Assembly rules


Community assembly rules are a set of controversial rules in ecology, first proposed by Jared Diamond.[1]


The rules were developed after more than a decade of research into the avian assemblages on islands near New Guinea. The rules assert that competition is responsible for determining the patterns of assemblage composition.

Rule 1: Forbidden species combinationsEdit

The first rule is "forbidden species combinations". Diamond's hypothesis was that competition, not random immigration, was the main force structuring the species composition of islands.

So for example, the Bismarck black myzomela (Myzomela pammelaena) excludes the black sunbird (Nectarinia sericea). The Bismarck black myzomela lives on 23 of the 41 surveyed islands in the Bismarck Archipelago, but not on any of the 14 islands inhabited by the black sunbird. The two birds are about the same size, and both use their curved bills to sip nectar; Diamond argued that competition affects their distribution.[2]

Rule 2: Reduced niche overlapEdit

Case tested the assembly rule that species occurring together on islands should have less niche overlap than random assemblages because they have undergone specialization.[3] His study measured niche overlap of lizards on 37 islands near Baja California and compared niche overlap to the median niche overlap of computer generated random species assemblages. Case found that 30 of the 37 islands had lower niche overlap than the random assemblages and that some of the competition is due to interspecific competition.

Testing and alternative theoriesEdit

Testing the assembly rules is a complex process that often uses computer simulations to compare experimental data with characteristics of random assemblages of species. The rules are generally regarded as hypotheses that need to be tested on an individual basis, not as accepted conclusions. This is the reason why Diamond's results sparked nearly two decades worth of controversy in the literature, from the late seventies through the late nineties and is considered a turning point in community ecology.[4] Daniel Simberloff led the arguments against these rules claiming that theory as the one developed by Diamond "has generated predictions that are either practically untestable, by virtue of unmeasurable parameters or unrealizable assumptions, or trivially true".[4] The controversy was developed in a number of scientific papers (see for instance [5] and [6]) and contributed to the development of null[7] and neutral models in community ecology, which are nowadays widely used to test the significance of ecological patterns.[8]


  1. ^ Cody ML, Diamond JM, ed. (1975). Ecology and Evolution of Communities. Cambridge, MA: Belknap Press, Harvard University Press. pp. 342–444.
  2. ^ Erik Stokstad (2009) 'On the Origin of Ecological Structure', Science 2 Oct 2009 pp. 33–35.
  3. ^ Case, Ted (1983). "Niche overlap and the assembly of island lizard communities". Oikos. 41 (3): 427–433. doi:10.2307/3544102. JSTOR 3544102.
  4. ^ a b Lewin, Roger (1983). "Santa Rosalia was a goat". Science. 221 (4611): 636–639. JSTOR 1691178.
  5. ^ Simberloff, Daniel (1978). "Using island biogeographic distributions to determine if colonization is stochastic". The American Naturalist. 112 (986): 713–726. doi:10.1086/283313.
  6. ^ Gilpin, Michael; Diamond, Jared (1982). "actors contributing to non-randomness in species co-occurrences on islands". Oecologia. 52 (1): 75–84. Bibcode:1982Oecol..52...75G. doi:10.1007/BF00349014. PMID 28310111. S2CID 22204293.
  7. ^ Gotelli (1999). "ECOLOGY: How Do Communities Come Together?". Science. 286 (5445): 1684–1685. doi:10.1126/science.286.5445.1684a. S2CID 152996619.
  8. ^ Gotelli, N. J.; McGill, B. J. (2006). "Null versus neutral models: what's the difference?". Ecography. 29 (5): 793–800. doi:10.1111/j.2006.0906-7590.04714.x.

Further readingEdit