The ecology and evolution of intergroup conflict in animal societies: theory and tests
Lead Research Organisation:
UNIVERSITY OF EXETER
Department Name: Biosciences
Abstract
The history of life on earth has been dominated by a series of cooperative transitions in biological organisation, in which lower-level entities have banded together to form higher-level cooperative groups. The evolution of cooperation creates opportunities for productive coordination and teamwork within groups, but also the potential for destructive conflict and violence between groups. Humans, for example, are capable of astonishing feats of altruism and cooperation, but are at once among the most murderous and violent animals we know. In other animals there is great variation in the intensity and form of intergroup conflict, but we have a very poor understanding of the causes of this variation. The link between intergroup conflict and cooperation is also unclear: in some species intergroup conflict leads to greater social cohesion, whereas in other species it leads to greater within-group aggression.
Our project combines new theory and empirical testing to explain these conflicting observations and to advance conceptual understanding of the biological roots of intergroup conflict. Our theoretical models will predict the ecological and social factors that promote or inhibit intergroup aggression, and how individuals and groups should respond to changes in levels of intergroup conflict. We will carry out the first tests of these models on an ideally suited wild mammal system, the banded mongoose Mungos mungo, which we have studied for 20 years at site in Uganda. Banded mongooses live in highly cooperative, territorial groups which engage in frequent intergroup fights. Intergroup aggression is more severe in banded mongooses than in meerkats, chimpanzees, or any other non-human mammal. Yet there is also great individual and group variation in levels of intergroup aggression, providing an opportunity to test our theoretical predictions using experiments and long-term observations.
The research will provide a better understanding of patterns of competition in nature, and clarify the link between intergroup conflict and cooperation. We will leave a testable theoretical framework that can be applied to a very wide range of species, from microbes to humans. The project will be of much interdisciplinary interest, with the potential to influence future research in evolution, psychology, economics and political science.
Our project combines new theory and empirical testing to explain these conflicting observations and to advance conceptual understanding of the biological roots of intergroup conflict. Our theoretical models will predict the ecological and social factors that promote or inhibit intergroup aggression, and how individuals and groups should respond to changes in levels of intergroup conflict. We will carry out the first tests of these models on an ideally suited wild mammal system, the banded mongoose Mungos mungo, which we have studied for 20 years at site in Uganda. Banded mongooses live in highly cooperative, territorial groups which engage in frequent intergroup fights. Intergroup aggression is more severe in banded mongooses than in meerkats, chimpanzees, or any other non-human mammal. Yet there is also great individual and group variation in levels of intergroup aggression, providing an opportunity to test our theoretical predictions using experiments and long-term observations.
The research will provide a better understanding of patterns of competition in nature, and clarify the link between intergroup conflict and cooperation. We will leave a testable theoretical framework that can be applied to a very wide range of species, from microbes to humans. The project will be of much interdisciplinary interest, with the potential to influence future research in evolution, psychology, economics and political science.
Planned Impact
Who will benefit from this research?
A major group of beneficiaries is other academics, with whom we will engage via standard science communication activities (publishing our research in interdisciplinary and high profile journals, presenting scientific seminars, attending conferences). Our research topic has broad cross-disciplinary relevance so we will actively seek engagement with economists, psychologists, and political scientists working on intergroup conflict in human society.
In addition, the main non-academic beneficiaries from the proposed research will be 1) the general public, secondary teachers and their students; and 2) the Uganda Wildlife Authority and Uganda Tourist Board.
How will they benefit
We will contribute to an improved conceptual understanding of an issue that is of great interdisciplinary interest in academia, and great topical interest for most people. Researchers in economics, psychology and political science will benefit from a tested theoretical framework that is capable of predicting adaptive behavioural responses to changes in the ecological and social environment. While our research is largely curiosity-driven, our findings and theoretical work will have relevance and influence in these disciplines that link directly to societal policy and strategy.
The topic and research subjects are of intrinsic interest to the general public, and we aim to capitalise on this interest to improve the public understanding of science. The general public, teachers and their students will benefit through education, international cultural exchange, and engagement with scientific research.
The project teach has long-standing relations with the Uganda Wildlife Authority and Uganda Tourist board who will benefit from through the information exchange aspects of the project, and the interest in our project from international wildlife filmmakers. Our media and knowledge sharing activities will help to promote ecotourism and investment in Uganda's protected areas, and increase local and international awareness of the value of Uganda's natural resources.
Finally, the project team including the early career researchers (the RCo-I and PDRA) will benefit from developing new skills and experience with outreach and engagement with beneficiaries of the research, which will enhance career development.
A major group of beneficiaries is other academics, with whom we will engage via standard science communication activities (publishing our research in interdisciplinary and high profile journals, presenting scientific seminars, attending conferences). Our research topic has broad cross-disciplinary relevance so we will actively seek engagement with economists, psychologists, and political scientists working on intergroup conflict in human society.
In addition, the main non-academic beneficiaries from the proposed research will be 1) the general public, secondary teachers and their students; and 2) the Uganda Wildlife Authority and Uganda Tourist Board.
How will they benefit
We will contribute to an improved conceptual understanding of an issue that is of great interdisciplinary interest in academia, and great topical interest for most people. Researchers in economics, psychology and political science will benefit from a tested theoretical framework that is capable of predicting adaptive behavioural responses to changes in the ecological and social environment. While our research is largely curiosity-driven, our findings and theoretical work will have relevance and influence in these disciplines that link directly to societal policy and strategy.
The topic and research subjects are of intrinsic interest to the general public, and we aim to capitalise on this interest to improve the public understanding of science. The general public, teachers and their students will benefit through education, international cultural exchange, and engagement with scientific research.
The project teach has long-standing relations with the Uganda Wildlife Authority and Uganda Tourist board who will benefit from through the information exchange aspects of the project, and the interest in our project from international wildlife filmmakers. Our media and knowledge sharing activities will help to promote ecotourism and investment in Uganda's protected areas, and increase local and international awareness of the value of Uganda's natural resources.
Finally, the project team including the early career researchers (the RCo-I and PDRA) will benefit from developing new skills and experience with outreach and engagement with beneficiaries of the research, which will enhance career development.
Publications

