Fighting over Food

Why do chimpanzees get into fights with their neighbors? As we report in this month’s Animal Behaviour, one of the major reasons seems to be food.

Since the 1970s, researchers have known that male chimpanzees defend group territories, and that fights between groups can be deadly. But what triggers these fights? Do chimpanzees go looking for trouble? Do they get into fights over mates? Or do they fight over some other resource, such as food? And when they do meet the neighbors, what determines whether they fight or flee? Are they more likely to respond aggressively if they are defending mates, or young infants, or food? Or does strength in numbers matter more? We sought to answer these questions, using 15 years of data on social behavior and ecology.

These data come from the Kanyawara community of chimpanzees in Kibale National Park, Uganda, where I did my PhD dissertation research as a student of Richard Wrangham and Marc Hauser. For the past decade, I’ve been doing fieldwork in Tanzania instead of Uganda, but I’ve continued to collaborate with Richard and his team. Such collaboration is made easier now that the long-term data have been entered into an Access database, and the original datasheets are all scanned, making it a simple matter to consult the original records from anywhere with good internet access.

Working with these data brought back many vivid memories, such as the day I first saw chimpanzees in the wild, back in June 1996. I had come to Kibale to spend a summer doing pilot research. We hiked for hours in the forest to find the chimpanzees, following narrow trails up and down steep hills, stepping carefully along narrow logs lain across boggy patches in the valley bottoms. Everything looked green and wet. Towering rainforest trees hung with vines. Dense stands of wild ginger and other leafy tropical plants crowded the understory.

As we crept along a steep trail, an opening in the trees showed the misty green expanse of the valley below. At last, I caught my first fleeting glimpse of a chimpanzee – I think it was Stout, an adult male – as he crossed the trail. His long black hair glistened with rain as he knuckle-walked calmly but quickly through the forest, looking more like a gorilla than I had expected, having only seen chimpanzees in movies and zoos.

During my first weeks at Kanyawara, the chimps traveled in large parties with many males. I saw most members of the community, including old Lamy, with his crippled foot, new mothers Outamba and Tongo carrying their first infants, matriarch Lope and her family, including the gangly teenager Makoku and young Rosa, grey-bearded Stocky, tough young males Imoso and Johnny, and Big Brown, who was still the alpha male.

During my first few days in the field, the chimps hunted and killed a red colobus monkey, and had a brief, noisy, and (for me) completely confusing encounter with their neighbors, during which the stranger chimps came within sight of us, crashing through the underbrush, displaying at and running towards our chimps before running they turned and ran away. As it turns out, I had come just in time for the last weeks of Uvariopsis season.

Uvariopsis congensis is a small tree that grows in large groves in the understory of the forest. It tends to fruit synchronously, usually in May and June. The fruits are small and red, and look rather like fleshy red peanuts, with two or three seeds.

At Kanyawara, the Uvariopsis groves occur mainly in the southeast of the range, along the border with another chimp community, which has not been habituated. And it turns out that that these Uvariopsis groves are a key focus of intergroup competition.

Back when I was doing my dissertation work, the number of intergroup encounters that had been observed at Kanyawara was still too small for meaningful statistical analysis. But now, in the 15-year dataset, we have records of 120 intergroup encounters. As is typical for chimpanzees, most of these encounters were shouting matches rather than full-fledged fights – chimpanzees gave loud pant-hoots and waa-barks and other calls, challenging their neighbors, and then ran either towards or away from them. But when chimpanzees came close enough to see each other, they were always hostile. Two chimpanzees were killed, and several others badly injured.  We found that these encounters happened mainly when chimpanzees visited border areas, especially the southern border.

So why did males visit border areas in the first place? To find mates, when few females in their own community were receptive? To find neighbors to attack, whenever they could gather in big enough parties to have a good chance of beating the neighbors? Or to look for food?

To answer these questions, we needed to use multivariate statistics to consider all these factors at once: mates, party composition, and food. We knew that male chimpanzees were highly motivated to search for mates, and we had previously found that parties that visited the range periphery had more males, presumably because that provided safety in numbers. But we also suspected that food would be an important part of the answer.

Chimpanzees spend most of their time searching for food, eating food, or digesting food. Though chimpanzees eat a variety of things, including monkeys, honey, ants, termites, leaves and stems, they spend most of their time eating ripe fruit from trees. In a tropical forest, the availability of tree fruits varies in a complex way over space and time. Some trees occur in groves in particular areas, whereas others are scattered about the forest. Some tree species, like figs, may produce food throughout the year, but other species fruit synchronously, with a big crop occurring every one or two years. To test how food influences chimpanzee movements, we needed data on both where and when key foods were located.

We had lots of data on feeding behavior. But it could be that chimpanzees travel to different parts of their range for social reasons (such as looking for mates or checking on the neighbors), and just happen to eat whatever they find along the way. So we needed to test whether the feeding data reflected where food was actually located. We therefore examined data from vegetation plots, and confirmed that both feeding data and vegetation plot data agreed that certain tree species were more common in the north, while others were more common in the south. For key species, we also found a good correlation between feeding behavior and independent measures of whether a sample of those trees had fruit in a given month. Additionally, the strong correlation between feeding records and counts of seeds from dung samples gave us confidence that our data on feeding behavior provided a good measure of what chimps were actually eating.

So what did we find?

