Animal Cognition

, Volume 15, Issue 5, pp 955–961 | Cite as

Putting the elephant back in the herd: elephant relative quantity judgments match those of other species

  • Bonnie M. Perdue
  • Catherine F. Talbot
  • Adam M. Stone
  • Michael J. Beran
Original Paper


The ability to discriminate between quantities has been observed in many species. Typically, when an animal is given a choice between two sets of food, accurate performance (i.e., choosing the larger amount) decreases as the ratio between two quantities increases. A recent study reported that elephants did not exhibit ratio effects, suggesting that elephants may process quantitative information in a qualitatively different way from all other nonhuman species that have been tested (Irie-Sugimoto et al. in Anim Cogn 12:193–199, 2009). However, the results of this study were confounded by several methodological issues. We tested two African elephants (Loxodonta africana) to more thoroughly investigate relative quantity judgment in this species. In contrast to the previous study, we found evidence of ratio effects for visible and nonvisible sequentially presented sets of food. Thus, elephants appear to represent and compare quantities in much the same way as other species, including humans when they are prevented from counting. Performance supports an accumulator model in which quantities are represented as analog magnitudes. Furthermore, we found no effect of absolute magnitude on performance, providing support against an object-file model explanation of quantity judgment.


Relative quantity judgment Ratio effects Elephants Accumulator model Object-file model 



We would like to thank the Zoo Atlanta elephant care staff, including Nathan Elgart, Kathryn Hunt, Steve Crews, as well as Rebecca Snyder and Megan Wilson. This work was approved by the Georgia Tech (A10040) and Georgia State University (A11028) IACUC as well as the Zoo Atlanta Scientific Review Committee. This work was supported in part by NICHD Grant P01HD060563 and NSF Grant BCS 0924811.