Birch G
(2019)
Behavioural response of workers to repeated intergroup encounters in the harvester ant Messor barbarus
in Insectes Sociaux


Green PA
(2022)
Fighting force and experience combine to determine contest success in a warlike mammal.
in Proceedings of the National Academy of Sciences of the United States of America

Johnstone RA
(2020)
Exploitative leaders incite intergroup warfare in a social mammal.
in Proceedings of the National Academy of Sciences of the United States of America

Johnstone RA
(2019)
Evolution of menopause.
in Current biology : CB

Marshall HH
(2018)
Data collection and storage in long-term ecological and evolutionary studies: The Mongoose 2000 system.
in PloS one

Nichols HJ
(2021)
A double pedigree reveals genetic but not cultural inheritance of cooperative personalities in wild banded mongooses.
in Ecology letters

Padget RFB
(2021)
Marking through molts: An evaluation of visible implant elastomer to permanently mark individuals in a lower termite species.
in Ecology and evolution

Padget RFB
(2023)
Us, them, and the others: Testing for discrimination amongst outgroups in a single-piece nesting termite, Zootermopsis angusticollis.
in Ecology and evolution

Sheppard CE
(2018)
Intragroup competition predicts individual foraging specialisation in a group-living mammal.
in Ecology letters
Description | Intergroup conflict has been proposed as a major force promoting the evolution of altruism in animals as diverse as insects and primates. A long-standing hypothesis is that groups under attack from rivals should become more altruistic. While previous research has examined the impact of intergroup conflict on cohesion and affiliative behavior, no research to date has tested the effect of conflict on altruism, such as helping behaviour in cooperative breeders. We have carried out the first such test using long term data on wild banded mongooses, a species that features both intense intergroup conflict and conspicuous helping behaviour. Contrary to expectations, contributions to babysitting pups at the den decreased following intergroup fights, resulting in elevated pup mortality. Contributions to 'escorting', the care of offspring after they emerge form the den, remained unchanged. These different responses may reflect a fundamental difference in the personal costs of the two forms of cooperation, since babysitters are vulnerable to intergroup attack, while escorts enjoy the safety of the group. Our findings suggest that intergroup conflict may promote or inhibit altruism, depending on the nature of costs involved. We have begun to develop new theory to understand how intergroup conflict shapes the evolution of cooperation, and test the predictions of this theory using mammal and insect systems. In 2020 we published a paper showing that female banded mongooses initiate conflict, while males pay the cost. We also developed new theory showing that this decoupling of leaders from the costs that they incite, which we term 'exploitative leadership' can explain the evolution of damaging and severe intergroup conflict in this species. The findings were published in a top journal (PNAS) and received much media attention, resulting in an altmetric score of over 1200. The paper was downloaded over 10,000 times in the first 60 days since publication. |
Exploitation Route | The results show that the impact of conflict on internal group cohesion depends on the form and nature of the costs of cooperation. This finding in a non-human animal mirrors theories in psychology developed in the 19th Century, and further highlights the parallels between the study of the evolutionary origins of intergroup conflict and its role in human behaviour and evolution. Our theory and empirical work will continue to have a major impact in drawing together and synthesising findings from evolutionary biology and social sciences, with relevance to a very broad scientific audience. |
Sectors | Education Environment Financial Services and Management Consultancy Government Democracy and Justice Security and Diplomacy |
Description | The research contributed to several papers on the evolution of warfare that have received widespread media attention digitally and in the broadcast media. The banded mongoose research has been featured in over 20 wildlife TV documentaries, and we are currently negotiating with film crews interested in filming the behaviours that our research has uncovered, particularly concerning the causes and consequences of intergroup violence in this system. In addition, we have been contacted by an award winning wildlife filmmaker who wants to spend 6 months at the site to film a conservation documentary. All of these media impacts have societal benefits via education and conservation awareness, locally in Uganda and internationally. |
