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 and Keith Begg of the Niassa Carnivore Project. A key focus to date 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 by a Consolidator Grant from the European Research Council, 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 crucial to explain how and why the outcome of species interactions varies in space and time, and to predict 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 the local community to collect data and assist with our field sampling and experiments.

Activities in 2022

  • Fieldwork was limited in 2021 owing to COVID-19, but David Lloyd-Jones was able to make three short visits to Niassa to support our field team of honey-hunter colleagues, who continued to collect excellent data on their interactions with honeyguides
  • We hosted a two-day online workshop on “Human-Wildlife Mutualisms”, involving 40 colleagues from 14 countries, including biologists, anthropologists, historians, conservation practitioners and honey-hunters.
  • CB MSc student Eliupendo Alaitetei Laltaika completed his MSc on honeyguide-human interactions in four cultures in northern Tanzania. We welcome Laltaika as a PhD student in 2022.
  • Honours student Rion Cuthill studied the effects of honeyguide-human mutualism on fire ecology, co-supervised by Prof. Sally Archibald. Rion will continue to work with the team in 2022.
  • Honours student Cameron Blair completed his research project on the developmental origins of the Greater Honeyguide’s guiding call, co-supervised by Jessica van der Wal. Cameron will join the brood parasite coevolution team as an MSc student in 2022.
  • Our citizen science project,, managed by Jessica van der Wal, continues to receive records of Greater Honeyguides, which will enable us to map the changes in the extent of guiding behaviour and help to shed light on how honeyguides acquire their ability to engage with humans (see also our website:
  • We established a new field site at Honeywood Farm near Grootvadersbosch Forest in the Western Cape, South Africa, allowing us to address some of our honeyguide research objectives closer to base.


  • We submitted two collaborative review papers arising from our “Human-Wildlife Mutualisms” workshop, on the ecology and evolution of human-wildlife cooperation, and on its safeguarding; these are currently under review.
  • Eliupendo Laltaika (MSc), Rion Cuthill (Honours) and Cameron Blair (Honours) all received Distinctions for their research dissertations. 
  • Jessica van der Wal and Claire Spottiswoode presented their research findings at the “Interspecies Conversations” workshop, an interdisciplinary meeting involving biologists, anthropologists, computer scientists, linguists, artists and others. Claire Spottiswoode presented a plenary talk on the honeyguide team’s research at the International Congress of Zoology.
  • Jessica van der Wal collaborated with Cape Town artist Jane Solomon to produce a ‘capulana’ (a traditional Mozambican sarong) depicting a honey-hunting scene in Niassa, to provide a form of media for honey-hunters to talk to, and hopefully enthuse others in the village about their profession. We thank the Eric Hosking Trust for their support of this initiative.

Impact of the project

This project involves rural communities in understanding a unique human-animal relationship. 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 develops. 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; National Geographic Society; Max Planck Institute for Evolutionary Anthropology; DSI-NRF CoE grant; British Ecological Society; Association for the Study of Animal Behaviour; American Ornithological Society.

Research team 2022
Prof. Claire Spottiswoode (FIAO, UCT / U. Cambridge)
Dr Jessica van der Wal (FIAO, UCT)
Dr Dominic Cram (U. Cambridge)
Dr Brian Wood (U. California, Los Angeles)
Prof. Sally Archibald (University of the Witwatersrand)
Prof. Timm Hoffman (Biological Sciences, UCT)
Dr Colleen Begg (Niassa Carnivore Project)
Keith Begg (Niassa Carnivore Project)

Students: David Lloyd-Jones (MSc, UCT); Eliupendo Alaitetei Laltaika (CB MSc, UCT); Rion Cuthill (Honours, UCT); Cameron Blair (Honours, UCT).

Research Assistants: Musaji Muemede, Carvalho Issa Nanguar, Iahaia Buanachique, Seliano Alberto Rucunua, with data collection by many others.