Large brains and cognition: Where do elephants fit in?
Abstract
Among terrestrial mammals, elephants share the unique status, along with humans and great apes, of having large brains, being long-lived and having offspring that require long periods of dependency. Elephants have the largest brains of all terrestrial mammals, including the greatest volume of cerebral cortex. In contrast to what one might expect from such a large-brained species, the performance of elephants in cognitive feats, such as tool use, visual discrimination learning and tests of "insight" behavior, is unimpressive in comparison to the performance by chimpanzees and, of course, humans. Where elephants do seem to excel is in long-term, extensive spatial-temporal and social memory. In addition, elephants appear to be somewhat unique among non-human species in their reactions to disabled and deceased conspecifics, exhibiting behaviors that are mindful of "theory-of-mind" phenomena. Information gleaned from studies on the neural cytoarchitecture of large brains reveals that the neurons of the cerebral cortex of elephants are much less densely populated than in large-brained primates. The interactions between cortical neurons would appear to be more global and less compartmentalized into local areas, and cortical information processing slower, than in great apes and humans. Although focused neural cytoarchitecture studies on the elephant are needed, this comparative perspective on the cortical neural cytoarchitecture appears to relate to differences in behavior between elephants and their primate counterparts.
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Elephants have large brains with a neuron density three times that of humans (257 billion neurons).
“Elephants have large brains with a neuron density three times that of humans (257 billion neurons; Hart et al., 2008; Herculano-Houzel et al., 2014; Shoshani et al., 2006)”
From Do elephants really never forget? What we know about elephant memory and a call for further investigation (2025) · cites this paper as Hart et al., 2008
Cites
The descent of man and selection in relation to sex
チャールス, ダ., & 茂穂, 田. (1907) · Tokyo Jinrui Gakkai zasshi · doi:10.1537/ase1887.22.495 · Cited by 10,909 · see citing works
The human hippocampus and spatial and episodic memory
Burgess, N., Maguire, E. A., & O'Keefe, J. (2002) · Neuron · doi:10.1016/s0896-6273(02)00830-9 · Cited by 2,401 · see citing works
EVOLUTION OF THE BRAIN AND INTELLIGENCE
Radinsky, L., & Jerison, H. J. (1975) · Evolution · doi:10.2307/2407154 · Cited by 1,982 · see citing works
Gallup, G. G. (1970) · Science · doi:10.1126/science.167.3914.86 · Cited by 1,794 · see citing works
Megaherbivores: The Influence of Very Large Body Size on Ecology
Owen‐Smith, N. (1988) · Cited by 882 · see citing works
Behavioral adaptations to pathogens and parasites: Five strategies
Hart, B. L. (1990) · Neuroscience & Biobehavioral Reviews · doi:10.1016/s0149-7634(05)80038-7 · Cited by 734 · see citing works
Self-citationA universal scaling law between gray matter and white matter of cerebral cortex
Zhang, K., & Sejnowski, T. J. (2000) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.090504197 · Cited by 723 · see citing works
Mirror self-recognition in the bottlenose dolphin: a case of cognitive convergence
Reiss, D., & Marino, L. (2001) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.101086398 · Cited by 711 · see citing works
Self-recognition in an Asian elephant
Plotnik, J. M., de Waal, F. B. M., & Reiss, D. (2006) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.0608062103 · Cited by 709 · see citing works
Matriarchs As Repositories of Social Knowledge in African Elephants
McComb, K., Moss, C., Durant, S. M., Baker, L., & Sayialel, S. (2001) · Science · doi:10.1126/science.1057895 · Cited by 651 · see citing works
Manufacture and use of hook-tools by New Caledonian crows
Hunt, G. R. (1996) · Nature · doi:10.1038/379249a0 · Cited by 645 · see citing works
Shaping of Hooks in New Caledonian Crows
