Animal Cognition

, Volume 15, Issue 4, pp 699–710 | Cite as

The role of numerical competence in a specialized predatory strategy of an araneophagic spider

Original Paper


Although a wide range of vertebrates have been considered in research on numerical competence, little is known about the role of number-related decisions in the predatory strategies of invertebrates. Here, we investigate how numerical competence is expressed in a highly specialized predatory strategy adopted by the small juveniles of Portiaafricana when practicing communal predation, with the prey being another spider, Oecobius amboseli. Two or more P. africana juveniles sometimes settle by the same oecobiid nest and then share the meal after one individual captures the oecobiid. Experiments were designed to clarify how these predators use number-related cues in conjunction with non-numerical cues when deciding whether to settle at a nest. We used lures (dead spiders positioned in lifelike posture) arranged in a series of 24 different scenes defined by the type, configuration and especially number of lures. On the whole, our findings suggest that P. africana juveniles base settling decisions on the specific number of already settled conspecific juveniles at the nest and express a preference for settling when the number is one instead of zero, two or three. By varying the size of the already settled juveniles and their positions around the nest, we show that factors related to continuous variables and stimulus configuration are unlikely explanations for our findings.


Numerosity discrimination Numerical ability Prototype matching Communal predation Araneophagy Salticidae 


  1. Agrillo C, Dadda M, Serena G (2008) Do fish count? Spontaneous discrimination of quantity in female mosquitofish. Anim Cogn 11:495–503PubMedCrossRefGoogle Scholar
  2. Anderson DE, Vogel EK, Awh E (2011) Precision in visual working memory reaches a stable plateau when individual item limits are exceeded. J Neurosci 31:1128–1138PubMedCrossRefGoogle Scholar
  3. Awh E, Barton B, Vogel EK (2007) Visual working memory represents a fixed number of items regardless of complexity. Psychol Sci 18:622–628PubMedCrossRefGoogle Scholar
  4. Baddeley A (1986) Working memory. Oxford University Press, OxfordGoogle Scholar
  5. Baddeley A (2003) Working memory: looking back and looking forward. Nat Rev Neurosci 4:829–839PubMedCrossRefGoogle 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 82:743–752CrossRefGoogle Scholar
  7. Beran MJ (2001) Summation and numerousness judgments of sequentially presented sets of items by chimpanzees (Pan troglodytes). J Comp Psychol 115:181–191PubMedCrossRefGoogle Scholar
  8. Beran MJ (2008) Capuchin monkeys (Cebus apella) succeed in a test of quantity conservation. Anim Cogn 11:109–116PubMedCrossRefGoogle Scholar
  9. Beran MJ (2010) Chimpanzees (Pan troglodytes) accurately compare poured liquid quantities. Anim Cogn 13:641–649PubMedCrossRefGoogle Scholar
  10. Beran MJ, Evans TA, Harris EH (2008) Perception of food amounts by chimpanzees based on the number, size, contour length and visibility of items. Anim Behav 75:1793–1802PubMedCrossRefGoogle Scholar
  11. Boysen ST, Berntson GG (1989) Numerical competence in a chimpanzee (Pan troglodytes). J Comp Psychol 103:23–31PubMedCrossRefGoogle Scholar
  12. Boysen ST, Berntson GG (1995) Responses to quantity: perceptual versus cognitive mechanisms in chimpanzees (Pan troglodytes). J Exp Psychol Anim Behav Process 21:82–86PubMedCrossRefGoogle Scholar
  13. Brady TF, Konkle T, Alvarez GA (2011) A review of visual memory capacity: beyond individual items and toward structured representations. J Vis 11:1–34CrossRefGoogle Scholar
