Evidence base first: the eight supplied records contain the methodological machinery for this analysis β multivariate ordination, NDVI/NMRI/fractional-cover remote sensing, variance partitioning/SEM, and dynamic vegetation modelling β but none reports dark-diversity data itself, and the primary dark-diversity literature is absent. Species-pool specifics below are therefore a blueprint to be sourced, not cited values.
Assemble a sites Γ species presence matrix from natural-vegetation plots, build regional co-occurrence pools, score suitability of absent species, and define dark diversity as suitable-but-absent species per site. Minimal working core:
import numpy as np
# X: sites x species presence (0/1) matrix
C = X.T @ X; np.fill_diagonal(C, 0) # co-occurrence counts
P = C / np.maximum(C.sum(0), 1) # conditional association
S = X @ P # site suitability (Beals-type)
pool = (X == 0) & (S > np.median(S, axis=1, keepdims=True))
dark = pool.sum(1) # dark diversity per site
ratio = dark / np.maximum(X.sum(1), 1) # impoverishment indexKnown unknown: the suitability threshold and pool definitions cannot be cited from the supplied records and must be sourced before running.
Reuse the Tibetan Plateau framework (Mann-Kendall trends, variance partitioning, LMG importance, SEM) to attribute dark diversity to climate, edaphic, and land-use covariates. Its headline result rejects temperature-only models: non-temperature factors explained about 45% of active-layer thickness and top-of-permafrost variation versus roughly 18β22% for air temperature, and SEM gave vegetation cover a β38.8% direct effect on permafrost state . Ordination is equally necessary: at two Vancouver Island marshes, elevation explained under half of vegetation variation, and dyking rewrote edaphic filters β organic content diverged by 27.000000000000004 points at Kokish (n=26/26) yet only -1.6999999999999993 points at Cluxewe (n=27/25) . GAMs on 1,920 one-metre arctic-alpine plots add nonlinear soil-temperature and moisture response surfaces .
Values as reported (approximate); categories may overlap within the 45% non-temperature total.
Reported means; Kokish shows extreme dyking divergence, Cluxewe does not.
Project future pool shifts with individual-based dynamic vegetation modelling integrating fire, phenology, and carbon allocation, as in aDGVM across African grasslandβsavannaβforest mosaics . Validate with held-out plots, per-biome cross-validation, and sensitivity analysis across pool thresholds. Falsification test: if estimated dark diversity is randomly distributed across land-use and socio-economic gradients β unlike Yangtze degradation, which tracked them β the anthropogenic-impoverishment hypothesis fails .
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