Weir, A. A. S., Chappell, J., & Kacelnik, A. (2002) · Science · doi:10.1126/science.1073433 · Cited by 625 · see citing works
Haug, H. (1987) · American Journal of Anatomy · doi:10.1002/aja.1001800203 · Cited by 481 · see citing works
The conditions for tool use in primates: implications for the evolution of material culture
van Schaik, C. P., Deaner, R. O., & Merrill, M. Y. (1999) · Journal of Human Evolution · doi:10.1006/jhev.1999.0304 · Cited by 463 · see citing works
(2003) · Harvard University Press eBooks · doi:10.4159/harvard.9780674419131 · Cited by 431 · see citing works
The living elephants: evolutionary ecology, behavior, and conservation
(2004) · Choice Reviews Online · doi:10.5860/choice.41-4051 · Cited by 379 · see citing works
Unusually extensive networks of vocal recognition in African elephants
McComb, K., Moss, C., Sayialel, S., & Baker, L. (2000) · Animal Behaviour · doi:10.1006/anbe.2000.1406 · Cited by 348 · see citing works
Evolution of the Brain and Intelligence
Gould, S. J. (1974) · Science · doi:10.1126/science.185.4149.400 · Cited by 329 · see citing works
Megaherbivores: The influence of very large body size on ecology
Albon, S. D. (1989) · Trends in Ecology & Evolution · doi:10.1016/0169-5347(89)90043-8 · Cited by 328 · see citing works
Genetic Evidence for Two Species of Elephant in Africa
Roca, A. L., Georgiadis, N., Pecon‐Slattery, J., & O’Brien, S. J. (2001) · Science · doi:10.1126/science.1059936 · Cited by 326 · see citing works
Long-distance communication of acoustic cues to social identity in African elephants
McComb, K., Reby, D., Baker, L., Moss, C. J., & Sayialel, S. (2003) · Animal Behaviour · doi:10.1006/anbe.2003.2047 · Cited by 316 · see citing works
Povinelli, D. J., Rulf, A. B., Landau, K. R., & Bierschwale, D. T. (1993) · Journal of Comparative Psychology · doi:10.1037/0735-7036.107.4.347 · Cited by 309 · see citing works
The organization of neocortex in mammals: are species differences really so different?
Krubitzer, L. A. (1995) · Trends in Neurosciences · doi:10.1016/0166-2236(95)93938-t · Cited by 298 · see citing works
On the evolution and geometry of the brain in mammals
Hofman, M. A. (1989) · Progress in Neurobiology · doi:10.1016/0301-0082(89)90013-0 · Cited by 295 · see citing works
Why is Brain Size so Important:Design Problems and Solutions as Neocortex Gets Biggeror Smaller
Kaas, J. H. (2000) · Brain and Mind · doi:10.1023/a:1010028405318 · Cited by 225 · see citing works
Behavioural reactions of elephants towards a dying and deceased matriarch
Douglas-Hamilton, I., Bhalla, S., Wittemyer, G., & Vollrath, F. (2006) · Applied Animal Behaviour Science · doi:10.1016/j.applanim.2006.04.014 · Cited by 219 · see citing works
Hippocampal Function in Cognition
Sweatt, J. D. (2004) · Psychopharmacology · doi:10.1007/s00213-004-1795-9 · Cited by 217 · see citing works
Elephant memories. Thirteen years in the life of an elephant family
(1993) · Environment International · doi:10.1016/0160-4120(93)90102-n · Cited by 215 · see citing works
Tower, D. B. (1954) · Journal of Comparative Neurology · doi:10.1002/cne.901010103 · Cited by 203 · see citing works
Evolutionary Anatomy of the Primate Cerebral Cortex
(2001) · Cambridge University Press eBooks · doi:10.1017/cbo9780511897085 · Cited by 196 · see citing works
Shoshani, J., Kupsky, W. J., & Marchant, G. H. (2006) · Brain Research Bulletin · doi:10.1016/j.brainresbull.2006.03.016 · Cited by 186 · see citing works
Elephant memories. Thirteen years in the life of an elephant family
Eltringham, S. K. (1989) · Animal Behaviour · doi:10.1016/0003-3472(89)90080-8 · Cited by 179 · see citing works
The Ai project: historical and ecological contexts
Matsuzawa, T. (2003) · Animal Cognition · doi:10.1007/s10071-003-0199-2 · Cited by 164 · see citing works
Annotation: The savant syndrome
Heaton, P. F., & Wallace, G. L. (2004) · Journal of Child Psychology and Psychiatry · doi:10.1111/j.1469-7610.2004.t01-1-00284.x · Cited by 143 · see citing works
The Savant Syndrome: Intellectual impairment and exceptional skill.