  14. Brannon EM (2002) The development of ordinal numerical knowledge in infancy. Cognition 83:223–240PubMedCrossRefGoogle Scholar
  15. Brannon EM (2006) The representation of numerical magnitude. Curr Opin Neurobiol 16:222–229PubMedCrossRefGoogle Scholar
  16. Brannon EM, Abbott S, Lutz DJ (2004) Number bias for the discrimination of large visual sets in infancy. Cognition 93:B59–B68PubMedCrossRefGoogle Scholar
  17. Breukelaar JWC, Dalrymple-Alford JC (1998) Timing ability and numerical competence in rats. J Exp Psychol 24:84–97Google Scholar
  18. Burr DC, Ross J (2008) A visual sense of number. Curr Biol 18:425–428PubMedCrossRefGoogle Scholar
  19. Burr DC, Turi M, Anobile G (2010) Subitizing but not estimation of numerosity requires attentional resources. J Vis 10(6):1–10CrossRefGoogle Scholar
  20. Butterworth B (1999) A head for figures. Science 284:928–929PubMedCrossRefGoogle Scholar
  21. Butterworth B (2008) Numerosity perception: how many speckles on the hen? Curr Biol 16:R388–R389CrossRefGoogle Scholar
  22. Cantlon JF, Brannon EM (2007) Basic math in monkeys and college students. PLoS ONE 5:e328Google Scholar
  23. Carazo P, Font E, Forteza-Behrendt E (2009) Quantity discrimination in Tenebrio molitor: evidence of numerosity discrimination in an invertebrate? Anim Cogn 12:463–470PubMedCrossRefGoogle Scholar
  24. Carey S (1998) Knowledge of number: its evolution and ontogeny. Science 282:641–642PubMedCrossRefGoogle Scholar
  25. Carey S (2001) Cognitive foundations of arithmetic: evolution and ontogenesis. Mind Lang 16:37–55Google Scholar
  26. Castelli F, Glaser DE, Butterworth B (2006) Discrete and analogue quantity processing in the parietal lobe: a functional MRI study. Proc Nat Acad Sci USA 103:4693–4698PubMedCrossRefGoogle Scholar
  27. Charnov EL (1976) Optimal foraging, the marginal value theorem. Theor Popul Biol 9:129–136PubMedCrossRefGoogle Scholar
  28. Chittka L, Geiger K (1995) Can honey-bees count landmarks? Anim Behav 49:159–164CrossRefGoogle Scholar
  29. Church RM, Broadbent HA (1990) Alternative representations of time, number, and rate. Cognition 37:55–81PubMedCrossRefGoogle Scholar
  30. Clearfield MW, Mix KS (1999) Number versus contour length in infants’ discrimination of small visual sets. Psychol Sci 10:408–411CrossRefGoogle Scholar
  31. Cowan N (1998) Visual and auditory working memory capacity. Trends Cogn Sci 2:77–78PubMedCrossRefGoogle Scholar
  32. Cowan N (2000) The magical number 4 in short-term memory: a reconsideration of mental storage capacity. Behav Brain Sci 24:87–185CrossRefGoogle Scholar
  33. Cowan N (2010) The magical mystery four: how is working memory capacity limited, and why? Curr Dir Psychol Sci 19:51–57PubMedCrossRefGoogle Scholar
  34. Curio E (1976) The ethology of predation. Springer, BerlinGoogle Scholar
  35. Dacke M, Srinivasan MV (2008) Evidence for counting in insects. Anim Cogn 11:683–689PubMedCrossRefGoogle Scholar
  36. Dadda M, Piffer L, Agrillo C, Biazza A (2009) Spontaneous number representation in mosquitofish. Cognition 112:343–348PubMedCrossRefGoogle Scholar
  37. Davis H, Memmott J (1982) Counting behavior in animals: a critical evaluation. Psychol Bull 92:547–571CrossRefGoogle Scholar
  38. Davis H, MacKenzie KA, Morrison S (1989) Numerical discrimination by rats (Rattus norvegicus) using body and vibrissal touch. J Comp Psychol 103:45–53CrossRefGoogle Scholar
  39. Dehaene S (1992) Varieties of numerical abilities. Cognition 44:1–42PubMedCrossRefGoogle Scholar
  40. Dehaene S (2001) Precis of the number sense. Mind Lang 16:16–36Google Scholar
  41. Feigenson L, Dehaene S, Spelke E (2004) Core systems of number. Trends Cogn Sci 8:307–314PubMedCrossRefGoogle Scholar
