Data from: Site characteristics set the trajectory for biocrust restoration: Identical treatments produce differing outcomes in a North American desert
Data files
Sep 17, 2026 version files 473.43 KB
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Griffen.et.al_2026_AGG_STABILITY.xlsx
60.08 KB
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Griffen.et.al_2026_BIOCRUST_COVER.xlsx
180.64 KB
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Griffen.et.al_2026_CHLOROPHYLL.xlsx
37.75 KB
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Griffen.et.al_2026_PEN_TOR.xlsx
62.31 KB
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Griffen.et.al_2026_VEG.xlsx
115.70 KB
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README.md
16.95 KB
Abstract
Biocrusts enhance ecosystem function across multiple scales, stabilizing soils, increasing fertility, and protecting ecosystem health. Human land use and other disturbances have reduced late-successional biocrust cover globally, yet methods for restoring late-successional biocrusts, and particularly the interactive effects of biocrust inoculum source, application rate, and delivery method, remain poorly understood. We compared wild-harvested versus cultivated biocrust inoculum applied in a continuous regression design (rates of 0–40% cover) across three Sonoran Desert sites with different disturbance histories. We crossed these treatments with a range of 0–34 g of psyllium husk, intended to retain inoculum. We subsequently measured biocrust cover, soil chlorophyll-a content, and indices of erosion resistance. Most treatment combinations increased biocrust cover, but efficacy varied by site and individual treatment. In a structural equation model, wild inoculum produced more late-successional biocrust cover than cultivated. Higher inoculation rates produced more biocrust. Psyllium husk had mixed effects. Late-successional biocrust cover correlated with higher soil aggregate stability at two sites, suggesting enhanced erosion resistance. Chlorophyll-a content did not respond strongly to treatment. Wild-inoculated plots were more lichen- and moss-rich at two sites (p = 0.026; p = 0.009). We conclude that site characteristics determine the degree of restoration success, but wild-collected biocrust inoculum applied at high rates consistently produced more diverse, higher-coverage biocrusts with benefits for ecosystem function. Synthesis and applications: Though rooted in the Sonoran Desert, our results provide insights for biocrust restoration worldwide. Despite innovations in biocrust cultivation methods, wild-harvested biocrust inoculum is still more effective for late-successional biocrust restoration than cultivated, and higher application rates produce greater biocrust cover. Since wild-harvesting disturbs intact biocrusts, practitioners should seek opportunities for biocrust salvage and incorporate cultivated inoculum as cultivation methods improve to mitigate this impact. Our study also confirms the site-dependency of treatment success; we suggest that practitioners should conduct pilot studies and tailor treatments to fit site conditions. Based on our findings, wetter sites or those disturbed by fire may be better candidates for restoration using high-rate, wild-harvested biocrust inoculation, while drier or more physically disturbed sites may require different strategies.
Dataset DOI: 10.5061/dryad.pc866t25t
Description of the data and file structure
All data were collected between Dec. 2022 and Mar. 2025 as part of a biological soil crust restoration experiment at Tonto National Monument (TONT), Casa Grande Ruins National Monument (CAGR), and Organ Pipe National Monument (ORPI) in Arizona, USA. There are a total of 120 plots in the dataset, which were visited multiple times throughout the study period.
All treatments were applied a single time in December 2022.
Data files include detailed biocrust cover data at the functional group, genus, and/or species level; vegetation cover data at the genus-to-species level; soil penetration and shear strength data; soil aggregate stability data; and measurements of soil chlorophyll-a content.
Files and variables
File: Griffen.et.al_2026_BIOCRUST_COVER.xlsx
Description: Biocrust cover data for the entirety of the experiment. These data were collected using the grid-point-intercept method: a pin was dropped along a grid in the center of the plot at each of 25 intersections, with the soil surface cover at the location observed at each pin drop. Biocrust organisms or cover types observed in the grid but not directly intercepted were included as "0.5" hits; thus, some plots have "Total Hit" values above 25.
Data were not collected at Organ Pipe in Mar. 2024 due to safety and access concerns.
The "METADATA" tab of this file provides definitions of each variable.
Lichen and moss specimens listed as "unknown" were unable to be identified in the field and could not be further identified using photo or physical vouchers. In many cases, remaining unknown species were not able to be vouchered in the field due to being only present in the research plots.
Missing values are denoted as "NA."
Variables:
- Date Visited: Date that the plot measurement was taken
- Site: Site where data were collected. Options are CAGR (Casa Grande Ruins National Monument), TONT (Tonto National Monument), or ORPI (Organ Pipe National Monument).
- Plot: Plot number where measurements were recorded.