Miller, L. K. (1999) · Psychological Bulletin · doi:10.1037/0033-2909.125.1.31 · Cited by 142 · see citing works
Buss, I. O., Hamilton, I. D., & Hamilton, O. D. (1976) · Journal of Wildlife Management · doi:10.2307/3799973 · Cited by 131 · see citing works
Cognitive behaviour in Asian elephants: use and modification of branches for fly switching
Hart, B. L., Hart, L. A., McCoy, M., & Sarath, C. R. (2001) · Animal Behaviour · doi:10.1006/anbe.2001.1815 · Cited by 130 · see citing works
Self-citationPovinelli, D. J. (1989) · Journal of comparative psychology · doi:10.1037/0735-7036.103.2.122 · Cited by 126 · see citing works
Animal Social Complexity: Intelligence, Culture, and Individualized Societies
Megan, S. (2004) · Complicity An International Journal of Complexity and Education · doi:10.29173/cmplct8720 · Cited by 120 · see citing works
Patterns of molecular genetic variation among African elephant populations
Comstock, K. E., Georgiadis, N., Pecon‐Slattery, J., et al. (2002) · Molecular Ecology · doi:10.1046/j.1365-294x.2002.01615.x · Cited by 114 · see citing works
Tool use by wild and captive elephants
Chevalier-Skolnikoff, S., & Liska, J. (1993) · Animal Behaviour · doi:10.1006/anbe.1993.1183 · Cited by 111 · cited 1× · see citing works
“The repertoire of tool use in Asian and African elephants also includes scratching with a stick and throwing sticks or stones at rodents competing for fruit beneath a tree (Chevalier-Skolnikoff and Liska, 1993; Hart and Hart, 1994; Kurt and Hartl, 1995; Wickler and Seibt, 1997).”
BackgroundAfrican elephants show high levels of interest in the skulls and ivory of their own species
McComb, K., Baker, L., & Moss, C. (2006) · Biology Letters · doi:10.1098/rsbl.2005.0400 · Cited by 98 · see citing works
Movements and patterns of habitat utilization of elephants in Tsavo National Park, Kenya
Leuthold, W., & Sale, J. B. (1973) · African Journal of Ecology · doi:10.1111/j.1365-2028.1973.tb00099.x · Cited by 88 · see citing works
[Cerebral amyloid angiopathy].
Almas, T., Vedeler, C. A., Moen, G., & Mörk, S. (1998) · PubMed · Cited by 87 · see citing works
Viljoen, P. J. (1989) · Journal of Zoology · doi:10.1111/j.1469-7998.1989.tb02561.x · Cited by 85 · see citing works
Humle, T., & Matsuzawa, T. (2001) · Folia Primatologica · doi:10.1159/000049924 · Cited by 85 · see citing works
Creative or created: Using anecdotes to investigate animal cognition
Bates, L. A., & BYRNE, R. (2007) · Methods · doi:10.1016/j.ymeth.2006.11.006 · Cited by 83 · see citing works
Glickman, S. E., Short, R. V., & Renfree, M. B. (2005) · Hormones and Behavior · doi:10.1016/j.yhbeh.2005.07.013 · Cited by 82 · see citing works
The Scaling of White Matter to Gray Matter in Cerebellum and Neocortex
Bush, E. C., & Allman, J. M. (2003) · Brain, Behavior and Evolution · doi:10.1159/000068880 · Cited by 81 · see citing works
Parcellation and Area-Area Connectivity as a Function of Neocortex Size
Changizi, M. A., & Shinsuke Shimojo, A. (2005) · Brain Behavior and Evolution · doi:10.1159/000085942 · Cited by 80 · see citing works
3. Sources of Social Complexity in the Three Elephant Species
Payne, Κ. (2003) · Harvard University Press eBooks · doi:10.4159/harvard.9780674419131.c5 · Cited by 80 · see citing works
Feelings and Emotions: The Amsterdam Symposium
(2004) · doi:10.1017/cbo9780511806582 · Cited by 79 · see citing works
Scaling laws in the mammalian neocortex: Does form provide clues to function?
