The spitting behavior of two species of spitting cobras


Spitting cobras defend themselves by spitting their venom in the face of a harasser. Although it is common belief that spitting cobras direct their venom at the eyes of an aggressor, this has never been investigated. Here, we show that the spitting act of cobras (Naja nigricollis and N. pallida) can readily be triggered by a moving human face or by a moving real size photo of a human face. In contrast, a stationary human face (real or photo) or a moving or stationary human hand does not trigger the spitting act. If threatened, spitting cobras aim their venom, ejected either in two distinct jets (N. pallida) or in a fine spray (N. nigricollis), either between the eyes or at one eye. In both cobra species investigated, the width and height of the area hit by the venom was independent of eye distance (test range 5.5 cm and 11 cm). During the spitting act the cobras performed fast undulating head movements that lead to a larger distribution of their venom. This behavior increases the probability that at least one eye of the aggressor is hit.

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  1. Bogert CM (1943) Dentitional phenomena in cobras and other elapids, with notes on adaptive modification of fangs. Bull Am Museum Nat Hist 81:285–360

  2. Broadley DG (1968) A review of the African cobras of the genus Naja (Serpentes: Elapinae). Arnoldia Rhod 3:1–14

  3. Broadley DG (1974) A review of the cobras of the Naja nigricollis complex in southwestern Africa (Serpentes: Elapidae). Cimbebasia 2:155–162

  4. Bullock TH, Diecke FPJ (1956) Properties of an infra-red receptor. Science 124:281–282

  5. Carpenter CC, Ferguson GW (1977) Variation and evolution of stereotyped behavior in reptiles. In: Gans C, Tinkle DW (eds) Biology of the reptilia 7. Academic, New York, pp 335–554

  6. Cascardi J, Young BA, Husic HD, Sherma J (1999) Protein variation in the venom spat by the red spitting cobra, Naja pallida (Reptilia: Serpentes). Toxicon 37:1271–1279

  7. Freyvogel T, Honegger C (1965) Der ‘Speiakt‘ von Naja nigricollis. Acta Trop 22:289–302

  8. Goring J (1900) Can a cobra eject its venom? J Bombay Nat Hist Soc 13:376

  9. Greene H (1988) Antipredator mechanisms in reptiles. In: Gans C, Huey RB (eds) Biology of the reptilia 16. Academic, New York, pp 1–152

  10. Greene H, Fogden M, Fogden P (1997) Snakes: the evolution of mystery in nature. University of California Press, Berkeley

  11. Ismail M, al-Bekairi AM, el-Bedaiwy AM, Abd-el Salam MA (1993a) The ocular effects of spitting cobras: I. The ringhals cobra (Hemachatus haemachatus) venom-induced corneal opacification syndrome. J Toxicol Clin Toxicol 31:31–41

  12. Ismail M, al-Bekairi AM, el-Bedaiwy AM, Abd-el Salam MA (1993b) The ocular effects of spitting cobras: II. Evidence that cardiotoxins are responsible for the corneal opacification syndrome. J Toxicol Clin Toxicol 31:45–62

  13. Kardong VK, Keine TL, Johnson EK (1997) Proximate factors affecting the predatory behavior of the red spitting cobra, Naja mossambica pallida. J Herpetol 31:66–71

  14. Mattison C (1995) The encyclopedia of snakes. Blandford, London

  15. Rasmussen S., Young B, Krimm H (1995) On the ‘spitting‘ behavior in cobras (Serpentes: Elapidae). J Zool 237:27–35

  16. Slowinski JB, Wüster W (2000) A new cobra (Elapidae: Naja) from Myanmar (Burma). Herpetol 56:257–270

  17. Slowinski JB, Knight A, Rooney A (1997) Inferring species trees from gene trees: a phylogenetic analysis of the Elapidae (serpentes) based on the amino acid sequences venom protein. Mol Phylogenet Evol 8:349–362

  18. Spawls S, Branch B (1995) The dangerous snakes of Africa. Blandford, London

  19. Spawls S, Howell K, Drewes R, Ashe J (2002) A field guide to the reptiles of East Africa. Academic, London

  20. Sweeney RCH (1971) Snakes of Nyasaland. A. Asher and Co, Amsterdam

  21. Warrell D, Ormerod L (1976) Snake venom ophthalmia and blindness caused by the spitting cobra (Naja nigricollis) in Nigeria. Am J Trop Med Hyg 25:525–529

  22. Wüster W, Broadley DG (2003) A new species of spitting cobra from northeastern Africa (Serpentes: Elapidae: Naja). J Zool 259:345–359

  23. Wüster W, Thorpe RS (1992) Dentitional phenomena in cobras revisited: spitting and fang structure in the Asiatic species of Naja (Serpentes: Elapidae). Herpetol 48:424–434

  24. Young BA, Dunlap K, Koenig K, Singer M (2004) The buccal buckle: the functional morphology of venom in spitting cobras. J Exp Biol 207:3483–3494

  25. Zug GR, Vitt LJ, Caldwell JP (2001) Herpetology. Academic, San Diego

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We would like to acknowledge Boris Chagnaud for his technical help with the high-speed video filming and Michael Hofmann for his help with the analysis of spitting pattern centres. We thank Wolfgang Alt and Gabriele Uhl for help with statistics. We also thank Joachim Mogdans and two anonymous referees for critically reading the manuscript and for many helpful suggestions. This study was supported by the German Science Foundation (DFG). The experiments comply with the ``Principles of animal care'', publication No. 86–23, revised 1985 of the National Institute of Health, and also with the current laws of Germany.

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Correspondence to G. Westhoff.

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Westhoff, G., Tzschätzsch, K. & Bleckmann, H. The spitting behavior of two species of spitting cobras. J Comp Physiol A 191, 873–881 (2005) doi:10.1007/s00359-005-0010-8

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  • Snake
  • Behavior
  • Reptile
  • Defensive behavior
  • Venom