Data and code from: Climate change will exacerbate the mismatch between plant diversity hotspots and protected areas in the Hindu Kush Himalaya
Data files
Sep 14, 2026 version files 19.41 MB
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README.md
16.48 KB
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Supplementary_Information_and_R_codes_R2.zip
19.39 MB
Abstract
Biodiversity conservation strategies often fall short of long-term planning. To address this gap, we integrated a completed phylogeny of 22,058 vascular plant species and 464,407 distribution records across the Hindu Kush Himalaya with climate projections. This revealed the highest species richness, phylogenetic diversity, phylogenetic endemism, and weighted endemism (plant diversity and endemism) in southern low- to mid-elevations during the Last Glacial Maximum and the current period. The centers of neo-endemism were located in the southeast, while paleo-endemism grids were distributed throughout the study area. Future projections (2040-2100) revealed clear northward and upward shifts in plant diversity and endemism hotspots, driven by changes in precipitation (explained variance > 30 %) and temperature extremes (explained variance > 20 %). These shifts increased the current 54 % mismatch between diversity hotspots and protected areas to over 80 %. By overlaying current and future hotspots with current protected areas, we identified 17 significant conservation gaps: 8 require immediate attention, and 9 are essential for future conservation planning. Exploratory path analysis revealed that 39 % of species richness and 16 % of phylogenetic endemism variation were explained, with evolution showing the strongest association with species richness and climate with phylogenetic endemism. Our findings emphasize the need for spatio-temporal conservation planning to ensure the establishment of climate-resilient protected areas.
Dataset DOI: 10.5061/dryad.h9w0vt4wz
Description of the data and file structure
This README describes the supporting datasets, analytical outputs, R codes, supplementary materials, and methodological information associated with the manuscript:
“Climate change will exacerbate the mismatch between plant diversity hotspots and protected areas in the Hindu Kush Himalaya.”
Prepared for Submission to: Dr. Yad
1. Overview
This dataset supports the study evaluating spatial and temporal patterns of vascular plant diversity, phylogenetic diversity, and endemism across the Hindu Kush Himalaya.
The dataset includes:
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464,407 vascular plant distribution records;
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22,058 accepted vascular plant species;
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a completed analytical phylogeny containing 22,058 species;
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current, Last Glacial Maximum, and future climate data;
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spatial taxonomic and phylogenetic diversity metrics;
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Random Forest and Generalized Linear Model projections;
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multidimensional biodiversity hotspot analyses;
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current-period CANAPE analysis;
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protected-area gap assessment;
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species-level complementarity analysis;
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exploratory Partial Least Squares Path Modeling (PLS-PM); and
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ecological vulnerability assessment.
External data sources are listed below with links/DOIs. Users should obtain raw data directly from the original providers under their terms.
Source Use in this study Link / DOI Included in Dryad? POWO Taxonomic standardization https://powo.science.kew.org/ No raw data; standardized names only GBIF Occurrence records https://www.gbif.org/ ; download DOIs are listed in Supporting Information Table S2 No raw records; derived grid presence only WorldClim 2.1 Current and future bioclimatic variables https://www.worldclim.org/data/worldclim21.html No raw layers; derived grid summaries WorldClim CMIP6 / MIROC6 Future projections under SSP2-4.5 https://www.worldclim.org/data/cmip6/cmip6_clim30s.html No raw layers; derived projections PaleoClim / NCAR-CCSM4 Last Glacial Maximum bioclimatic variables https://www.paleoclim.org/ No raw layers; derived grid summaries WDPA Protected-area gap analysis https://www.protectedplanet.net/en/thematic-areas/wdpa?tab=WDPA No raw boundaries; derived protected according to study area map. PREDICTS Biodiversity Intactness Index PREDICTS database (Hudson et al., 2017) No raw data; derived grid values NASA LP DAAC MODIS NDVI, EVI, and vegetation quality layers NASA LP DAAC at the USGS EROS center (MODI3Q1), NASA LP DAAC at the USGS EROS center (MODI3A2), NASA LP DAAC No raw layers; derived grid values CGIAR-CSI Aridity index Global Aridity Index and Potential Evapotranspiration database version 3.1 (Zomer et al., 2022) No raw layers; derived grid values Human footprint Anthropogenic pressure Gassert et al. 2023 (Global 100 m resolution dataset) No raw layers; derived grid values Population density Anthropogenic pressure Global 1-km gridded population dataset (L. Liu et al., 2024) No raw layers; derived grid values Habitat heterogeneity Habitat heterogeneity indices Metrics of spectral habitat heterogeneity. (Tuanmu & Jetz, 2015) No raw layers; derived grid values Land use / land cover Global Human Modification Land Use/Land Cover classification (Kennedy et al., 2020) No raw layers; derived grid values R packages Analysis and visualization CoordinateCleaner, V.PhyloMaker, picante, Biodiverse, caret, glmulti, iml, VennDiagram, ggupset, dplyr, tidyr, lmtest, broom Code only
2. Core Workflow Summary
The analysis was conducted through the following sequential workflow.
