Biology & Philosophy

, 24:215 | Cite as

San Marco and evolutionary biology

  • Alasdair I. Houston


Gould and Lewontin use San Marco, Venice, to criticise the adaptationist program in biology. Following their lead, the architectural term “spandrel” is now widely used in biology to denote a feature that is a necessary byproduct of other aspects of the organism. I review the debate over San Marco and argue that the spandrels are not necessary in the sense originally used by Gould and Lewontin. I conclude that almost all the claims that Gould makes about San Marco are wrong and that it is reasonable to view the architectural spandrel as an adaptation. The spandrels example has not provided a good illustration of why adaptive explanations should be avoided. In fact, it can be used as an example of how adaptive explanations can be dismissed even when there is evidence in their favour. I also discuss the use of the concept of a spandrel in biology.


Adaptation Constraint Mosaic Necessity San Marco Spandrel Symmetry 



I thank Lutz Fromhage, Robert Mark, Andrew Phillips and Sean Rands for helpful comments on previous versions of the ms and Tim Colborn for producing the figures. This work was supported by a Leverhulme Research Fellowship.


  1. Aerts P, Van Damme R, D’Aout K, Van Hooydonck B (2003) Bipedalism in lizards: whole-body modelling reveals a possible spandrel. Philos Trans R Soc Lond B Biol Sci 358:1525–1533. doi: 10.1098/rstb.2003.1342 CrossRefGoogle Scholar
  2. Botha RP (2001) How much of language, if any, came about in the same sort of way as the brooding chamber in snails? Lang Commun 21:225–243. doi: 10.1016/S0271-5309(01)00002-7 CrossRefGoogle Scholar
  3. Buss DM, Haselton MG, Shackelford TK, Bleske AL, Wakefield JC (1998) Adaptations, exaptations, and spandrels. Am Psychol 53:533–548. doi: 10.1037/0003-066X.53.5.533 CrossRefGoogle Scholar
  4. Conway Morris S (1998) The crucible of creation. Oxford University Press, OxfordGoogle Scholar
  5. Creswell KAC (1979) Early muslim architecture. Hacker Art Books, New YorkGoogle Scholar
  6. Crouch DP, Johnson JG (2001) Traditions in architecture. Oxford University Press, New YorkGoogle Scholar
  7. Demus O (1950) The mosaics of Norman Sicily. Routledge & Kegan Paul, LondonGoogle Scholar
  8. Demus O (1984) The mosaics of San Marco in Venice. University of Chicago Press, ChicagoGoogle Scholar
  9. Demus O (1988) The mosaic decoration of San Marco, Venice. University of Chicago Press, ChicagoGoogle Scholar
  10. Dennett DC (1995) Darwin’s dangerous idea. Simon & Schuster, New YorkGoogle Scholar
  11. De Sousa R (2004) Is art an adaptation? Prospects for an evolutionary perspective on beauty. J Aesth Art Crit 62:109–118. doi: 10.1111/j.1540-594X.2004.00144.x CrossRefGoogle Scholar
  12. Dupre J (2001) Human nature and the limits of science. Oxford University Press, OxfordCrossRefGoogle Scholar
  13. Gazzaniga MS (1994) Nature’s mind. Penguin, HammondsworthGoogle Scholar
  14. Gould SJ (1997) The exaptive excellence of spandrels as a term and prototype. Proc Natl Acad Sci USA 94:10750–10755. doi: 10.1073/pnas.94.20.10750 CrossRefGoogle Scholar
  15. Gould SJ (2002) The structure of evolutionary theory. Harvard University Press, CambridgeGoogle Scholar
  16. Gould SJ, Lewontin RC (1979) The spandrels of San Marco and the panglossian paradigm: a critique of the adaptationist programme. Proc R Soc Lond B Biol Sci 205:581–598CrossRefGoogle Scholar
  17. Gould SJ, Vrba ES (1982) Exaptation—a missing term in the science of form. Paleobiology 8:4–15Google Scholar
  18. Hamilton JA (1933) Byzantine architecture and decoration. Batsford, LondonGoogle Scholar
  19. Hampton SJ (2004) Domain mismatches, scruffy engineering, exaptations and spandrels. Theory Psychol 14:147–166. doi: 10.1177/0959354304042014 CrossRefGoogle Scholar
  20. Houston AI (1997) Are the spandrels of San Marco really panglossian pendentives? Trends Ecol Evol 12:125. doi: 10.1016/S0169-5347(96)20112-0 CrossRefGoogle Scholar
  21. Howard D (2002) The architecture of Venice. Yale University Press, New HavenGoogle Scholar
  22. Jackendoff R, Lerdahl F (2006) The capacity for music: what is it, and what’s special about it? Cognition 100:33–72. doi: 10.1016/j.cognition.2005.11.005 CrossRefGoogle Scholar
  23. Kraus H (1979) Gold was the mortar: the economics of cathedral building. Routledge & Kegan Paul, BostonGoogle Scholar