  1. Anderson US, Stoinski TS, Bloomsmith MA, Marr MJ, Smith AD, Maple TL (2005) Relative numerousness judgment and summation in young and old Western Lowland Gorillas. J Comp Psychol 119:285–295. doi: 10.1037/0735-7036.119.3.285 PubMedCrossRefGoogle Scholar
  2. Anderson US, Stoinski TS, Bloomsmith MA, Maple TL (2007) Relative numerousness judgment and summation in Young, middle-aged, and older adult Orangutans (Pongo pygmaeus ablii and Pongo pygmaeus pygmaeus). J Comp Psychol 121:1–11. doi: 10.1007/s10071-008-0185-9 PubMedCrossRefGoogle Scholar
  3. Baker JM, Shivik J, Jordan KE (2011) Tracking of food quantity by coyotes (Canis latrans). Behav Process 88:72–75. doi: 10.1016/j.beproc.2011.08.00 CrossRefGoogle Scholar
  4. Barth H, Kanwisher N, Spelke ES (2003) The construction of large number representations in adults. Cognition 86:201–221. doi: 10.1016/S0010-0277(02)00178-6 PubMedCrossRefGoogle Scholar
  5. Bates LA, Poole JH, Byrne RW (2008) Elephant cognition. Curr Biol 18:R544–R546PubMedCrossRefGoogle Scholar
  6. Benson-Amram S, Heinen VK, Dryer SL, Holekamp KE (2011) Numerical assessment and individual call discrimination by wild spotted hyaenas, Crocuta crocuta. Anim Behav 82:743–752. doi: 10.1016/j.anbehav.2011.07.004 CrossRefGoogle Scholar
  7. Beran MJ (2001) Summation and numerousness judgments of sequentially presented sets of items by chimpanzees (Pan troglodytes). J Comp Psychol 115:181–191. doi: 10.1037/0735-7036.115.2.181 PubMedCrossRefGoogle Scholar
  8. Beran MJ (2007) Rhesus monkeys (Macaca mulatta) enumerate sequentially presented sets of items using analog numerical representations. J Exp Psychol Anim Behav Process 33:42–54. doi: 10.1037/0097-7403.33.1.42 PubMedCrossRefGoogle Scholar
  9. Beran MJ, Taglialatela LA, Flemming TJ, James FM, Washburn DA (2006) Nonverbal estimation during numerosity judgments by adult humans. Q J Exp Psychol 59:2065–2082. doi: 10.1080/17470210600701171 CrossRefGoogle Scholar
  10. Boysen ST, Bernston GG, Mukobi KL (2001) Size matters: impact of item size and quantity on array choice by chimpanzees (Pan troglodytes). J Comp Psychol 115:106–110. doi: 10.1037/0735-7036.115.1.106 PubMedCrossRefGoogle Scholar
  11. Brannon EM (2006) The representation of numerical magnitude. Curr Opin Neurobiol 16:222–229. doi: 10.1016/j.conb.2006.03.002 PubMedCrossRefGoogle Scholar
  12. Brannon EM, Roitman JD (2003) Nonverbal representations of time and number in animals and human infants. In: Meck WH (ed) Functional and neural mechanisms of interval timing. CRC Press, Boca Raton, pp 147–182Google Scholar
  13. Brannon EM, Terrace HS (2000) Representation of the numerosities 1–9 by rhesus macaques (Macaca mulatta). J Exp Psychol Anim Behav Process 26:31–49. doi: 10.1037/0097-7403.26.1.31 PubMedCrossRefGoogle Scholar
  14. Brannon EM, Cantlon JF, Terrace HS (2006) The role of reference points in ordinal numerical comparisons by rhesus macaques (Macaca mulatta). J Exp Psychol Anim Behav Process 32:120–134. doi: 10.1037/0097-7403.32.2.120 PubMedCrossRefGoogle Scholar
  15. Call J (2000) Estimating and operating on discrete quantities in orangutans (Pongo pygmaeus). J Comp Psychol 114:136–147. doi: 10.1037/0735-7036.114.2.136 PubMedCrossRefGoogle Scholar
  16. Cantlon JF, Brannon EM (2006) Shared system for ordering small and large numbers in monkeys and humans. Psychol Sci 17:401–406. doi: 10.1111/j.1467-9280.2006.01719.x PubMedCrossRefGoogle Scholar
  17. Evans TA, Beran MJ, Harris EH, Rice DF (2009) Quantity judgments of sequentially presented food items by capuchin monkeys (Cebus apella). Anim Cogn 12:97–105. doi: 10.1007/s10071-008-0174-z PubMedCrossRefGoogle Scholar
  18. Feigenson L, Carey S, Hauser MD (2002) The representations underlying infants’ choice of more: object files versus analog magnitudes. Psychol Sci 13:150–156. doi: 10.1111/1467-9280.00427 PubMedCrossRefGoogle Scholar
  19. Foerder P, Galloway M, Barthel T, Moore DE III, Reiss D (2011) Insightful problem solving in an Asian Elephant. PLoS ONE 6:e23251Google Scholar
  20. Gaalema DE, Perdue BM, Kelling AS (2011) Food choice, keeper ratings, and reinforcer effectiveness in exotic animals: the value of systematic testing. J Appl Anim Welfare Sci 14:33–41CrossRefGoogle Scholar
