The evolution, ecology and conservation of honeyguide–human mutualism

This project focuses on a unique mutualism: the foraging partnership between Greater Honeyguides Indicator indicator and human honey-hunters whom they guide to bees’ nests. Honeyguides know where bees’ nests are located and like to eat beeswax; humans know how to subdue the bees using fire, and open nests using axes. By working together, the two species can overcome the bees’ defences, with benefits to both. Remarkably, this relationship has evolved through natural selection, and provides a wonderful opportunity to study the ecology and evolution of mutualisms in nature, because human and honeyguide populations vary strikingly in how they interact, and we can readily manipulate these interactions.

Claire Spottiswoode and her team at the Fitz and the University of Cambridge have been studying human–honeyguide interactions in the Niassa National Reserve of northern Mozambique since 2013, collaborating with the honey-hunting community of Mbamba village, and receiving crucial support from the Mariri Environmental Centre led by Dr Colleen Begg, Keith Begg and Agostinho Jorge of the Niassa Carnivore Project. One key focus has been investigating reciprocal communication between the two parties: not only do honeyguides signal to humans, but in many different cultures, humans signal back to honeyguides, giving special calls to attract honeyguides and maintain their attention while following them. The Yao honey-hunters of northern Mozambique give a loud trill followed by a grunt. A 2016 experiment showed that honeyguides were twice as likely to initiate a cooperative interaction with humans who made this sound compared to humans giving control sounds, and three times as likely to lead such humans to honey.

Supported in the recent past by a Consolidator Grant from the European Research Council and currently the Max Planck–UCT Centre for Behaviour and Coevolution, we now ask  whether learning is involved in maintaining a geographical mosaic of honeyguide adaptation to local human cultures; how such reciprocal communication between humans and honeyguides mediates their interactions; what the effects of cultural co-extinctions may be on each partner and their ecosystems; and ultimately, how quickly such cultures can be re-ignited following their loss. In so doing we hope to test whether reciprocal learning can give rise to matching cultural traits between interacting species. Understanding the role of such phenotypic plasticity is important to explaining how and why the outcome of species interactions varies in space and time, and to predicting how they will respond to a rapidly changing world.

Our project, known as ‘Projecto Sego’ (‘sego’ is Greater Honeyguide in the Yao language), has the support of the community and traditional chiefs of the Mbamba and Nkuti villages. We cooperate closely with Mbamba’s honey-hunting community to carry out field sampling, experiments and documentation of Indigenous knowledge. We also regularly carry out honeyguide fieldwork in several parts of Tanzania, again in collaboration with local honey-hunting communities, and at field sites in Zambia and South Africa. Since 2022, we have been documenting honey-hunting cultures in over ten countries as part of a pan-African collaborative effort, the Honey-hunter Research Network, led by Honorary Research Affiliate, Jessica van der Wal (postdoctoral fellow 2019-2024), primarily funded by a Cultural Evolution Society Transformation Grant, and now by the Max Planck–UCT Centre for Behaviour and Coevolution. 

Activities in 2025 

•    Lailat Guta continued her MSc work. She made a successful field trip to Mbamba together with Jessica van der Wal to conduct interviews with our honey-hunter collaborators and others in the community about how people, bees and plants rely on one another (often via honeyguides). Lailat is supervised by Claire Spottiswoode, Dr Colleen Seymour from SANBI, and Jessica van der Wal. Unfortunately, the further fieldwork the honeyguide team had planned in 2025 in the Niassa Special Reserve had to be postponed due to a very difficult period of insecurity for Niassa’s people, but we plan to return in mid-2026. 

•    Even with the insecurity at the Niassa Special Reserve in Mozambique, a team of five honey-hunters continued to collect bee samples from the bee colonies they naturally harvest, resulting in a bee genetics dataset which allows us to tackle questions on the ecology of wild honeybees and how this may be affected by honeyguide–human mutualism. 

•    Eliupendo Alaitetei Laltaika finished his PhD fieldwork in Tanzania and will return to the Fitz in early 2026 to write up his thesis.

•    Prudence Tegueu carried out a successful three months of fieldwork in southeastern Cameroon for her Conservation Biology MSc project, studying the associations between Baka honey-hunters and several forest honeyguide species in the Congo Basin rainforest. In December she was joined in the field by Claire Spottiswoode and Jess Lund. Prudence is supervised by Claire Spottiswoode and Jessica van der Wal, together with Dr Mazi Sanda from the University of Ngaoundéré, Cameroon, and Dr Edmond Dounias from the French National Research Institute for Sustainable Development. 

•    Daniella Mhangwana continued her MSc research project investigating why and how several other bird species besides honeyguides also eat wax. Daniella is supervised by Claire Spottiswoode, Dr Celiwe Ngcamphalala and Dr Susan Miller. 

•    Dr Julia Cramer from the Max Planck Institute for Biological Intelligence visited from Germany to conduct sweet-taste test experiments in a variety of birds, with support from our team. These birds included the Lesser Honeyguide Indicator minor, and the data are helping us to understand the evolution of the unusual diet preferences of honeyguides. 