Harrison, K. H., Hof, P. R., & Wang, S. S.-H. (2002) · Journal of Neurocytology · doi:10.1023/a:1024178127195 · Cited by 78 · see citing works
Not all chimpanzees (Pan troglodytes) show self-recognition
Swartz, K. B., & Evans, S. (1991) · Primates · doi:10.1007/bf02381939 · Cited by 78 · see citing works
Home range and seasonal movement of elephants in the Kunene Region, northwestern Namibia
Leggett, K. E. A. (2006) · African Zoology · doi:10.3377/1562-7020(2006)41[17:hrasmo]2.0.co;2 · Cited by 76 · see citing works
Arealization of the Neocortex in Mammals: Genetic and Epigenetic Contributions to the Phenotype
Krubitzer, L. A., & Huffman, K. J. (2000) · Brain Behavior and Evolution · doi:10.1159/000006667 · Cited by 75 · see citing works
Molecules remodel the mammalian tree
JONG, W. W. D. (1998) · Trends in Ecology & Evolution · doi:10.1016/s0169-5347(98)01335-4 · Cited by 72 · see citing works
Brain evolution in hominids: are we at the end of the road?
Hofman, M. A. (2001) · Cambridge University Press eBooks · doi:10.1017/cbo9780511897085.009 · Cited by 67 · see citing works
Brain of the African elephant (Loxodonta africana): Neuroanatomy from magnetic resonance images
Hakeem, A. Y., Hof, P. R., Sherwood, C. C., Switzer, R. C., Rasmussen, L. E. L., & Allman, J. M. (2005) · The Anatomical Record Part A Discoveries in Molecular Cellular and Evolutionary Biology · doi:10.1002/ar.a.20255 · Cited by 56 · see citing works
Fly switching by Asian elephants: tool use to control parasites
Hart, B. L., & Hart, L. A. (1994) · Animal Behaviour · doi:10.1006/anbe.1994.1209 · Cited by 53 · cited 3× · see citing works
“More recently, studies have documented the efficacy of fly switching in repelling flies (Hart and Hart, 1994) and the modification of branches to use as switches (Hart et al., 2001).”
BackgroundSelf-citationCozzi, B., Spagnoli, S., & Bruno, L. (2001) · Brain Research Bulletin · doi:10.1016/s0361-9230(00)00456-1 · Cited by 50 · see citing works
Elephant life: fifteen years of high population density
(1991) · Choice Reviews Online · doi:10.5860/choice.29-0304 · Cited by 45 · see citing works
On the Possibility of Animal Empathy
de Waal, F. Β. Μ. (2004) · Cambridge University Press eBooks · doi:10.1017/cbo9780511806582.022 · Cited by 39 · see citing works
SIMULTANEOUS VISUAL DISCRIMINATION IN ASIAN ELEPHANTS
Nissani, M., Hoefler-Nissani, D., Lay, U. T., & Htun, U. W. (2005) · Journal of the Experimental Analysis of Behavior · doi:10.1901/jeab.2005.34-04 · Cited by 39 · see citing works
Reconstructing the Organization of Neocortex of the First Mammals and Subsequent Modifications
Kaas, J. (2007) · Evolution of Nervous Systems · doi:10.1016/b0-12-370878-8/00057-4 · Cited by 32 · see citing works
Scaling the Brain and Its Connections
Changizi, M. (2007) · Evolution of Nervous Systems · doi:10.1016/b0-12-370878-8/00061-6 · Cited by 32 · see citing works
Evolution of Longevity in Ungulates and Carnivores
Cutler, R. G. (2009) · Gerontology · doi:10.1159/000212324 · Cited by 29 · see citing works
Do asian elephants (Elephas maximus) apply causal reasoning to tool-use tasks?