  42. Flombaum JI, Junge JA, Hauser MD (2005) Rhesus monkeys (Macaca mulatta) spontaneously compute addition operations over large numbers. Cognition 97:315–325PubMedCrossRefGoogle Scholar
  43. Fougnie D, Marois R (2011) Distinct capacity limits for attention and working memory. Psychol Sci 17:526–534CrossRefGoogle Scholar
  44. Franks NR, Dornhaus A, Metherell BG, Nelson TR, Lanfear SAJ, Symes WS (2006) Not everything that counts can be counted: ants use multiple metrics for a single nest trait. Proc R Soc B 273:165–169PubMedCrossRefGoogle Scholar
  45. Gallistel CR, Gelman R (2000) Non-verbal numerical cognition: from reals to integers. Trends Cogn Sci 4:59–65PubMedCrossRefGoogle Scholar
  46. Gelman R, Gallistel CR (1978) The child’s understanding of number. Harvard University Press, Cambridge, MAGoogle Scholar
  47. Giaquinto M (2001) What cognitive systems underlie arithmetical abilities? Mind Lang 16:56–68Google Scholar
  48. Gross HJ, Pahl M, Si A, Zhu H, Tautz J, Zhang S (2009) Number-based visual generalisation in the honeybee. PLoS ONE 4(1):e4263PubMedCrossRefGoogle Scholar
  49. Hager MC, Helfman GS (1991) Safety in numbers: shoal size choice by minnows under predatory threat. Behav Ecol Sociobiol 29:271–276CrossRefGoogle Scholar
  50. Hanus D, Call J (2007) Discrete quantity judgments in the great apes (Pan paniscus, Pan troglodytes, Gorilla gorilla, Pongo pygmaeus): the effect of presenting whole sets versus item-by-item. J Comp Psychol 121:241–249PubMedCrossRefGoogle Scholar
  51. Harland DP, Jackson RR (2001) Prey classification by Portia fimbriata, a salticid spider that specializes at preying on other salticids: species that elicit cryptic stalking. J Zool 255:445–460CrossRefGoogle Scholar
  52. Haun DBM, Jordan FM, Vallortigara G, Clayton NS (2010) Origins of spatial, temporal and numerical cognition: insights from comparative psychology. Trends Cog Sci 14:552–560CrossRefGoogle Scholar
  53. Hauser MD, Carey S (2003) Spontaneous representations of small numbers of objects by rhesus macaques: examination of content and format. Cognit Psychol 47:367–491PubMedCrossRefGoogle Scholar
  54. Hoare DJ, Couzin ID, Godin GJ, Krause J (2004) Context-dependent group size choice in fish. Anim Behav 67:155–164CrossRefGoogle Scholar
  55. Hunt S, Low J, Burns KC (2008) Adaptive numerical competency in a food-hoarding songbird. Proc R Soc B 275:2373–2379PubMedCrossRefGoogle Scholar
  56. Hurewitz F, Gelman R, Schnitzer B (2006) Sometimes area counts more than number. Proc Nat Acad Sci USA 103:19599–19604PubMedCrossRefGoogle Scholar
  57. Jackson RR, Cross FR (2011) Spider cognition. Adv Insect Physiol 41:115–174CrossRefGoogle Scholar
  58. Jackson RR, Nelson XJ (2012) Attending to detail by communal spider-eating spiders. Anim Cogn (in press). doi:10.1007/s10071-012-0469-y
  59. Jackson RR, Pollard SD, Salm K (2008) Observations of Portia africana, an araneophagic jumping spider, living together and sharing prey. N Z J Zool 35:237–242CrossRefGoogle Scholar
  60. Jakob EM, Christa D, Skow CD, Long S (2011) Plasticity, learning and cognition. In: Herberstein ME (ed) Spider behaviour: flexibility and versatility. Cambridge University Press, New York, pp 307–347Google Scholar
  61. Kahneman D, Treisman A, Gibbs BJ (1992) The reviewing of object files: object-specific integration of information. Cognit Psychol 24:175–219PubMedCrossRefGoogle Scholar
  62. Kaufman EL, Lord MW, Reese TW, Volkmann J (1949) The discrimination of visual number. Am J Psychol 62:498–525PubMedCrossRefGoogle Scholar
  63. Kitchen DM (2006) Alpha male black howler monkey responses to loud calls: effect of numeric odds, male companion behaviour and reproductive investment. Anim Behav 67:125–139CrossRefGoogle Scholar