- Crust_Type: Type of biocrust applied. Options are "Cultivated" or "Wild"
- Crust_Percent: Percent surface area of biocrust inoculum applied during the treatment phase.
- Crust_Inoc_Amount: Gram amount of biocrust inoculum applied during the treatment phase. Total gram amounts to be applied in Crust_Inoc_Amount were calculated based on Crust_Percent, assuming percent coverage of a 0.75 m² plot at a depth of 0.5 cm.
- Psyllium_Inoc_Amount: Gram amount of psyllium husk powder applied during the treatment phase.
- Bare_Soil: Hits of bare mineral soil.
- Dark._Cyano: Hits of dark cyanobacteria (identified by color and/or the presence of Nostoc nodules).
- Lichen: Hits of lichens not identifiable to species or genus. Primarily recorded in December 2022 when inoculum fragments were too small to key to species or genus.
- Light_Cyano: Hits of "light" cyanobacteria identified through the presence of soil filaments or color.
- Moss: Hits of mosses not taxonomically distinguishable to genus or species. Primarily recorded in December 2022 when inoculum fragments were too small to key.
- Bryum: Hits of mosses in the genus Bryum that were not identifiable to species.
- Syntrichia: Hits of mosses in the genus Syntrichia that were not identifiable to species.
- Psyllium: Hits of psyllium husk at the soil surface.
- Plant: Hits of live plant bases.
- Litter: Hits of plant litter.
- Bioturb_Bare: Hits of bare soil disturbed by biological activity (e.g., rodent burrows).
- Bioturb_Cyano: Hits of light or dark cyanobacterial crust disturbed by biological activity (e.g., rodent burrows).
- Peltula: Hits of lichens in the genus Peltula not identifiable to species.
- Heppia_conchiloba: Hits of the lichen Heppia conchiloba.
- Psorra_decipiens: Hits of the lichen Psora decipiens.
- Scat: Hits of animal scat.
- Trash_Human: Hits of human trash.
- Mini_Rock: Hits of rocks <1 mm to 2 mm.
- Small_Rock: Hits of rocks >2 mm to 10 mm.
- Medium_Rock: Hits of rocks >10 mm to 3 cm.
- Large_Rock: Hits of rocks >3 cm.
- Liverwort: Hits of liverworts not identifiable to genus or species.
- Heppia: Hits of lichens in the genus Heppia not identifiable to species.
- Enchylium: Hits of lichens in the genus Enchylium not identifiable to species.
- Placidium: Hits of lichens in the genus Placidium not identifiable to species.
- LC_01: Hits of an unknown lichen first observed at CAGR in December 2024. Notes: "Very small, pink fruiting bodies in a black matrix. Took photo; could not find outside the plot for voucher."
- LC_02: Hits of an unknown lichen first observed at CAGR in December 2024. Notes: "Scaly, crustose. Brownish, turns yellowish when wet. Maybe just Peltula?"
- LC_05: Hits of an unknown lichen first observed at ORPI in March 2025. Notes: "Olive green, looks like Peltula with upraised and wavy margins?" Could not be vouchered.
- LC_06: Hits of an unknown lichen first observed at ORPI in March 2025. Notes: "Brighter green. Looks like a fringed Peltula observed by MB at CAGR. Margin is almost toothed."
- LC_08: Hits of an unknown lichen first observed at TONT in March 2025. Notes: "Bright green, squamules puff up (more jelly-like) when wet, with red apothecia."
- LC_10: Hits of an unknown lichen first observed at TONT in March 2025. Notes: "Brown lichen, stays olive-brown when wet, looks kind of 'cabbage-y.' Big pile of puffy squamules. Placidium?"
- LC_11: Hits of an unknown lichen first observed at TONT in March 2025. Notes: "Round, very tan/brown squamules, similar in color to Endocarpon but with a different gestalt. No visible black perithecia under a hand lens. Dark black margin."
- Hyphae: Hits of fungal hyphae.
- Nostoc: Hits of Nostoc.
- Endocarpon: Hits of lichens in the genus Endocarpon not identifiable to species.
- M_01: Unknown moss first observed at CAGR in December 2024. Notes: "Upright moss with long awns." Could not be vouchered or photographed.
- Psorra_iterica: Hits of the lichen Psora icterica.
- Yellow..Termite.Activity..TA.: Observations of yellow termite activity. Recorded only when observed, but not included in cover quantification.
- Season_Year: Identifier based on season and year. March/April observations are classified as Spring; November/December observations are classified as Winter.
- Peltula_patellata: Hits of the lichen Peltula patellata.