•    Dr David Lloyd-Jones and Claire Spottiswoode started a new collaboration with Prof. L. Mahadevan and Dr Golnar Gharooni Fard at Harvard University on modelling honeyguide–human communication. 

•    Vhuawelo Simba joined the Honey-hunter Research Network as a collaborator and conducted research on honey-hunting with honeyguides in Venda-Limpopo in northeastern South Africa. She visited the Fitz in September to present her findings. 

•    David Lloyd-Jones gave a seminar at the Fitz summarising the diverse findings from his PhD on honeyguide–human mutualism, carried out in close collaboration with the honey-hunting community of Mbamba village, Mozambique. 

•    Jessica van der Wal delivered a guest lecture for an undergraduate evolutionary anthropology course at the University of Amsterdam on the mosaic of mutualism between humans and honeyguides, including ongoing research in the Honey-hunter Research Network. 

•    Claire Spottiswoode gave departmental seminars as a representative of the Honeyguide Research Project at the University of Helsinki, Finland; the University of Mainz, Germany; the University of the Witwatersrand, South Africa; and Ludwig Maximilians University, Munich, Germany. 

•    Our citizen science project, Honeyguiding.me, continues to receive records of Greater Honeyguides, which will enable us to map the extent of changes in guiding behaviour and help to shed light on how honeyguides acquire their ability to engage with humans. Please share your Greater Honeyguides records too – we’d love to have them! (See Honeyguiding.me.) 

•    Jessica van der Wal met in person in Nairobi with collaborator Dr Hussein Isack, the pioneer in research on honeyguide–human mutualism, and they were joined online by Claire Spottiswoode and David Lloyd-Jones for a day of inspiring conversations on the past, present and future of research on this unique interspecies relationship. 

•    Jessica van der Wal continues as a Research Fellow at both the Fitz and the Max Planck Institute for Biological Intelligence, funded by the recently established Max Planck–UCT Centre for Behaviour and Coevolution. 

•    Jessica van der Wal coauthored a large collaborative study published in BMC Biology titled ‘Same data, different analysts: variation in effect sizes due to analytical decisions in ecology and evolutionary biology’ examining how different researchers analyse the same data in different ways.

Highlights 

•    David Lloyd-Jones graduated with his PhD in September with warm feedback from his international examiners. His thesis was entitled ‘Cooperation, ecology and behaviour in the honeyguide-human mutualism’ and was based on collaborative research with Claire Spottiswoode and colleagues, including a large team of honey-hunter collaborators in Mozambique. David continues his honeyguide research at the Fitz as a postdoc, based in Tanzania. 

•    David Lloyd-Jones, Claire Spottiswoode and honey-hunting colleague Musaji Muamedi have published a new study in Ecology and Evolution presenting evidence that honeyguides do occasionally (but only rarely) guide humans to animals other than bees and testing various hypotheses for why they do so. They were delighted to publish on a topic which has fascinated people for centuries, goes hand-in-hand with stories about honeyguides, and corroborates centuries of reports from a wide variety of human cultures across Africa. 

•    Jessica van der Wal and colleagues published a paper in Proceedings of the Royal Society of London B paper on recreational honey-hunting with honeyguides in Eswatini – how it works and why it still thrives. The team, led by Swati researcher Sanele Nhlabatsi and coauthored by Celiwe Ngcamphalala and Sylvester Gcina Dlamini, found that human–honeyguide mutualism is widespread across the country, especially among young cattle-herders who team up with the Greater Honeyguide to find wild honey as a sweet snack. Even though the number of honey-hunters has declined due to jobs, schooling, and habitat loss, the tradition continues, sustained by culture and community rather than economic gain, as opposed to places like Niassa Special Reserve, where honey-hunting is driven largely by its economic value. 

•    Former CB student Wiro-Bless Kamboe published a paper together with Jessica van der Wal, Claire Spottiswoode and Fitz graduate Dr Timothy Khan Aikins of the University for Development Studies, Ghana, published in Philosophical Transactions of the Royal Society B. The study documents honey-hunting practices in northern Ghana and explores the involvement of Greater Honeyguides. We found that while mutualism persists, it occurs at lower levels than those documented in eastern and southern Africa. Honey-hunters in Ghana often visit known bees’ nests without honeyguides’ help, and discarded beeswax continues to supplement the birds’ diet. We found no clear evidence that socioeconomic changes, such as increased access to motorised transport, have disrupted this relationship. This paper appears in a theme issue titled ‘Transforming cultural evolution research and its application to global futures’. The theme issue features work supported by the Cultural Evolution Society Transformation Fund, awarded to Jessica in 2022, and also includes an opinion piece coauthored by Wiro-Bless and Jessica on advancing equity in collaborative research. 

•    Claire gave an inaugural lecture on honeyguide–human mutualism at the start of her appointment as Honorary Professor of African Evolutionary Ecology at the University of Cambridge. Part of Claire’s job in this role is to help act as a bridge between us at the Fitz and UCT and colleagues in the Department of Zoology in Cambridge and the Cambridge Conservation Initiative. 