  64. Kobayashi T, Hiraki K, Mugitani R, Hasegawa T (2004) Baby arithmetic: one object plus one tone. Cognition 91:B23–B34PubMedCrossRefGoogle Scholar
  65. Krusche P, Uller C, Dicke U (2010) Quantity discrimination in salamanders. J Exp Biol 213:1822–1828PubMedCrossRefGoogle Scholar
  66. Le Corre M, Carey S (2008) Why the verbal counting principles are constructed out of representations of small sets of individuals: a reply to Gallistel. Cognition 107:650–662PubMedCrossRefGoogle Scholar
  67. Lipton JS, Spelke ES (2003) Origins of number sense: large-number discrimination in human infants. Psychol Sci 14:396–401PubMedCrossRefGoogle Scholar
  68. Lyon BE (2003) Egg recognition and counting reduce costs of avian conspecific brood parasitism. Nature 422:495–499PubMedCrossRefGoogle Scholar
  69. Mandler G, Shebo BJ (1982) Subitizing: an analysis of its conceptual processes. J Exp Psychol 111:1–22Google Scholar
  70. Matsuzawa T (1985) Use of numbers by a chimpanzee. Nature 315:57–59PubMedCrossRefGoogle Scholar
  71. McComb K, Packer C, Pusey A (1994) Roaring and numerical assessment in contests between groups of lions, Panthera leo. Anim Behav 47:379–387CrossRefGoogle Scholar
  72. McCrink K, Wynn K (2004) Large-number addition and subtraction by 9-month-old infants. Psychol Sci 15:776–781PubMedCrossRefGoogle Scholar
  73. Meck WH, Church RM (1983) A mode control model of counting and timing processes. J Exp Psychol Anim Behav Process 9:320–334PubMedCrossRefGoogle Scholar
  74. Nelson XJ, Jackson RR (2011) Flexibility in the foraging strategies of spiders. In: Herberstein ME (ed) Spider behaviour: flexibility and versatility. Cambridge University Press, New York, pp 31–56Google Scholar
  75. Owen AM (2004) Working memory: imaging the magic number four. Curr Biol 14:R573–R574PubMedCrossRefGoogle Scholar
  76. Pepperberg IM (2006) Grey parrot numerical competence: a review. Anim Cogn 9:377–391PubMedCrossRefGoogle Scholar
  77. Piazza M, Mechelli A, Butterworth B, Price CJ (2002) Are subitizing and counting implemented as separate or functionally overlapping processes? NeuroImage 15:435–446PubMedCrossRefGoogle Scholar
  78. Piazza M, Fumarola A, Chinello A, Melcher D (2011) Subitizing reflects visuo-spatial object individuation capacity. Cognition 121:147–153PubMedCrossRefGoogle Scholar
  79. Pyke GH, Pulliam HR, Charnov EL (1977) Optimal foraging: a selective review of theory and tests. Q Rev Biol 52:137–154CrossRefGoogle Scholar
  80. Pylyshyn ZW (1989) The role of location indexes in spatial perception: a sketch of the FINST spatial-index model. Cognition 32:65–97PubMedCrossRefGoogle Scholar
  81. Pylyshyn ZW (2001) Visual indexes, preconceptual objects, and situated vision. Cognition 80:127–158PubMedCrossRefGoogle Scholar
  82. Railo H, Koivisto M, Revonsuo A, Hannula MM (2008) The role of attention in subitizing. Cognition 107:82–104PubMedCrossRefGoogle Scholar
  83. Reznikova Z, Ryabko B (2011) Numerical competence in animals, with an insight from ants. Behaviour 148:405–434CrossRefGoogle Scholar
  84. Riggs KJ, Ferrand L, Lancelin D, Fryziel L, Dumur G (2006) Subitizing in tactile perception. Psychol Sci 17:271–272PubMedCrossRefGoogle Scholar
  85. Rodríguez RL, Gamboa E (2000) Memory of captured prey in three web spiders (Araneae: Araneidae, Linyphiidae, Tetragnathidae). Anim Cogn 3:91–97CrossRefGoogle Scholar
  86. Rodríguez RL, Gloudeman MD (2011) Estimating the repeatability of memories of captured prey formed by Frontinella communis spiders (Araneae: Linyphiidae). Anim Cogn 14:675–682PubMedCrossRefGoogle Scholar
  87. Rosch E (1975) Cognitive representations of semantic categories. J Exp Psychol 104:192–233Google Scholar
