Companion to the paper Karsdorp, F., Kandler, A., Kestemont, M., Romanowska, I. & Stapel, R. (2026). Correcting collection bias in comparative studies of diversity. Journal of the Royal Society Interface 23: 20260163. doi:10.1098/rsif.2026.0163

Comparing diversity under collection bias

Two assemblages, A and B, are recorded from synthetic populations under different collection biases. Pick a bias scenario and a standardization strategy, and see which one still recovers the true richness ratio.

Sample size → observed richness

Coverage → observed richness

Assemblage A Assemblage B standardization point (Ĉmax / nmin)
Collection-bias scenario:

Assemblage A

Assemblage B

Shared populations

unbiased (γ = 0)over-records rare (γ = 3) →
True richness ratio  A : B set by the populations — independent of how they were sampled
Bars show each estimate’s deviation from the true ratio · centre line = exact.

The two populations share one abundance shape, so a difference in diversity is real, not an artefact of structure. Sample sizes capture population-size and productivity (“rich-get-richer”) biases — some assemblages simply attract more records. The rarity-bias slider captures Stromer’s Riddle: records are drawn with weight ∝ count(1−γ), so larger γ over-records rare, novel variants. Coverage-based standardization corrects all three; size- and rarefaction-based comparisons do not.

Richness accumulation and sample coverage use the Chao / iNEXT estimators — the same estimate_coverage and rarefaction_extrapolation as copia — and the standardization follows the reference code (coverage at Ĉmax = the smaller of the two sample coverages; rarefaction at nmin), averaged over many resamples. For clarity the two populations share one log-normal abundance shape, where the paper’s full study draws populations from a neutral (Wright–Fisher) model. Its accuracy is governed primarily by the effective standardized sample size — the number of records retained at the comparison point — not by the coverage level reached (n ≥ 30 sufficed in the paper’s simulations).