Nissani, M. (2006) · Journal of Experimental Psychology: Animal Behavior Processes · doi:10.1037/0097-7403.32.1.91 · Cited by 28 · see citing works
Neuronal correlates of corticalization in mammals: A theory
Hofman, M. A. (1985) · Journal of Theoretical Biology · doi:10.1016/s0022-5193(85)80117-x · Cited by 27 · see citing works
CURRENT ISSUES AND EMERGING THEORIES IN ANIMAL COGNITION
Boysen, S. T., & Himes, G. T. (1999) · Annual Review of Psychology · doi:10.1146/annurev.psych.50.1.683 · Cited by 27 · see citing works
Spatial and Social Cognition in Corvids: An Evolutionary Approach
Balda, R. P., & Kamil, A. C. (2002) · The MIT Press eBooks · doi:10.7551/mitpress/1885.003.0020 · Cited by 26 · see citing works
A two-component theory of encephalization in mammals
Hofman, M. A. (1982) · Journal of Theoretical Biology · doi:10.1016/0022-5193(82)90211-9 · Cited by 23 · see citing works
Aimed Object‐throwing by a Wild African Elephant in an Interspecific Encounter
Wickler, W., & Seibt, U. (1997) · Ethology · doi:10.1111/j.1439-0310.1997.tb00152.x · Cited by 23 · see citing works
Captured in the Net of Space and Time: Understanding Cortical Field Evolution
Krubitzer, L. A., & Hunt, D. L. (2007) · Evolution of Nervous Systems · doi:10.1016/b0-12-370878-8/00058-6 · Cited by 19 · see citing works
Theory of Mind and Insight in Chimpanzees, Elephants, and Other Animals?
Nissani, M. (2004) · Comparative Vertebrate Cognition · doi:10.1007/978-1-4419-8913-0_7 · Cited by 19 · see citing works
Zähmung und Dressurleistungen indischer Arbeitselefanten
Koehler, O., & Altevogt, B. R. U. R. (2010) · Zeitschrift für Tierpsychologie · doi:10.1111/j.1439-0310.1954.tb02173.x · Cited by 16 · see citing works
Evolution of the Elephant Brain: A Paradox between Brain Size and Cognitive Behavior
Hart, B. L., & Hart, L. A. (2007) · Evolution of Nervous Systems · doi:10.1016/b0-12-370878-8/00343-8 · Cited by 12 · see citing works
Self-citationThe Biology of Large African Mammals in Their Environment.
Leader‐Williams, N., Jewell, P. A., & Maloiy, G. M. O. (1990) · Journal of Applied Ecology · doi:10.2307/2404404 · Cited by 9 · see citing works
Dr. Skinner on the 'disintegration of perceptual principles' of psychotics.
