 4 centimetres), the ants selected mainly small soil grains to transport to and drop into the food. These grains are most easily transportable.  If all piles were at greater distance  (12 centimetres), the ants were less selective. And if one type of material was close to the food bait while the other types were not, they used that material relatively more often. So, the choice of carrier material is not fixed. The ants prefer small soil grains, but if something else is discovered sooner or more readily accessible, they will use that, maximizing their efficiency.

Leave fragments were used only infrequently, even if it could be found close to the food bait. This material is difficult to handle.

Ants that retrieved food-soaked material from the food bait, mainly picked up small soil grains. From honey, they also collected many pieces of sponge. As the authors suggest, these may be easy to pick up because of their buoyancy. They also observed that after a while, the ants started to tear the pieces of sponge into smaller fragments before using them.

Into water, objects were dropped infrequently, and no objects were retrieved from it.

So, workers of Aphaenogaster subterranea show flexibility in foraging tool use, and they even modify some material, which is unique among insects as far as is known.

Willy van Strien

Large: Aphaenogaster subterranea. Christophe Quintin (via Flickr, cropped; Creative Commons CC BY-NC 2.0)
Small: The ants covered a drop of food with absorbent material. ©Gábor Lörinczi

Lőrinczi, G., G. Módra, O. Juhász & I. Maák, 2018. Which tools to use? Choice optimization in the tool-using ant, Aphaenogaster subterranea. Behavioral Ecology, online August 1. Doi: 10.1093/beheco/ary110
Maák, I., G. Lőrinczi, P. Le Quinquis, G. Módra, D. Bovet, J. Call & P. d’Ettore, 2017. Tool selection during foraging in two species of funnel ants. Animal Behaviour 123: 207-216. Doi: 10.1016/j.anbehav.2016.11.005

First aid

Hunting ant workers rescue lightly injured nest-mates

termite-hunting ant Megaponera analis rescues lightly-wounded nest-mates

Groups of the African ant Megaponera analis undertake hunting parties that are risky because of the fierce resistance of the termites that are attacked. Some ant workers get injured, but they are carried back to the nest and treated if possible, Erik Frank and colleagues report.

Workers of the large African ant Megaponera analis, also known as Matabele ant, face a heavy task. The ants prey on termites that they detect and overpower at their foraging sites. The ants approach the termites in a column formation consisting of hundreds of individuals. When the first ants reach a site, they wait until all participants have arrived and then they attack. The large individuals, the majors, break open the protective layer of earth that covers the termites’ foraging site; the small ants, the minors, then go inside to seize, kill, and pull out the termites.

Emergency signal

And that is a risky job, as Erik Frank and colleagues write. Termite soldiers with strong head and mandibles will fight fiercely. Some ants, almost all of them being minors, get injured; some ants are bitten off one or more legs or antennas, others are hindered by a termite that clings to them.

The ants limit losses by rescuing many injured nest-mates. After the fight, the ants gather before they jointly return to the nest, because an ant travelling alone easily falls prey to predators, for example spiders. Majors run over the place to pick up and carry dead termites and nest-mates that lag behind. If all ants have joined the column, they start walking. But ants that lost one or two legs and ants with a termite clinging to them are unable to keep up with the group, according to observations and experiments in the field and in the lab. By excreting certain substances they signal to others that they need help.

Majors that are not yet carrying anything will pick up these lightly-injured nest-mates, which tuck in their legs to facilitate transportation.

Ants who are severely injured and can no longer stand on their legs, don’t emit an emergency signal and they don’t let themselves to be picked up: they keep on twisting and turning. These unhappy ants are left behind, so that only victims that have a chance to recover are taken home. Almost all of them safely reach the nest, whereas without help many injured ants would not be able to complete the journey.


As soon as the victims are brought into the nest, they are taken care of. A termites that clings to an ant mostly is pulled off successfully and the ant doesn’t suffer any long-term consequences of the adventure. An ant that lost a leg or antenna receives a thorough treatment: nest mates groom the open wound for a long time, cleaning it and probably also applying antimicrobial substances that they produce in special glands. Experiments show that an ant with an untreated open wound almost always dies, probably due to an infection. But when treated, it usually survives and it will learn to walk on four or five legs as fast as the others – and soon enough, it will join termite raiding parties again.

When heavily injured ants are brought in, which happens only infrequently, they will get no treatment, but are carried out of the nest instead. The ants only help injured nest mates that will survive.

The rescue behaviour in Megaponera analis is unique. It could develop in these ants because they conduct short and space-limited raids on a dangerous prey. There are many casualties, but the injuries are rarely fatal when the victims get help – and help is worth the effort. Without rescuing behaviour, the colony would be much smaller and fewer workers would be available to join a raiding party. To give an idea of the importance: the number of ants that are rescued on a day roughly equals the number that is born.

Willy van Strien

Photo: Megaponera analis: major carrying injured nest mate back to the nest. ETF89 (Wikimedia Commons, Creative Commons CC BY-SA 4.0)

Videos of Megaponera ants that carry and treat injured nest mates

Frank, E.T., M. Wehrhahn & K.E. Linsenmair, 2018. Wound treatment and selective help in a termite-hunting ant. Proceedings of the Royal Society B 285: 20172457. Doi: 10.1098/rspb.2017.2457
Frank, E.T., T. Schmitt, T. Hovestadt, O. Mitesser, J. Stiegler, K.E. Linsenmair, 2017. Saving the injured: Rescue behavior in the termite-hunting ant Megaponera analis. Science Advances 3: e1602187. Doi: 10.1126/sciadv.1602187