Dr David Lloyd-Jones

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Biography

I am a behavioural ecologist, ornithologist and post-doctoral researcher at the FitzPatrick Institute for African Ornithology, University of Cape Town. I grew up in southern Tanzania and this biodiverse setting nurtured my passion for natural history and field research. I completed my undergraduate studies at the University of Canterbury, New Zealand. During this period, I became fascinated in avian brood parasitism, co-evolutionary interactions and avian communication and I have conducted research exploring these topics. After returning to East Africa I worked on biodiversity surveys and aerial surveys of large mammals, while also keeping bees and experimenting with beekeeping techniques.

In 2016 I first went to northern Mozambique’s stunning Niassa Special Reserve for work on an aerial survey. In 2017 I returned to working there – in collaboration with Prof. Claire Spottiswoode – to study the remarkable mutualism between human honey-hunters and the greater honeyguides (Indicator indicator). I have been involved in honeyguide research since then, completing my PhD in 2025, and continue to be inspired by complex species interactions and the challenge of studying them using field experiments. It continues to be a profound privilege spending extended time with people who coexist alongside wildlife inside Niassa’s vast wilderness.

Research focus

My current focus is on the ecology and economics of human-honeyguide cooperation and in particular, gaining a better understanding of the foraging behaviour and decision-making of both honeyguides and human honey-hunters at the population level. More specifically, I am looking at how the mutualism shifts the costs and benefits of honey-hunting, using spatial data for humans and measuring the rewards (wax, honey) for both parties. I am also testing whether honeyguides learn to recognise cheating honey-hunters, and whether they punish them?

With the help of a honey-hunting community I am collecting a wide range of data on natural honey-hunting journeys and their payoffs. I am trying to discover whether humans learn to recognise unskilled honeyguides, and whether they avoid them? I am also investigating questions relating to honeyguide-human communication, and the ecosystem effects of honey-hunting at a landscape level. Our multi-year dataset enables us to ask how honeyguides change human movement and foraging patterns.