  24. Landweber LF, Pokrovskaya ID (1999) Emergence of a dual-catalytic RNA with metal-specific cleavage and ligase activities: the spandrels of RNA evolution. Proc Natl Acad Sci USA 96:173–178. doi: 10.1073/pnas.96.1.173 CrossRefGoogle Scholar
  25. Lassus J (1966) The early christian and Byzantine world: landmarks of the world’s art. Paul Hamlyn, LondonGoogle Scholar
  26. Lauder GV (1996) The argument from design. In: Rose MR, Lauder GV (eds) Adaptation. Academic Press, New York, pp 55–91Google Scholar
  27. Mainstone RJ (1999) Structure in architecture: history, design and innovation: variorum collected studies series, v. CS659. Ashgate, AldershotGoogle Scholar
  28. Mark R (1982) Experiments in gothic structure. MIT, CambridgeGoogle Scholar
  29. Mark R (1990) Light, wind, and structure. MIT, CambridgeGoogle Scholar
  30. Mark R (1996) Architecture and evolution. Am Sci 84:383–389Google Scholar
  31. Mayr E (1983) How to carry out the adaptationist program. Am Nat 121:324–334. doi: 10.1086/284064 CrossRefGoogle Scholar
  32. Nuttegns P (1983) The story of architecture. Phaidon, OxfordGoogle Scholar
  33. Ousterhout R (1992) Originality in Byzantine architecture + the Chois Katholikon − the Case of Nea-Moni. J Soc Archit Hist 51:48–60. doi: 10.2307/990640 CrossRefGoogle Scholar
  34. Ousterhout R (1999) Master builders of Byzantium. Princeton University Press, PrincetonGoogle Scholar
  35. Peretz I (2006) The nature of music from a biological perspective. Cognition 100:1–32. doi: 10.1016/j.cognition.2005.11.004 CrossRefGoogle Scholar
  36. Pinker S (1994) The language instinct. Penguin, LondonGoogle Scholar
  37. Pinker S, Bloom P (1990) Natural language and natural selection. Behav Brain Sci 13:707–784Google Scholar
  38. Rose S (1997) Lifelines. Allen Lane, LondonGoogle Scholar
  39. Schacter DL, Dodson CS (2001) Misattribution, false recognition and the sins of memory. Philos Trans R Soc Lond B Biol Sci 356:1385–1393. doi: 10.1098/rstb.2001.0938 CrossRefGoogle Scholar
  40. Segerstråle U (2000) Defenders of the truth. Oxford University Press, OxfordGoogle Scholar
  41. Seilacher A (1972) Divaricate patterns in pelecypod shells. Lethaia 5:325–343. doi: 10.1111/j.1502-3931.1972.tb00862.x CrossRefGoogle Scholar
  42. Seilacher A (1973) Fabricational noise in adaptive morphology. Syst Zool 22:451–465. doi: 10.2307/2412952 CrossRefGoogle Scholar
  43. Selzer J (1993) Understanding scientific prose. University of Wisconsin Press, MadisonGoogle Scholar
  44. Shubnikov AV, Koptsik VA (1974) Symmetry in science and art. Plenum, New YorkGoogle Scholar
  45. Sole RV, Valverde S (2006) Are network motifs the spandrels of cellular complexity? Trends Ecol Evol 21:419–422. doi: 10.1016/j.tree.2006.05.013 CrossRefGoogle Scholar
  46. Stephens DW, Krebs JR (1986) Foraging theory. Princeton University Press, PrincetonGoogle Scholar
  47. Stevens PS (1980) Handbook of regular patterns. MIT, CambridgeGoogle Scholar
  48. Tenner E (1996) Why things bite back. Forth Estate, LondonGoogle Scholar
  49. Turnbull D (2000) Gothic tales of spandrels, hooks and monsters: complexity and association in the explanation of technological change. In: Ziman J (ed) Technological innovation as an evolutionary process. Cambridge University Press, CambridgeGoogle Scholar
  50. Van Leeuwen DSN (2007) The spandrels of self-deception: prospects for a biological theory of a mental phenomenon. Philos Psychol 20:329–348. doi: 10.1080/09515080701197148 CrossRefGoogle Scholar
  51. Weiss MA, Nakagawa SH, Jia WH, Xu B, Hua QX, Chu YC, Wang RY et al (2002) Protein structure and the spandrels of San Marco: insulin’s receptor-binding surface is buttressed by an invariant leucine essential for its stability. Biochemistry 41:809–819. doi: 10.1021/bi011839+ CrossRefGoogle Scholar
  52. Williams GC (1966) Adaptation and natural selection: a critique of some current evolutionary thought. Princeton University Press, PrincetonGoogle Scholar
  53. Williams GC (1992) Natural selection: domains, levels, and challenges. Oxford University Press, OxfordGoogle Scholar
  54. Willis D (1995) The sand dollar and the slide rule. Adison-Wesley, ReadingGoogle Scholar
  55. Wilson DS (2002) Darwin’s cathedral. University of Chicago, ChicagoGoogle Scholar

Copyright information

© Springer Science+Business Media B.V. 2008

Authors and Affiliations

  1. 1.School of Biological SciencesUniversity of BristolBristolUK

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