2.1 Species-occurrence compilation
Plant distribution records were compiled from:
- national and regional floras;
- taxonomic checklists;
- herbarium datasets;
- published occurrence databases; and
- Global Biodiversity Information Facility (GBIF) downloads.
The complete list of national and regional floras and all GBIF download DOIs is provided in the Supporting Information.
2.2 Taxonomic standardization
Species names and synonyms were standardized against Plants of the World Online, facilitated by the Royal Botanic Gardens, Kew.
Invalid coordinates, zero coordinates, non-numeric coordinates, and marine records were removed. Verified occurrence records were converted into presence records within 50 × 50 km grid cells across the Hindu Kush Himalaya.
2.3 Phylogenetic-tree preparation
The source phylogeny contained 22,152 species and was derived from a previously published dated phylogeny of the Pan-Himalayan flora.
After matching the occurrence dataset with the source phylogeny:
- the pruned backbone tree contained 19,944 species;
- 2,114 missing species were incorporated using V.PhyloMaker; and
- the final analytical phylogeny contained 22,058 species.
Ten replicate completed trees were generated to evaluate sensitivity to random phylogenetic grafting.
2.4 Spatial phylogenetic analyses
Grid-level taxonomic and phylogenetic diversity metrics were calculated using Biodiverse and R.
The calculated indices included:
- species richness;
- phylogenetic diversity;
- phylogenetic endemism;
- weighted endemism;
- relative phylogenetic diversity;
- relative phylogenetic endemism;
- standardized effect size of phylogenetic diversity;
- net relatedness index; and
- nearest taxon index.
2.5 Environmental variables and temporal projections
Current, Last Glacial Maximum, and future climate variables were prepared for diversity projections.
Random Forest-based variable selection and Generalized Linear Models were used to project four principal diversity indices:
- species richness;
- phylogenetic diversity;
- phylogenetic endemism; and
- weighted endemism.
Random Forest models were fitted using 500 trees. Predictor selection was performed using Random Forest-based recursive feature elimination with ten-fold cross-validation.
Model performance was assessed using:
- coefficient of determination (R²);
- root mean square error (RMSE); and
- mean absolute error (MAE).
Future projections were generated after refitting the selected models using the complete current dataset.
2.6 Hotspot identification
Multidimensional biodiversity hotspots were identified based on the spatial overlap of:
- species richness;
- phylogenetic diversity;
- phylogenetic endemism; and
- weighted endemism.
Two hotspot thresholds were assessed:
- top 5 %, representing the highest-priority hotspot areas; and
- top 10 %, representing broader hotspot regions.
Temporal changes in hotspots were evaluated through:
- spatial loss;
- gain;
- retention;
- turnover;
- directional shifts; and
- elevational shifts.
2.7 CANAPE analysis
CANAPE was conducted only for the current period to identify centers of:
- neo-endemism;
- paleo-endemism; and
- mixed-endemism.
These analyses characterized current evolutionary patterns but were not incorporated into the Level I or Level II conservation-gap framework.
2.8 Conservation-gap analysis
Current and future multidimensional biodiversity hotspots were overlaid with the existing World Database on Protected Areas.
Conservation gaps were defined as hotspot areas occurring outside the current protected-area network.
2.9 Complementarity analysis
Species-level complementarity analysis was conducted to evaluate representation of plant species within existing protected areas and identified conservation gaps.
The analysis:
- identified species with less than 10 % of occupied grid cells represented within protected areas;
- quantified their representation within Level I and Level II conservation gaps;
- identified narrow-endemic species occupying five or fewer grid cells; and
- evaluated evolutionary distinctiveness among poorly represented species.
2.10 Driver and vulnerability analyses
Exploratory Partial Least Squares Path Modeling (PLS-PM) was used to examine relationships among:
- evolutionary factors;
- vegetation;
- climate;
- topography;
- anthropogenic pressure;
- species richness; and
- phylogenetic endemism.
An ecological vulnerability index (EcoVI) was calculated using pressure, resilience, and sensitivity indicators.
The abbreviation EVI refers only to the remotely sensed enhanced vegetation index.