Sector | Digital/Communication/Information Technologies (including Software),Education |
Impact Types | Cultural Societal |
Description | Leaders of war: the evolution of collective decision-making in the face of intergroup conflict |
Amount | £647,771 (GBP) |
Funding ID | NE/S009914/1 |
Organisation | Natural Environment Research Council |
Sector | Public |
Country | United Kingdom |
Start | 03/2020 |
End | 05/2024 |
Title | Data supporting Johnstone, Cant, Cram & Thompson (2020) Exploitative leaders incite intergroup warfare in a social mammal |
Description | This data supports the following publication:
Rufus A. Johnstone, Michael A. Cant, Dominic L. Cram & Faye J. Thompson (2020) Exploitative leaders incite intergroup warfare in a social mammal. Proceedings of the National Academy of Sciences. Please read the "Read Me.txt" file for a full description of the data contained in each data set |
Type Of Material | Database/Collection of data |
Year Produced | 2020 |
Provided To Others? | Yes |
URL | https://figshare.com/articles/dataset/Data_supporting_Johnstone_Cant_Cram_Thompson_2020_Exploitative... |
Title | Early life asymmetries and the allocation of cooperative care in banded mongooses in Queen Elizabeth National Park, Uganda 2013-2016 |
Description | This is a dataset of the effect of pre-natal provisioning (supply with extra food) on offspring weight and growth, contributions to offspring care, and offspring survival in banded mongooses on the Mweya Peninsula, Queen Elizabeth National Park, Uganda, collected between 2013 and 2016. Pre-natal condition of females was experimentally manipulated by provisioning half of pregnant females in each social group during the second half of pregnancy. The other half of pregnant females did not receive any extra provisioning and acted as matched controls. Through field observations we measured female weight change during pregnancy, weight change during the pup care period following birth (the escorting period), and female contributions to escorting. We also measured pup weight at birth, weight change during the escorting period, and pup survival. Data were collected to examine how extra resources are used by females to level up inequalities in pup weight via the allocation of cooperative care. |
Type Of Material | Database/Collection of data |
Year Produced | 2021 |
Provided To Others? | Yes |
URL | https://catalogue.ceh.ac.uk/id/9c5ce831-aeda-4a59-950f-921f820d28b7 |
Title | Parentage assignments from a genetic pedigree of a wild population of banded mongooses in Queen Elizabeth National Park, Uganda, 2000-2019 |
Description | The data contains the genetic identity of parents (maternal and paternal identities and assignment probabilities) identified from DNA extracted from tail tips analysed using the MASTERBAYES program, for individual banded mongooses in a wild population on the Mweya Peninsula, Queen Elizabeth National Park, Uganda between 2000-2019. A nine generation deep genetic pedigree was constructed from which maternity and paternity assignments were calculated. This data was used to calculate lifetime reproductive success for individuals in the population who were exposed to conflict with rival groups to determine the fitness costs and benefits of intergroup conflict. In addition the type of microsatellite panel used to genotype the DNA samples is recorded. |
Type Of Material | Database/Collection of data |
Year Produced | 2022 |
Provided To Others? | Yes |
URL | https://catalogue.ceh.ac.uk/id/f397e842-b411-4256-b507-a4aa4647b914 |
Description | Collaboration on developing and testing the theory of intergroup conflict |
Organisation | University of California, Santa Cruz |
Country | United States |
Sector | Academic/University |
PI Contribution | Through our interest in intergroup conflict we have set up a collaboration with Prof Suzanne Alonzo at the University of Santa Cruz, to develop and test new theory about the role of intergroup conflict in social evolution. The theoretical predictions outlined in the grant can be tested using controlled experiments on a model insect system, the dampwood termite Zootermopsis angusticollis, which complements our field research objectives on social mammals and provides a strong test of the generality of the theoretical models we are developing. |
Collaborator Contribution | Prof Alonzo has provided advice on logistics and marking of animals, and has provided access to her laboratory facilities at the University of Santa Cruz as required. |
Impact | We are in the process of discussing and developing theoretical models. |
Start Year | 2019 |