Multivariate statistics were conducted using the adonis (multivariate analysis of variance) and metaMDS (ordination) functions in the package ‘vegan’ (Oksanen et al., 2016). Ordination was conducted on arcsine-square root transformed lipid biomass values using Euclidian distances. The maximum number of random starts was increased to 200 to ensure a stable solution was always returned. For each ordination, metaMDS was called iteratively six times, each time using the previous best solution as the starting point.