Yacorzynski, G. K. (1943) · Journal of Experimental Psychology · doi:10.1037/h0059425 · Cited by 3 · see citing works
Peal, S. E. (1879) · Nature · doi:10.1038/021034b0 · Cited by 2 · see citing works
Wiese (2000) · doi:10.1002/1098-2361(2000)19:5
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Millan, M. J., Agid, Y., Brüne, M., Bullmore, E. T., Carter, C. S., Clayton, N. S., Connor, R., Davis, S., Deakin, B., DeRubeis, R. J., et al. (2012) · Nature Reviews Drug Discovery · doi:10.1038/nrd3628 · Cited by 1,292 · see citing works
BODDY, A. M., McGOWEN, M. R., SHERWOOD, C. C., GROSSMAN, L. I., GOODMAN, M., & WILDMAN, D. E. (2012) · Journal of Evolutionary Biology · doi:10.1111/j.1420-9101.2012.02491.x · Cited by 207 · see citing works
Millan, M. J., Goodwin, G. M., Meyer-Lindenberg, A., & Ove Ögren, S. (2015) · European Neuropsychopharmacology · doi:10.1016/j.euroneuro.2015.01.016 · Cited by 132 · see citing works
Perdue, B. M., Talbot, C. F., Stone, A. M., & Beran, M. J. (2012) · Animal Cognition · doi:10.1007/s10071-012-0521-y · Cited by 115 · see citing works
Insightful problem solving in an Asian elephant
Foerder, P., Galloway, M., Barthel, T., Moore, D. E., & Reiss, D. (2011) · PLoS ONE · doi:10.1371/journal.pone.0023251 · Cited by 107 · see citing works
Successful acquisition of an olfactory discrimination test by Asian elephants, Elephas maximus
Arvidsson, J., Amundin, M., & Laska, M. (2012) · Physiology & Behavior · doi:10.1016/j.physbeh.2011.08.021 · Cited by 49 · see citing works
Evidence of means–end behavior in Asian elephants (Elephas maximus)
Irie-Sugimoto, N., Kobayashi, T., Sato, T., & Hasegawa, T. (2008) · Animal Cognition · doi:10.1007/s10071-007-0126-z · Cited by 48 · see citing works
Sharma, N., Pokharel, S. S., Kohshima, S., & Sukumar, R. (2020) · Primates · doi:10.1007/s10329-019-00739-8 · Cited by 42 · see citing works
Physiological plasticity in elephants: highly dynamic glucocorticoids in African and Asian elephants
Pokharel, S. S., & Brown, J. L. (2023) · Conservation Physiology · doi:10.1093/conphys/coad088 · Cited by 23 · see citing works
Novel behaviour shown by an Asian elephant in the context of allomothering
Vidya, T. N. C. (2014) · acta ethologica · doi:10.1007/s10211-013-0168-y · Cited by 13 · see citing works
Hope, S. F., Willgohs, K. R., Dittakul, S., & Plotnik, J. M. (2025) · Learning & Behavior · doi:10.3758/s13420-024-00655-y · Cited by 8 · see citing works
Long-Term Olfactory Memory in African Elephants
Hoerner, F., Lawrenz, A., Oerke, A.-K., Müller, D. W. H., Azogu-Sepe, I., Roller, M., Damerau, K., & Preisfeld, A. (2023) · Animals · doi:10.3390/ani13040679 · Cited by 8 · see citing works
Doyle, C. (2018) · The Palgrave Handbook of Practical Animal Ethics · doi:10.1057/978-1-137-36671-9_12 · Cited by 5 · see citing works
- Authors
- Hart, B. L., Hart, L. A., & Pinter-Wollman, N.
- Affiliations
- University of California, Davis (US)
- Corresponding author
- Hart, Benjamin L.
- Venue
- Neuroscience & Biobehavioral Reviews
- Publisher
- Elsevier BV
- Published
- 2008-01
- Volume
- 32
- Issue
- 1
- Pages
- 86-98
- ISSN
- 0149-7634 (ISSN-L)
- OpenAlex
- W2075821884
- Cited by
- 185
- References
- 82
- Citations by year
- 2025: 8 · 2024: 10 · 2023: 12 · 2022: 16 · 2021: 10 · 2020: 9 · 2019: 9 · 2018: 7 · 2017: 29 · 2016: 6 · 2015: 13 · 2014: 16 · 2013: 5 · 2012: 7
- FWCI
- 0.78
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- Neuroscience & Biobehavioral Reviews (publishedVersion)
- PubMed (publishedVersion)
- Topics
- Social Sciences › Psychology › Social Psychology › Primate Behavior and Ecology
- Health Sciences › Veterinary › Small Animals › Animal Behavior and Welfare Studies
- Health Sciences › Medicine › Urology › Comparative Animal Anatomy Studies
- Keywords
- elephants, brain size, cognition, cerebral cortex, spatial memory, social memory, tool use, visual discrimination learning, theory of mind, long-term memory, primate comparison
- MeSH terms
- Animals, Behavior, Animal, Brain, Cognition, Elephants, Humans, Memory, Organ Size, Tool Use Behavior
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