- Peltula_richardsii: Hits of the lichen Peltula richardsii.
- Total_Cover: Total hits within a plot. Values greater than 25 indicate inclusion of 0.5-point observations for organisms present in the frame but not directly intercepted by the pin.
- Total_Lichen: Total lichen hits.
- Total_Moss: Total moss hits.
- Total_Liverwort: Total liverwort hits.
- Total_Bare: Total bare soil hits.
- Total_BSC: Total hits of biocrust organisms, defined here as dark cyanobacteria, lichens, mosses, and liverworts, but excluding light cyanobacteria.
- not_habitat: Hits of non-habitat categories including plant bases, litter, scat, rocks of any size, and human trash.
- crust_habitat: Total hits in a plot minus non-habitat hits.
- crust_niche_occupancy: Percentage calculated as (Total_BSC ÷ crust_habitat) × 100.
- cno: crust_niche_occupancy expressed as a decimal between 0 and 1 (percentage divided by 100)
File: Griffen.et.al_2026_PEN_TOR.xlsx
Description: This file contains data on soil penetration resistance and shear strength. Penetration resistance was tested using an AMS Pocket Penetrometer. Shear strength was tested using a Durham Geo Slope Indicator Torvane device. The "METADATA" tab of this document provides definitions of each variable.
Data were not collected at ORPI in Dec. 2023 due to safety issues.
Occasionally, rocks or other obstacles prevented us from taking all three data points in a given plot. Missing values are denoted as "NA."
Variables
- Site: The site where data were collected. Options are CAGR (Casa Grande Ruins National Monument), TONT (Tonto National Monument), or ORPI (Organ Pipe National Monument).
- Date.Visited: Date that the plot measurement was taken.
- Plot: Plot number where the measurements were recorded.
- Crust_Type: Type of biocrust applied. Options are "Cultivated" or "Wild"
- Crust_Percent: Percent surface area of biocrust inoculum applied during the treatment phase.
- Crust_Inoc_Amount: Gram amount of biocrust inoculum applied during the treatment phase. Total gram amounts to be applied in Crust_Inoc_Amount were calculated based on Crust_Percent, assuming percent coverage of a 0.75 m² plot at a depth of 0.5 cm.
- Psyllium_Inoc_Level: Gram amount of psyllium husk powder applied during the treatment phase.
- Penetrometer..kg.cm.2.: Soil penetration resistance measured in kg/cm².
- Torvane..kg.cm.2.: Soil shear strength measured in kg/cm².
- Season_Year: Identifier based on season and year. March/April observations are classified as Spring; November/December observations are classified as Winter.
File: Griffen.et.al_2026_CHLOROPHYLL.xlsx
Description: This file contains chlorophyll-a data collected from soil samples from each plot. Samples were collected as six 1 cm-deep subsamples per plot, which were aggregated before extraction. We used a double extraction with spectrophotometric-grade ethanol following Castle et al. (2011).
We did not collect soil samples from ORPI in Dec. 2023 or Mar. 2024 due to safety issues. Those locations/dates are thus not present in this dataset. Missing data are otherwise denoted as "NA."
References:
Castle, S. C., Morrison, C. D., & Barger, N. N. (2011). Extraction of chlorophyll a from biological soil crusts: A comparison of solvents for spectrophotometric determination. Soil Biology and Biochemistry, 43(4), 853–856. https://doi.org/10.1016/j.soilbio.2010.11.025
Variables
- Season_Year: The season and year in which the sample was collected. March/April observations are classified as Spring; November/December observations are classified as Winter.
- Date_Extracted: The specific date on which a sample extraction was performed.
- Site: The site where data were collected. Options are CAGR (Casa Grande Ruins National Monument), TONT (Tonto National Monument), or ORPI (Organ Pipe National Monument).
- Plot: The specific plot from which the soil sample was collected and from which soil chlorophyll-a was extracted.
- Crust_Type: Type of biocrust applied. Options are "Cultivated" or "Wild"
- Crust_Percent: Percent surface area of biocrust inoculum applied during the treatment phase.
- Crust_Inoc_Amount: Gram amount of biocrust inoculum applied during the treatment phase. Total gram amounts to be applied in Crust_Inoc_Amount were calculated based on Crust_Percent assuming percent coverage of a 0.75 m² plot at a depth of 0.5 cm.
- Psyllium_Inoc_Level_g: Gram amount of psyllium husk powder applied during the treatment phase.
- SAMPLE.ID: Unique identifier assigned to each individual sample.
- tot_chl_a_no_acid: Micrograms of chlorophyll-a per gram of soil in the sample (µg chl-a/g soil). Data reflect an unacidified state.
