Cross‐scale scaling‐law analyses for the heterogeneity and diversity of animal gut microbiomes from community to landscape

Diversity and heterogeneity often are conflated but are fundamentally different. An aphorism proposed by Shavit and Ellison (2021; J. Phil. 118: 525–548) for distinguishing them is that ‘a zoo is diverse whereas an ecosystem is heterogeneous.' That is, a zookeeper measuring diversity simply enumerates the different types of animals; interactions are not expected to occur between animals separated by fences or other barriers. In contrast, measures of heterogeneity ought to include both interspecific interactions and relationships between species and their heterogeneous habitats. Here, we use cross‐scale, dual scaling‐law analyses of heterogeneity and diversity of animal gut microbiomes (AGMs) to address three objectives: 1) estimate the spatial heterogeneity and diversity of animal‐gut microbiomes; 2) analyze influences of phylogeny and diets on scaling of diversity and heterogeneity; 3) explore mechanistic differences between diversity and heterogeneity in AGMs. From 4903 AGM samples collected from 318 animal species covering all six classes of vertebrates and four major classes of invertebrates, we estimated that ≈640 000 operational taxonomic units (OTUs or ‘species') make up the pool of microbial species that could inhabit animal guts, among which ≈8000 are relatively common and ≈800 are dominant. The gut of any single animal, however, includes only 0.01–0.5% of the total species pool. We applied Ma's diversity‐area relationship for scaling diversity, and extend Taylor's power law and Luna et al.'s (2020; Diversity 12: 86) interaction diversity for scaling heterogeneity. At the community scale, phylogeny significantly influenced heterogeneity, but diets did not. Phylogeny and diets had limited influence on diversity at both community and landscape scales. We demonstrated that diversity and heterogeneity measure two different properties. Further, differences in scaling of diversity and heterogeneity are a result of heterogeneity being primarily the result of evolved dispersal behavior and interspecific interactions, whereas diversity results from abundances controlled on ecological time scales.