TuskWise

For students and researchers: key elephant research papers

These are the 23 peer reviewed studies that matter most to the facts on TuskWise, from the DNA evidence that Africa has two elephant species to the 2024 discovery of name like calls. Each one is listed with its authors, year, journal and DOI, grouped by theme, with a one line summary and a link to the page where we use it.

Last reviewed 8 October 2026. Every DOI on this page was checked against the Crossref registry on 8 October 2026.

How to use this list

Each entry gives the authors, year, title, journal and DOI, plus one plain English sentence on what the study found and the TuskWise page that draws on it. Click a DOI to reach the publisher's page; many of these papers are open access. For a school report, cite the original paper rather than TuskWise when you use its findings, and use the Cite this page box on any TuskWise article when you quote our summary. Teachers will find classroom versions of these ideas in the TuskWise teachers hub.

Species and evolution

  1. Roca, A. L., Georgiadis, N., Pecon-Slattery, J., and O'Brien, S. J. (2001). Genetic Evidence for Two Species of Elephant in Africa. Science, 293(5534), 1473-1477. DOI: 10.1126/science.1059936

    DNA evidence that African savanna and forest elephants are separate species rather than one. Related page: African forest elephant.

  2. Rohland, N., Reich, D., Mallick, S., Meyer, M., Green, R. E., Georgiadis, N. J., Roca, A. L., and Hofreiter, M. (2010). Genomic DNA Sequences from Mastodon and Woolly Mammoth Reveal Deep Speciation of Forest and Savanna Elephants. PLoS Biology, 8(12), e1000564. DOI: 10.1371/journal.pbio.1000564

    Genome sequences, compared with mastodon and woolly mammoth DNA, showed a deep genetic split between forest and savanna elephants. Related page: Species compared.

Memory, families and leadership

  1. McComb, K., Moss, C., Durant, S. M., Baker, L., and Sayialel, S. (2001). Matriarchs As Repositories of Social Knowledge in African Elephants. Science, 292(5516), 491-494. DOI: 10.1126/science.1057895

    Families led by older matriarchs were better at telling familiar from unfamiliar elephant calls. Related page: Do elephants really never forget?, Behavior.

  2. McComb, K., Shannon, G., Durant, S. M., Sayialel, K., Slotow, R., Poole, J., and Moss, C. (2011). Leadership in elephants: the adaptive value of age. Proceedings of the Royal Society B: Biological Sciences, 278(1722), 3270-3276. DOI: 10.1098/rspb.2011.0168

    Older matriarchs responded more appropriately to the roars of male lions, the greater threat. Related page: Intelligence, Do elephants really never forget?.

  3. McComb, K., Baker, L., and Moss, C. (2006). African elephants show high levels of interest in the skulls and ivory of their own species. Biology Letters, 2(1), 26-28. DOI: 10.1098/rsbl.2005.0400

    African elephants showed more interest in elephant skulls and ivory than in other objects. Related page: Behavior: responses to the dead.

  4. Foley, C., Pettorelli, N., and Foley, L. (2008). Severe drought and calf survival in elephants. Biology Letters, 4(5), 541-544. DOI: 10.1098/rsbl.2008.0370

    During a severe drought in Tanzania, calf deaths rose sharply, and survival differed between families. Related page: Do elephants really never forget?.

Intelligence and communication

  1. Plotnik, J. M., de Waal, F. B. M., and Reiss, D. (2006). Self-recognition in an Asian elephant. Proceedings of the National Academy of Sciences, 103(45), 17053-17057. DOI: 10.1073/pnas.0608062103

    An Asian elephant at the Bronx Zoo passed the mirror mark test, evidence of self recognition. Related page: Intelligence.

  2. Hart, B. L., Hart, L. A., McCoy, M., and Sarath, C. R. (2001). Cognitive behaviour in Asian elephants: use and modification of branches for fly switching. Animal Behaviour, 62(5), 839-847. DOI: 10.1006/anbe.2001.1815

    Asian elephants modified branches to make more effective fly switches, a form of tool making. Related page: Intelligence.

  3. Plotnik, J. M., Lair, R., Suphachoksahakun, W., and de Waal, F. B. M. (2011). Elephants know when they need a helping trunk in a cooperative task. Proceedings of the National Academy of Sciences, 108(12), 5116-5121. DOI: 10.1073/pnas.1101765108

    Asian elephants learned to pull a rope together and waited for a partner when they could not succeed alone. Related page: Intelligence.

  4. McComb, K., Shannon, G., Sayialel, K. N., and Moss, C. (2014). Elephants can determine ethnicity, gender, and age from acoustic cues in human voices. Proceedings of the National Academy of Sciences, 111(14), 5433-5438. DOI: 10.1073/pnas.1321543111

    Elephants in Amboseli reacted differently to recorded voices depending on the speaker's ethnic group, gender and age. Related page: Intelligence.

  5. Pardo, M. A., Fristrup, K., Lolchuragi, D. S., Poole, J. H., Granli, P., Moss, C., Douglas-Hamilton, I., and Wittemyer, G. (2024). African elephants address one another with individually specific name-like calls. Nature Ecology & Evolution, 8(7), 1353-1364. DOI: 10.1038/s41559-024-02420-w

    Wild African elephants appear to address one another with individually specific, name like calls. Related page: Intelligence, Behavior.