Honey-hunting Research Network 

•    Jessica van der Wal continued coordinating efforts by the Honey-hunting Research Network, supported by the Max Planck–UCT Centre for Behaviour and Coevolution. The Network is documenting the remaining diversity of Africa’s endangered cultures of honey-hunting with honeyguide birds. 

•    All network researchers (Anap Afan, George Malembo, Sanele Nhlabatsi, Wiro-Bless Kamboe, Faroukou Wabi, David Garakva, Rochelle Mphetlhe, Ali Langa, Samson Zelleke, Vhuawelo Simba and Prudence Tegueu) have now finalised their data collection and entry, and the data are currently being merged. 

•    Wiro-Bless Kamboe’s study is now published in Philosophical Transactions of the Royal Society B: Biological Sciences, and is titled ‘Honey-hunting with honeyguides in northern Ghana: cultural continuity amid change’. 

•    Sanele Nhlabatsi’s study is now published in Proceedings of the Royal Society B: Biological Sciences and is titled ‘Recreational honey-hunting with honeyguides in the Kingdom of Eswatini’.

Impact of the project 

This project closely involves collaboration with honey-hunting communities and simultaneously relies on and showcases their knowledge and expertise. Data from the project on the ecological role of honey-hunting are currently contributing to conservation planning in the Niassa Special Reserve, Mozambique. More broadly, we hope to further our understanding of how mutualisms evolve, and specifically how learnt traits mediating mutualisms may coevolve. Understanding the evolution of mutualisms sheds light on the mechanisms that can maintain cooperation among unrelated individuals. It is also important for effective conservation because mutualisms can have a wide reach in ecological communities. The honeyguide–human mutualism has disappeared from large parts of Africa as the continent changes. It would be a tragedy if it vanished altogether before we fully understood this part of our own evolutionary history. 

Key co-supporters
European Research Council; Max Planck–UCT Centre for Behaviour and Coevolution; Cultural Evolution Society; National Geographic Society; Max Planck Institute for Evolutionary Anthropology; British Ecological Society; Association for the Study of Animal Behaviour; American Ornithological Society.

Research team 2025
Team leaders and collaborators:
Prof. Claire Spottiswoode (FIAO, UCT / U. Cambridge)
Dr Jessica van der Wal (FIAO, UCT / Max Planck Institute for Biological Intelligence)
Dr Susan Miller (FIAO, UCT)
Dr David Lloyd-Jones (FIAO, UCT)
Farisayi Dakwa (FIAO, UCT)
Dr Celiwe Ngcamphalala (Biological Sciences, UCT)
Assoc. Prof. Brian Wood (U. California, Los Angeles)
Prof. Sally Archibald (Wits University)
Agostinho Jorge (Niassa Carnivore Project)
Dr Colleen Begg (Niassa Carnivore Project)
Keith Begg (Niassa Carnivore Project)
Celestino Dauda (Niassa Carnivore Project)
Dr Yusuf Abdullahi Ahmed (U. Pretoria)
Prof. Robin Crewe (U. Pretoria)
Prof. Christian Pirk (U. Pretoria)
Prof. Robert Fleischer (Smithsonian Institution)
Dr Anne Kandler (Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany)
Dr Laurel Fogarty (Max Planck Institute for Anthropology, Leipzig, Germany)
Dr Julia Cramer (Max Planck Institute for Biological Intelligence)
Dr Eunbi Kwon (Max Planck Institute for Biological Intelligence)
Dr Mazi Sanda (U. Ngaoundéré, Cameroon)
Dr Edmond Dounias (French National Research Institute for Sustainable Development)
Dr Timothy Khan Aikins (University for Development Studies, Ghana)
Dr Rodrigue Idohou (Université Nationale d'Agriculture, Benin)
Prof. L. Mahadevan (Harvard University, USA)
Dr Golnar Gharooni Fard (Harvard University, USA)
Dr Hussein Isack (Kivulini Trust, Kenya)

Honey-hunting research network 2025:
Anap Afan (APLORI, Nigeria); George Malembo (Mzuzu University, Malawi); Sanele Nhlabatsi (Eswatini); Wiro-Bless Kamboe (UCT); Faroukou Wabi (Benin); David Garakva (U. Ngaoundéré, Cameroon); Rochelle Mphetlhe (UCT); Ali Langa (Chad); Samson Zelleke (Ethiopia); Vhuawelo Simba (Rhodes University, South Africa), Prudence Tegueu (UCT, Cameroon)

Students: 
Eliupendo Alaitetei Laltaika (PhD, UCT); David Lloyd-Jones (PhD, UCT); Jess Lund (PhD, U. Cambridge); Prudence Tegueu (CB MSc, UCT); Daniella Mhangwana (MSc, UCT); Lailat Guta (MSc, UCT)

Project Sego data collection team:
Fátima Balasani, Carvalho Issa Nanguar, Kambunga Jaime, Seliano Alberto Rucunua and Rui Francisco Ndala, and many others