File: Griffen.et.al_2026_AGG_STABILITY.xlsx
Description: This file contains soil aggregate stability measurements collected from each plot using a field soil aggregate stability kit and methods from Herrick et al. (2021). Three peds were collected and submerged per plot.
Occasionally, we were unable to find a suitable location for a sample. We did not collect any soil aggregate stability data in Dec. 2022, Mar. 2023, or Mar. 2024, and these dates/locations are thus not present in the dataset. Missing data from dates/locations where data was collected are indicated as "NA."
References:
Herrick, J. E., Whitford, W. G., de Soyza, A. G., Van Zee, J. W., Havstad, K. M., Seybold, C. A., & Walton, M. (2001). Field soil aggregate stability kit for soil quality and rangeland health evaluations. CATENA, Soil Aggregation in Arid and Semi-Arid Environments, 44(1), 27–35. https://doi.org/10.1016/S0341-8162(00)00173-9
Variables
- Site: The site where data were collected. Options are CAGR (Casa Grande Ruins National Monument), TONT (Tonto National Monument), or ORPI (Organ Pipe National Monument).
- Date.Visited: Date that the plot measurement was taken.
- Plot: Plot number where the measurements were recorded.
- Crust_Type: Type of biocrust applied. Options are "Cultivated" or "Wild"
- Crust_Percent: Percent surface area of biocrust inoculum applied during the treatment phase.
- Crust_Inoc_Amount: Gram amount of biocrust inoculum applied during the treatment phase. Total gram amounts to be applied in Crust_Inoc_Amount were calculated based on Crust_Percent, assuming percent coverage of a 0.75 m² plot at a depth of 0.5 cm.Psyllium_Inoc_Level: Gram amount of psyllium husk powder applied during the treatment phase.
- Position: Three soil peds were collected per plot in pre-selected positions designed to prevent observers from resampling previously disturbed areas. The numerical values refer to these assigned positions.
- Veg.Cover: If recorded, this category describes the type of nearby vegetation cover shading the ped collection location. These data were not collected consistently and were not used in analyses relevant to this study. Abbreviations include: "NC" = No Cover; "AG" = Annual Grass; "AF" = Annual Forb; "PG" = Perennial Grass; "PF" = Perennial Forb; and "Sh" = Shrub.
- Class: Soil aggregate stability class determined following Herrick et al. (2001), Field Soil Aggregate Stability Kit for Soil Quality and Rangeland Health Evaluations (CATENA, 44(1), 27-35; https://doi.org/10.1016/S0341-8162(00)00173-9).
- Season_Year: Identifier based on season and year. March/April observations are classified as Spring; November/December observations are classified as Winter.
File: Griffen.et.al_2026_VEG.xlsx
Description: This file contains data from ocular estimates of plant cover in each plot. Cover was estimated using the following cover classes: <1%, 1–5%, 6–25%, 26–50%, 51–75%, 76–95%, 95–100%.
The tab "METADATA" contains a full list of species codes and abbreviations.
We were unable to collect vegetation data at ORPI in Spring 2024 due to safety concerns. Data from this location/time are thus not included in the dataset. Missing data are entered as "NA." In some cases, plots where no plants were observed were not entered in the dataset--these plots have been confirmed to have had no cover for the non-entered location/time.
Variables
- Site.: The site where data were collected. Options are CAGR (Casa Grande Ruins National Monument), TONT (Tonto National Monument), or ORPI (Organ Pipe National Monument).
- Date.Visited.: Date that the plot measurement was taken.
- Plot...: Plot number where the measurements were recorded.
- Cover.: Four-letter code for the encountered species. Codes typically follow USDA PLANTS symbols, though not always. The Scientific.Name. column contains the authoritative identification. In some cases, unique field-generated identifiers were used instead and are defined elsewhere in the dataset documentation.
- Scientific.Name.: Scientific name of the recorded taxon. The taxonomic level reflects the level of identification possible by the field crew or through voucher specimens. Identifications may be recorded at the family, genus, or species level.
- Functional.Type.: Life form or functional group of the recorded species.
- Cover.Class.: Estimated percent cover of the species within the plot. Cover was recorded using the following classes: <1%, 1-5%, 6-25%, 26-50%, 51-75%, 76-95%, and 95-100%.
- Season_Year: Identifier based on season and year. March/April observations are classified as Spring; November/December observations are classified as Winter.
- veg_simp: Median value of the reported cover class (%). This value was used in statistical analyses.
Code/software
All data files are in an Excel workbook format. These can be freely viewed using Google Drive or LibreOffice, or imported to R using the package readxl.