  88. Rosch E, Mervis CB, Greay WD, Jonhson DM, Boyes-Braem P (1976) Basic objects in natural categories. Cognit Psychol 8:382–429CrossRefGoogle Scholar
  89. Schmitt V, Fischer J (2011) Representational format determines numerical competence in monkeys. Nat Commun 2:257. doi:10.1038/ncomms1262 PubMedCrossRefGoogle Scholar
  90. Seron X, Pesenti M (2001) The number sense theory needs more empirical evidence. Mind Lang 16:76–88Google Scholar
  91. Shettleworth SJ (2009) Cognition, evolution, and behaviour, 2nd edn. Oxford University Press, New YorkGoogle Scholar
  92. Slaughter V, Itakura S, Kutsuki A, Siegal M (2011) Learning to count begins in infancy: evidence from 18 month olds’ visual preferences. Proc R Soc B 278:2979–2984PubMedCrossRefGoogle Scholar
  93. Starkey P, Spelke ES, Gelman R (1983) Detection of intermodal numerical correspondences by human infants. Science 222:179–181PubMedCrossRefGoogle Scholar
  94. Stephens DW, Krebs JR (1986) Foraging theory. Princeton University Press, PrincetonGoogle Scholar
  95. Tardif SD, Layne DG, Smucny DA (2002) Can marmoset mothers count to three? Effect of litter size on mother-infant interactions. Ethology 108:825–836CrossRefGoogle Scholar
  96. 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 104:238–247PubMedCrossRefGoogle Scholar
  97. Thomas RK, Simmons LW (2010) Cuticular hydrocarbons influence female attractiveness to males in the Australian field cricket, Teleogryllus oceanicus. J Evol Biol 23:707–714PubMedCrossRefGoogle Scholar
  98. Tomonaga M (2008) Relative numerosity discrimination by chimpanzees (Pan troglodytes): evidence for approximate numerical representations. Anim Cogn 11:43–57PubMedCrossRefGoogle Scholar
  99. Treisman AM, Gelade G (1980) A feature integration theory of attention. Cogn Psychol 12:97–136PubMedCrossRefGoogle Scholar
  100. Trick LM (2005) The role of working memory in spatial enumeration: patterns of selective interference in subitizing and counting. Psychon Bull Rev 12:675–681PubMedCrossRefGoogle Scholar
  101. Trick LM, Pylyshyn ZW (1994) Why are small and large numbers enumerated differently? A limited-capacity preattentive stage in vision. Psychol Rev 101:80–102PubMedCrossRefGoogle Scholar
  102. Uller C, Hauser MD, Carey S (2001) Spontaneous representation of number in cotton-top tamarins. J Comp Psychol 115:1–10CrossRefGoogle Scholar
  103. Uller C, Jaeger R, Guidry G, Martin C (2003) Salamanders (Plethodon cinereus) go for more: rudiments of number in a species of basal vertebrate. Anim Cogn 6:105–112PubMedGoogle Scholar
  104. Wesley F (1961) The number concept: a phylogenetic review. Psychol Bull 53:420–428CrossRefGoogle Scholar
  105. Wood JN, Hauser MC, Glynn DD, Barner D (2007) Free-ranging rhesus monkeys spontaneously individuate and enumerate small numbers of non-solid portions. Cognition 106:207–221PubMedCrossRefGoogle Scholar
  106. Wynn K (1992) Addition and subtraction by human infants. Nature 358:749–750PubMedCrossRefGoogle Scholar
  107. Wynn K (1998) Psychological foundations of number: numerical competence in human infants. Trends Cogn Sci 2:296–303PubMedCrossRefGoogle Scholar
  108. Xu Y, Chun MM (2009) Selecting and perceiving multiple visual objects. Trends Cogn Sci 13:167–174PubMedCrossRefGoogle Scholar
  109. Xu Y, Spelke ES (2000) Large number discrimination in 6-month-old infants. Cognition 74:B1–B11PubMedCrossRefGoogle Scholar

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© Springer-Verlag 2012

Authors and Affiliations

  1. 1.School of Biological SciencesUniversity of CanterburyChristchurchNew Zealand
  2. 2.International Centre of Insect Physiology and EcologyMbita PointKenya

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