Peer-reviewed publications

Dunkley, K., Cantor, M., Afan, A.I., Ahlibi, D.S., Allen, S.J., Amphaeris, J., Atkins, S., Attwood, M.C., Bankhead, K., Blair, C.J., Bronstein, J.L., Camargo, Y.R.R., Carvalho, S., Channer, L.W., Cuthill, R.R.T., Das, J., Daura-Jorge, F.G., Deb, A.K., Dixit, T., Dounias, E., Dyble, M., Farine, D.R., Freymann, E., He, P., Hoffmann, L.S., Isack, H.A., Ilha, E.B., Kamboe, W.-B.W., Kilawi, A.O., Kingston, A., Laltaika, E.A., Lloyd-Jones, D.J., Lund, J., Machado, A.M.S., McGarvey, K., M’manga, G.M., Mphetlhe, R., Moreno, I.B., Ngcamphalala, C.A., Nhlabatsi, S.O., Nwaogu, C.J., Pierotti, R., Reeves, I.M., Robinson, E.J.H., Samad, I., Sanda, M., Serpa, N.B., Simões-Lopes, P.C., Spottiswoode, C.N., Soma, T., Sridhar, H., Tun, T., Uomini, N.T., Valle-Pereira, J.V.S., van Holstein, L., Wood, B.M., Cram, D.L., & van der Wal, J.E.M. 2026. The ecology and evolution of cues and signals in animal interspecies cooperation. Animal Behaviour, 237: 123611.
Lloyd-Jones, D.J., Muamedi, M. & Spottiswoode, C.N. 2025 To bees or not to bees: greater honeyguides sometimes guide humans to animals other than bees, but likely not as punishment. Ecology & Evolution 15:e71136.
van der Wal, J.E.M., Dauda, C., Lloyd-Jones, D.J., Murico, H., Begg, C.M., Begg, K.S., Jorge, A.A., Spottiswoode, C.N. 2025. The economic value of human-honeyguide mutualism in Reserva Especial do Niassa, Moçambique. Ecosystem Services 101696. Read abstract in English and Portuguese here
Cram, D.L., Lloyd-Jones, D.J., van der Wal, J.E.M., Lund, J., Buanachique, I.O., Muamedi, M., Nanguar, C.I., Ngovene, A., Raveh, S., Boner, W., Spottiswoode, C.N. 2023. Guides and cheats: producer-scrounger dynamics in the human-honeyguide mutualism. Proceedings Royal Society B 290, 20232024.
van der Wal, J.E.M., Afan, A.I., Anyawire, M., Begg, C.M., Begg, K.S., Dabo, G.A., Gedi, I.I., Harris, J.A., Isack, H.A., Ibrahim^,J.I., Jamie, G.A., Kamboe^,W.-B.W., Kilawi, A.O., Kingston, A., Laltaika, E.A., Lloyd-Jones, D.J., M’manga, G.M., Muhammad, N.Z., Ngcamphalala, C.A., Nhlabatsi, S.O., Oleleteyo, T.T., Sanda, M., Tsamkxao, L., Wood, B.M., Spottiswoode, C.N., Cram, D.L. 2023 Do honey badgers and greater honeyguide birds cooperate to access bees’ nests? Ecological evidence and honey-hunter accounts. Journal of Zoology 321: 22-32 https://doi.org/10.1111/jzo.13093 Read abstract in English and Kiswahili here
Lloyd-Jones, D.J., St Clair, J.J., Cram, D. L., Yassene, O., van der Wal, J.E. and Spottiswoode, C.N. 2022. When wax wanes: competitors for beeswax stabilize rather than jeopardize the honeyguide–human mutualism. Proceedings of the Royal Society B 289, 20221443.
Spottiswoode, C.N., & Lund, J. & Lloyd-Jones, D.J. 2022 Quick Guide: Honeyguides. Current Biology 32: R1072-R107
Torney, C. J., Laxton, M., Lloyd-Jones, D. J., Kohi, E. M., Frederick, H. L., Moyer, D. C., Mrisha, C., Mwita, M., & Hopcraft, J. G. C. 2022. Estimating the abundance of a group-living species using multi-latent spatial models. Methods in Ecology and Evolution, 14: 77-86.
Cram, D.L., van der Wal, J.E.M., Uomini, N.T., Cantor, M., Afan, A.I., Attwood, M.C., Amphaeris, J., Balasani, F., Blair, C.J., Bronstein, J.L., Buanachique, I.O., Cuthill, R.R.T., Das, J., Daura-Jorge, F.G., Deb, A., Dixit, T., Dlamini, G.S., Dounias, E., Gedi, I.I., Gruber, M., Hoffman, L.S., Holzlehner, T., Isack, H.A., Laltaika, A.E., Lloyd-Jones, D.J., Lund, J., Machado, A.M.S., Mahadevan, L., Moreno, I.B., Nwaogu, C.J., Pereira, V.L., Pierotti, R., Rucunua, S.A., dos Santos, W.F., Serpa, N., Smith, B.D., Sridhar, H., Tolkova, I., Tun, T., Valle-Pereira, J.V.S., Wood, B.M., Wrangham, R.W. & Spottiswoode, C.N. 2022 The ecology and evolution of human-wildlife cooperation. People and Nature 4: 841-855. Read abstract in English, Portuguese and Kiswahili here
van der Wal, J.E.M., Spottiswoode, C.N., Uomini, N.T., Cantor, M., Daura-Jorge, F.G., Afan, A.I., Attwood, M.C., Amphaeris, J., Balasani, F., Begg, C.M., Blair, C.J., Bronstein, J.L., Buanachique, I.O., Cuthill, R.R.T., Das, J., Deb, A., Dixit, T., Dlamini, G.S., Dounias, E., Gedi, I.I., Gruber, M., Hoffman, L.S., Holzlehner, T., Isack, H.A., Laltaika, A.E., Lloyd-Jones, D.J., Lund, J., Machado, A.M.S., Mahadevan, L., Moreno, I.B., Nwaogu, C.J., Pereira, V.L., Pierotti, R., Rucunua, S.A., dos Santos, W.F., Serpa, N., Smith, B.D., Tolkova, I., Tun, T., Valle-Pereira, J.V.S., Wood, B.M., Wrangham, R.W. & Cram, D.L. 2022 Safeguarding human-wildlife cooperation. Conservation Letters e12886 Read abstract in English, Portuguese and Kiswahili here
Torney, C.J.*, Lloyd-Jones, D.J.*, Chevallier, M., Moyer, D.C., Maliti, H.T., Mwita, M., Kohi, E.M. & Hopcraft, G.C. 2019 A comparison of deep learning and citizen science techniques for counting wildlife in aerial survey images. Methods in Ecology and Evolution 10: 779-787 *equal contributors
Torney CJ, Dobson AP, Borner F, Lloyd-Jones DJ, Moyer D, et al. (2016), Assessing rotation-Invariant feature classification for automated wildebeest population counts. Plos One 11: e0156342.
Lloyd-Jones, D.J. & Briskie, J.V. 2016 Mutual wattle ornaments in the South Island saddleback (Philesturnus carunculatus) function as armaments. Ethology 122: 61-71.
Khwaja, N. & Lloyd-Jones, D.J. 2015 Eurasian blackbird (Turdus merula) nest parasitised by song thrush (T. philomelos). Notornis 62: 41-44.
Lloyd-Jones, D.J. 2014 Ultrasonic harmonics in the calls of rock wren (Xenicus gilviventris). Nortornis 61: 165-169.

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News

New study shows that honey-hunter calls vary regionally like dialects

Jan 22, 2026

We have published a new study in People and Nature showing that people in northern Mozambique use regionally distinct “dialects” when communicating with honeyguides. Led by Jessica van der Wal, the paper shows that human–honeyguide communication varies across landscapes in ways that mirror regional variation in human languages. Despite these differences in calls, cooperation between people and honeyguides remains successful and important for human livelihoods across the Niassa Special Reserve, suggesting that both species adjust to one another across their shared landscape.

New paper on honey-hunting with honeyguides in western Africa

Dec 4, 2025

We are pleased to share our new paper on honey-hunting with honeyguides, in western Africa this time. Led by Wiro-Bless Kamboe as part of his MSc project, and co-authored with Claire Spottiswoode and Timothy Khan Aikins, with Jessica van der Wal as senior author, 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 socio-economic changes, such as increased access to motorised transport, have disrupted this relationship.