  21. Gallistel CR, Gelman R (1992) Preverbal and verbal counting and computation. Cognition 44:43–74. doi: 10.1016/0010-0277(92)90050-R PubMedCrossRefGoogle Scholar
  22. Gallistel CR, Gelman R (2000) Non-verbal numerical cognition: from reals to integers. Trends Cogn Sci 4:59–65. doi: 10.1016/S1364-6613(99)01424-2 PubMedCrossRefGoogle Scholar
  23. Gòmez-Laplaza LM, Gerlai R (2011) Can angelfish (Pterophyllum scalare) count? Discrimination between different shoal sizes follows Weber’s law. Anim Cogn 14:1–9. doi: 10.1007/s10071-010-0337-6 PubMedCrossRefGoogle Scholar
  24. Hart BL, Hart LA, Pinter-Wollman N (2008) Large brains and cognition: where do elephants fit in? Neurosci Biobehav Rev 32:86–98. doi: 10.1016/j.neubiorev.2007.05.012 PubMedCrossRefGoogle Scholar
  25. Hauser MD, Carey S, Hauser LB (2000) Spontaneous number representation in semi-free-ranging rhesus monkeys. Proc R Soc Lond Biol 267:829–833. doi: 10.1098/rspb.2000.1078 CrossRefGoogle Scholar
  26. Huntley-Fenner G (2001) Children’s understanding of number is similar to adults’ and rats’: numerical estimation by 5–7-year-olds. Cognition 78:B27–B40. doi: 10.1016/S0010-0277(00)00122-0 PubMedCrossRefGoogle Scholar
  27. Irie N, Hasegawa T (2012) Summation by Asian Elephants (Elephas maximus). Behav Sci 2:50–56. doi: 10.3390/bs2020050 CrossRefGoogle Scholar
  28. Irie-Sugimoto N, Kobayashi T, Sato T, Hasegawa T (2009) Relative quantity judgment by Asian elephants (Elephas maximus). Anim Cogn 12:193–199. doi: 10.1007/s10071-008-0185-9 PubMedCrossRefGoogle Scholar
  29. Krusche P, Uller C, Dicke U (2010) Quantity discrimination in salamanders. J Exp Biol 213:1822–1828. doi: 10.1242/jeb.039297 PubMedCrossRefGoogle Scholar
  30. Lipton JS, Spelke E (2004) Discrimination of large and small numerosities by human infants. Infancy 5:271–290CrossRefGoogle Scholar
  31. Olthof A, Roberts WA (2000) Summation of symbols by pigeons (Columba livia): the important number and mass of reward items. J Comp Psychol 114:158–166. doi: 10.1037/0735-7036.114.2.158 PubMedCrossRefGoogle Scholar
  32. Perusse R, Rumbaugh DM (1990) Summation in chimpanzees (Pan troglodytes): effects of amounts, number of wells, and finer ratios. Int J Primatol 11:425–437. doi: 10.1007/BF02196130 CrossRefGoogle Scholar
  33. Pisa PE, Agrillo C (2009) Quantity discrimination in felines: a preliminary investigation of the domestic cat (Felis silvestris catus). J Ethol 27:289–293. doi: 10.1007/s10164-008-0121-0 CrossRefGoogle Scholar
  34. Plotnik JM, de Waal FBM, Moore D, Reiss D (2010) Self-recognition in the Asian elephant and future directions for cognition research with elephants in zoological settings. Zoo Biol 29:179–191. doi: 10.1002/zoo.20257 PubMedCrossRefGoogle Scholar
  35. Sulkowski GM, Hauser MD (2001) Can rhesus monkeys spontaneously subtract? Cognition 79:239–262. doi: 10.1016/S0010-0277(00)00112-8 PubMedCrossRefGoogle Scholar
  36. Terrell DF, Thomas RK (1990) Number-related discrimination and summation by squirrel monkeys (Saimiri sciureus sciureus and S. boliviensus boliviensus) on the basis of the number of sides of polygons. J Comp Psychol 104:238–247. doi: 10.1037/0735-7036.104.3.238 PubMedCrossRefGoogle Scholar
  37. Thomas R, Chase L (1980) Relative numerousness judgments by squirrel monkeys. Bull Psychonomic Soc 16:79–82Google Scholar
  38. Uller C, Jaeger R, Guidry G, Martin C (2003) Salamanders (Plethodon cinereus) go for more: rudiments of number in an amphibian. Anim Cogn 6:105–112. doi: 10.1007/s10071-003-0167-x PubMedGoogle Scholar
  39. Ward C, Smuts BB (2007) Quantity-based judgments in the domestic dog (Canis lupus familiaris). Anim Cogn 10:71–80. doi: 10.1007/s10071-006-0042-7 PubMedCrossRefGoogle Scholar

Copyright information

© Springer-Verlag 2012

Authors and Affiliations

  • Bonnie M. Perdue
    • 1
    • 2
  • Catherine F. Talbot
    • 2
  • Adam M. Stone
    • 1
  • Michael J. Beran
    • 2
  1. 1.Zoo AtlantaAtlantaUSA
  2. 2.Language Research CenterGeorgia State UniversityAtlantaUSA

Personalised recommendations