  6. O'Connell-Rodwell, C. E. (2007). Keeping an "Ear" to the Ground: Seismic Communication in Elephants. Physiology, 22(4), 287-294. DOI: 10.1152/physiol.00008.2007

    A review of how elephants may send and detect low frequency signals through the ground. Related page: Behavior: communication.

Body, brain, sleep and reproduction

  1. Herculano-Houzel, S., Avelino-de-Souza, K., Neves, K., Porfirio, J., Messeder, D., Mattos Feijo, L., and 2 other authors (2014). The elephant brain in numbers. Frontiers in Neuroanatomy, 8. DOI: 10.3389/fnana.2014.00046

    An African elephant brain held about 257 billion neurons, about 98% of them in the cerebellum. Related page: Anatomy: brain.

  2. Gravett, N., Bhagwandin, A., Sutcliffe, R., Landen, K., Chase, M. J., Lyamin, O. I., Siegel, J. M., and Manger, P. R. (2017). Inactivity/sleep in two wild free-roaming African elephant matriarchs - Does large body size make elephants the shortest mammalian sleepers? PLOS ONE, 12(3), e0171903. DOI: 10.1371/journal.pone.0171903

    Two wild matriarchs in Botswana slept about two hours a day on average, often standing. Related page: Behavior: sleep.

  3. Lueders, I., Niemuller, C., Rich, P., Gray, C., Hermes, R., Goeritz, F., and Hildebrandt, T. B. (2012). Gestating for 22 months: luteal development and pregnancy maintenance in elephants. Proceedings of the Royal Society B: Biological Sciences, 279(1743), 3687-3696. DOI: 10.1098/rspb.2012.1038

    Ultrasound and hormone work showing how elephants sustain a pregnancy of about 22 months. Related page: How long are elephants pregnant?.

Ecology

  1. Berzaghi, F., Longo, M., Ciais, P., Blake, S., Bretagnolle, F., Vieira, S., and 3 other authors (2019). Carbon stocks in central African forests enhanced by elephant disturbance. Nature Geoscience, 12(9), 725-729. DOI: 10.1038/s41561-019-0395-6

    Forest elephant feeding shifts central African forests toward larger, denser trees that store more carbon. Related page: African forest elephant.

  2. Campos-Arceiz, A., and Blake, S. (2011). Megagardeners of the forest - the role of elephants in seed dispersal. Acta Oecologica, 37(6), 542-553. DOI: 10.1016/j.actao.2011.01.014

    A review of elephants as seed dispersers, carrying seeds long distances in their dung. Related page: Habitats and diet.

Conservation and welfare

  1. Campbell-Staton, S. C., Arnold, B. J., Gonçalves, D., Granli, P., Poole, J., Long, R. A., and Pringle, R. M. (2021). Ivory poaching and the rapid evolution of tusklessness in African elephants. Science, 374(6566), 483-487. DOI: 10.1126/science.abe7389

    Heavy ivory poaching during civil war in Mozambique's Gorongosa was followed by a rise in tuskless females. Related page: Conservation: poaching.

  2. Wittemyer, G., Northrup, J. M., Blanc, J., Douglas-Hamilton, I., Omondi, P., and Burnham, K. P. (2014). Illegal killing for ivory drives global decline in African elephants. Proceedings of the National Academy of Sciences, 111(36), 13117-13121. DOI: 10.1073/pnas.1403984111

    Illegal killing for ivory from 2010 to 2012 was high enough to drive a decline in African elephants. Related page: Conservation: poaching.

  3. Chase, M. J., Schlossberg, S., Griffin, C. R., Bouché, P. J. C., Djene, S. W., Elkan, P. W., and 8 other authors (2016). Continent-wide survey reveals massive decline in African savannah elephants. PeerJ, 4, e2354. DOI: 10.7717/peerj.2354

    The Great Elephant Census, an aerial survey of savanna elephants across 18 countries, reported a steep decline. Related page: How many elephants are left?.

  4. Maisels, F., Strindberg, S., Blake, S., Wittemyer, G., Hart, J., Williamson, E. A., and 56 other authors (2013). Devastating Decline of Forest Elephants in Central Africa. PLoS ONE, 8(3), e59469. DOI: 10.1371/journal.pone.0059469

    Surveys across central Africa found forest elephant numbers fell sharply between 2002 and 2011. Related page: African forest elephant.

  5. King, L. E., Douglas-Hamilton, I., and Vollrath, F. (2007). African elephants run from the sound of disturbed bees. Current Biology, 17(19), R832-R833. DOI: 10.1016/j.cub.2007.07.038

    Elephants moved away from recordings of disturbed bees, the basis of beehive fence research. Related page: Conservation: conflict.

  6. Clubb, R., Rowcliffe, M., Lee, P., Mar, K. U., Moss, C., and Mason, G. J. (2008). Compromised Survivorship in Zoo Elephants. Science, 322(5908), 1649-1649. DOI: 10.1126/science.1164298

    Elephants in European zoos had shorter lifespans than elephants in protected or working populations. Related page: Conservation: captivity.

Other sources we use

Population figures and conservation status come from the IUCN Red List and the reports of its African and Asian Elephant Specialist Groups, and from national surveys such as India's 2021 to 2025 DNA based count. Animal facts such as weights and daily food intake come from accredited zoos, mainly San Diego Zoo Wildlife Alliance and the Smithsonian's National Zoo and Conservation Biology Institute. Each article lists its own sources at the bottom of the page.