3. Repository structure
The complete supplementary materials are provided in:
Supplementary_Information_and_R_codes_R2.zip
This archive contains two folders:
Folder 1: R codes
The R scripts used for data processing, analyses, and visualization are provided as follows:
Code 1_VphyloMaker
R code for phylogenetic tree preparation and incorporation of missing species using V.PhyloMaker.
Code 2_SESPDNTIandNRI
R code for calculating phylogenetic structure metrics, including SES.PD, NTI, and NRI.
Code 3_Prediction-LGM-RF
R code for Random Forest modelling, recursive feature elimination, GLM modelling, and Last Glacial Maximum and future diversity projections.
Code 4_Upset plot and Venndiagrams
R code for generating UpSet plots and Venn diagrams.
Code 5_Plspm model
R code for Partial Least Squares Path Modeling (PLS-PM) analysis.
Code 6_EVI
R code for calculating the ecological vulnerability index (EcoVI).
Code 7_Complementarity analysis
R code for species-level complementarity analysis and conservation-gap assessment.
Folder 2: Supporting Information files 1–24
This folder contains supplementary datasets, tables, figures, and methodological information associated with the manuscript.
Supporting Information 1
Species list and locality records.
Supporting Information 2
List of fossil calibrations and records.
Supporting Information 3
Tactual_HKH.tre — initial phylogenetic tree for the Hindu Kush Himalaya dataset.
Supporting Information 4
List of species incorporated using V.PhyloMaker.
Supporting Information 5
Full completed phylogenetic tree.
Supporting Information 6
Variables used for the 2100 projection.
Supporting Information 7
Current plant diversity indices.
Supporting Information 8
Predictions from the Last Glacial Maximum model.
Supporting Information 9
Predictions for 2021–2040.
Supporting Information 10
Predictions for 2041–2060.
Supporting Information 11
Predictions for 2061–2080.
Supporting Information 12
Predictions for 2081–2100.
Supporting Information 13
Projected plant diversity indices for the Last Glacial Maximum and future periods.
Supporting Information 14
Gain and loss data among different periods.
Supporting Information 15
Multidimensional hotspot classifications for all periods using top 5 % and top 10 % thresholds.
Supporting Information 16
CANAPE categories.
Supporting Information 17
Phylogenetic Jaccard groups and clustering results.
Supporting Information 18
Protected and non-protected area grids.
Supporting Information 19
Variables used in correlation analysis for PLS-PM.
Supporting Information 20
Variables used in ecological vulnerability index (EcoVI) analysis.
Supporting Information 21
Complementarity analysis summary.
Supporting Information 22
Endemism summary.
Supporting Information 23
Poorly represented species.
Supporting Information 24
Poorly represented species with endemism information.
4. Variable descriptions
Variables occurring repeatedly across different temporal periods follow the same naming conventions.
Diversity variables
| Variable | Description |
|---|---|
| SR | Species richness |
| PD | Phylogenetic diversity |
| PE | Phylogenetic endemism |
| WE | Weighted endemism |
Temporal abbreviations
| Term | Description |
|---|---|
| LGM | Last Glacial Maximum |
| Cur | Current period |
| 2040 | 2021–2040 projection period |
| 2060 | 2041–2060 projection period |
| 2080 | 2061–2080 projection period |
| 2100 | 2081–2100 projection period |
Environmental variables
Environmental predictors retain their original names from the source datasets and analytical files.
Examples:
| Variable | Description |
|---|---|
| Bio variables | Bioclimatic variables |
| EVI | Enhanced Vegetation Index |
| NDVI | Normalized Difference Vegetation Index |
| HFP | Human Footprint |
| Popdensity | Population density |
| BII | Biodiversity Intactness Index |
Projection variables
| Variable | Description |
|---|---|
| SR2040 | Projected species richness for 2021–2040 |
| PD2040 | Projected phylogenetic diversity for 2021–2040 |
| PE2040 | Projected phylogenetic endemism for 2021–2040 |
| WE2040 | Projected weighted endemism for 2021–2040 |
Equivalent naming conventions apply for 2060, 2080, and 2100.
Conservation and complementarity variables
| Variable | Description |
|---|---|
| Total_Grids | Total occupied grid cells |
| Protected_Grids | Occupied grid cells within protected areas |
| Prop_Protected | Proportion of occupied grid cells protected |
| ED | Evolutionary distinctiveness |
| Narrow_Endemic | Narrow endemic classification |
| Top10_ED | Top 10 % evolutionary distinctiveness classification |
Variable definitions, transformations, and analytical procedures are further described in the associated manuscript Methods and Supporting Information.