Males tended to stay closer to the range center, and were less likely to travel to the south, when they were with more sexually receptive females. Party composition also mattered: as we had found previously, chimps were more likely to visit dangerous border areas when in parties with many males. And as we suspected, food had a big impact on chimpanzee movements. They visited the southern border mainly when southern fruits were in season. One species in particular had a strong effect on encounter rate: Uvariopsis. Most of the intergroup encounters took place in the southeast, in or near the Uvariopsis groves, and encounters were more likely to occur on days when chimps spent more time eating Uvariopsis.

So chimpanzees at Kanyawara visited borders for various reasons, but food seemed especially important. In particular, a good crop of Uvariopsis attracts chimps from both sides of the border, resulting in a spike in the rate of intergroup encounters during Uvariopsis season.

When encounters did occur, though, we found that the response depended mainly on the number of adult males that were present. Whether they were with mates, infants, or food didn’t matter as much as whether they had enough males to put up a good fight. These results were very much in line with what we had previously found with the playback experiments.

So is food generally the main driver of intergroup competition in chimpanzees? I’m currently working on data from Gombe to test whether similar factors apply there. It seems likely that in addition to food, the relative power of communities is a key factor. An interesting case in point is just 12 km from Kanyawara: the Ngogo community, studied by John Mitani and David Watts. This is the biggest chimpanzee community ever studied, with about 150 members. These chimps live in a range about the same size as Kanyawara’s, but in higher quality forest, with a lot more food, and three times the chimpanzees. And a lot more violence. The Ngogo chimps frequently patrol their boundaries, and in a ten-year period, Ngogo males killed 21 of their neighbors.

While the Ngogo chimps were killing their neighbors and expanding their range, the Kanyawara chimps were losing territory to their powerful southern neighbors (a group of unhabituated chimpanzees whose range appears to be sandwiched between Ngogo and Kanyawara). By 2006, the Kanyawara range was less than half as big as it had been in 1998, and the southern chimps had pushed the boundary a full kilometer north. It may be that chimps in a powerful community, like Ngogo, are more likely to go looking for trouble, whereas chimps in a weak community, like Kanyawara, visit dangerous border areas only when the abundance of food there makes the risk worthwhile.

7 thoughts on “Fighting over Food”

  1. Fascinating. I was initially shocked that the chimps in the area with more abundant food resources were actually more violent. But if a “scarcity mindset” were programmed into the species by evolutionary pressures, it could make sense that those who are fortunate enough to find themselves in an area with abundant resources would be eager to protect and expand their territory. Such a group could also be expected to thrive well enough on the abundant resources to grow strong and numerous enough to protect and expand its territory more effectively.

    I’m speculating, of course, and I certainly don’t share your scientific credentials nor your access to the data. In any case, the correlations you found are wonderfully interesting. Perhaps, if our ancestors faced similar environmental and evolutionary pressures, it might help to explain why our species is so keen on becoming violent and acquisitive with our neighbors, especially when we judge we are strong enough to do so.

  2. Glad you liked the post, Doug.

    As for scarcity — even in humans, it often seems to be those with abundant resources that most eagerly and effectively carry out war. The British Empire expanded across the glob as the Industrial Revolution made the British increasingly more wealthy and powerful than others. Industrialization also fueled imperial expansion by Germany and Japan. The relative wealth and power of the early United States compared to other powers in North America enabled the expansion of the US from the Atlantic seaboard across the entire continent to the Pacific. Like the Bible says, “For whosoever hath, to him shall be given, and he shall have more abundance: but whosoever hath not, from him shall be taken away even that he hath.” (Matthew 13:12)

  3. Mike – I’m just now catching up on blogs – glad you’re posting again. It seems to me that offensive (i.e. territory-expanding) aggression might depend on the availability of surplus resources – if you’re just scraping by on what you’ve got, it doesn’t seem sensible to squander it on a risky move to get more. But if you’re doing well, why not blow some of it on trying to get more? In the worst case, you lose some surplus, get rid of some of your more aggressive young males, and go on with business. Of course, if you’re truly desperate (i.e. what you’ve got just isn’t enough to survive), then maybe different rules apply. But perhaps by then most societies are too far gone to mount any attack at all?

  4. Mike – Great post. Came across it while searching for material for my intro bio anthro course. funny to come across it too because we actually met in the Tabora airport in TZ in 2000(?). I think both of us were heading back to Dar post field seasons. Glad to have come across this write up (and others) – excellent complement to your groups papers.


  5. Alex — yes, I think that’s a good point about offensive aggression being related to surplus resources. That certainly seems to be the case with the Ngogo chimpanzees. They live in the best part of the forest in terms of chimp food, and yet seem to be constantly seeking to expand their range.

    And of course with humans, and as I mentioned to Doug — increasing resource availability seems to be an important enabler of offensive aggression in humans. The European colonial project seems to have been fueled in large part by potatoes and maize — New World crops that increased agricultural productivity enormously, leading to population growth, and thus more potential soldiers (and more competition for opportunities at home, thereby making going overseas a more attractive option.)

  6. He also mentioned some pcsohclogiyal study that showed if two humans are looking at each other eye to eye for more than thirty seconds, they are either fighting, or thinking about fighting, (or at the very least feeling some kind of aggressive competition), or the opposite either engaged in sex, thinking about sex, or at the very least having sexual feelings. Just wondering if you could tell me more about the study. Trying to research eye contact, and not getting very far.Thanks,Jen

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