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The final step in food web construction is the generation of species-level networks (SLNs). A SLN is considered here a single potential pattern of community interactions in any given place at an instant of time, and may be constructed in two distinctly different ways. First, using empirical observations, one could construct the SLN of a community. This is typically the fashion in which SLNs are reconstructed for modern communities; workers observe and record the community’s trophic interactions. SLNs of this type are precise and without error, though usually taxonomically incomplete, and we cannot have similar confidence in their accuracies because of the sources of uncertainty described above. Capturing their variability requires repeated observations, which is possible under some circumstances. For example, there has been documentation of seasonal variation in food webs. Repeat observations are impossible for paleo-food webs. The best that can be done is to measure spatial or temporal variation in taxonomic composition. The latter of course could describe variability on only the longest of ecological timescales. Dunne et al. (2008) (see previous post) compiled SLNs of two Cambrian food webs derived from the Burgess Shale and Chenjiang lagerstatten, comprising 142 and 85 taxa respectively. The taxa in both these networks were subsequently aggregated into trophic species, 48 and 33 respectively, on the basis that species within the trophic species have identical consumers and resources. As argued above, it is impossible to validate this claim for fossil taxa. Trophic species-level links were ranked according to uncertainty in these networks, but there was no explicit attention paid to uncertainty at the level of species within the trophic species.

SLN derived from metanetwork in previous post

The CEG model takes an alternative approach to SLN reconstruction, generating multiple plausible SLNs from the metanetwork and hypothetical or underlying principles of food web networks as gleaned from modern food webs. This type of SLN generation requires a trophic in-link distribution for each guild. Recall that a trophic in-link distribution describes the number of prey per species within a guild. This number ranges from 1 (a heterotrophic species must prey upon at least 1 other species) up to the total species-richness of all guilds that are specified as prey of the guild in the metanetwork. SLN-generation requires initially that species within a guild be treated neutrally, that is, they have no distinguishing trophic properties. Stochastic draws from specific guild trophic link distributions then determine the number of prey to be assigned to each species. The prey species themselves are drawn randomly from the pool of prey guilds of the predatory species. The result is a directed graph or network in which each species in the community has been assigned prey, and many therefore also have predators (see figure). SLNs capture the uncertainty associated with the reconstruction of fossil food webs, and in fact any food web, in a manner in which static or unvarying trophic link determinations cannot. Repeated stochastic generation of SLNs accounts for the sources of uncertainty discussed earlier, namely uncertainty of the particular interactions of a species, and the temporal and spatial variability of a community type. Also, even though any two SLNs derived from any moderately complex metanetwork are unlikely to share the same exact topology (isomorphic), they are drawn from the same ensemble, as discussed earlier for Erdӧs-Renyi random graphs. Whether the argument can then be extended to claim that they will also have the same behavior on average, as with random graphs, is an interesting question, because the ensemble is the range of variation possible for a paleocommunity’s food web based on paleontological uncertainty. The next few posts will therefore deal with a description of the ensemble, and the ecological dynamics of the SLNs in an ensemble.