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Network analysis has revealed whole-network properties hypothesized to begeneral characteristics of ecosystems including pathway proliferation, andnetwork non-locality, homogenization, amplification, mutualism, and synergism.Collectively these properties characterize the impact of indirect interactionsamong ecosystem elements. While ecosystem networks generally trace athermodynamically conserved unit through the system, there appear to be severalmodel classes. For example, trophic (TRO) networks are built upon a food web,usually follow energy or carbon, and are the most abundant in the literature.Biogeochemical cycling (BGC) networks trace nutrients like nitrogen orphosphorus and tend to have more recycling than TRO. We tested (1) thehypothesized generality of the properties in BGC networks and (2) that theytend to be more strongly expressed in BGC networks than in the TRO networks dueto increased recycling. We compared the properties in 22 BGC and 57 TROecosystem networks from the literature using enaR. The results generallysupport the hypotheses. First, five of the properties occurred in all 22 BGCmodels, while network mutualism occurred in 86% of the models. Further, theseresults were generally robust to a $\pm$50% uncertainty in the modelparameters. Second, the mean network statistics for the six properties werestatistically significantly greater in the BGC models than the TRO models.These results (1) confirm the general presence of these properties in ecosystemnetworks, (2) highlight the significance of model types in determining propertyintensities, (3) reinforce the importance of recycling, and (4) provide a setof indicator benchmarks for future systems comparisons. Further, this workhighlights how indirect effects distributed by network connectivity cantransform whole-ecosystem functioning, and adds to the growing domain ofnetwork ecology.

Six General Ecosystem Properties are more Intense in Biogeochemical Cycling Networks than Food Webs