Data from: Tree compositional change in upland hardwood forests aligns with mycorrhizal type
Data files
Jun 05, 2026 version files 125.11 KB
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README.md
11.82 KB
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SizeClass_Density_Fire.csv
26.45 KB
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SizeClass_Density_Myco.csv
20.20 KB
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SizeClass_Density_Shade.csv
26.48 KB
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Small_DBH_Current.csv
19.75 KB
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Spp_Comp_Change.csv
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Abstract
Long-term monitoring in the eastern United States has documented widespread declines in fire-adapted oaks (Quercus spp.) and concurrent increases in mesophytic species such as red maple (Acer rubrum). While these changes are typically attributed to fire exclusion and shifting light and moisture regimes, belowground functional traits—particularly mycorrhizal association—may play a central role in reinforcing mesophication and altering forest resilience. To gain insight into this process, we analyzed nearly five decades of demographic data from upland oak–hickory forests on the Cumberland Plateau in southeastern USA to quantify shifts in species composition, aboveground biomass, size-class structure, and functional traits. We found that fire tolerance varied significantly by mycorrhizal type, with arbuscular mycorrhizal (AM) species more often classified as fire intolerant than ectomycorrhizal (EM) species, whereas shade tolerance was unrelated to mycorrhizal type and was instead higher in fire-intolerant than fire-tolerant species. Over time, AM and ericoid (ErM) species gained importance value (calculated by combining relative density, frequency, and aboveground total biomass) through increases in frequency and stem density, while EM species declined in importance value despite continued biomass accumulation. Fire-tolerant and intermediate shade-tolerant species exhibited similar declines, indicating a regeneration bottleneck in historically dominant EM fire-adapted trees. In contrast, AM-associated species, including broadly tolerant generalists, now dominate small stems and regeneration strata. Although total aboveground biomass increased across all functional groups, regeneration failure among EM species suggests a structural transition masked by demographic inertia of long-lived canopy individuals. These results provide evidence that shifts in mycorrhizal dominance and fire-related traits reinforce mesophication trajectories, constraining oak regeneration and reshaping belowground mycorrhizal dynamics. Integrating mycorrhizal strategies and disturbance-related traits into management frameworks may be critical for sustaining oak ecosystems and restoring fire-adapted forest structure in the eastern United States.
Dataset DOI: 10.5061/dryad.2ngf1vj3z
Description of the data and file structure
This dataset contains analysis-ready data used to evaluate long-term compositional and functional change in upland oak–hickory forests on the Cumberland Plateau of Tennessee, USA. The study uses nearly five decades of permanent-plot vegetation data from two forested watersheds: Cross Creek at Franklin State Forest and Camp Branch at Fall Creek Falls State Park. At Camp Branch, four 1-ha plots were censused in 1976, with three re-censused in 2016 and all four in 2024. At Cross Creek, four 1-ha plots were censused in 1977, three were re-censused in 1998 and 2005, and all four were re-censused in 2015 and 2023. Within each hectare, all woody stems ≥ 1.4 m tall were identified, mapped, and measured for stem diameter.
The manuscript evaluates changes in species composition, size-class structure, aboveground biomass, and functional traits. Species were classified by predominant mycorrhizal association, shade tolerance, and fire tolerance. Mycorrhizal association was classified as arbuscular mycorrhizal, ectomycorrhizal, or ericoid; shade tolerance was based on published continuous scores and collapsed into categorical classes; and fire tolerance was assigned from published sources and grouped into intolerant, intermediate, and tolerant categories. A complete species-level trait classification table is provided in the supplemental materials associated with the manuscript and is not duplicated here.
The Dryad files are organized into analysis-ready datasets that correspond to the major analyses in the manuscript: species-level changes in community composition, functional-group changes in frequency/density/aboveground biomass/importance value, and size-class distributions by mycorrhizal type, fire tolerance, and shade tolerance.
Files and variables
File: Spp_Comp_Change.csv
Description: Species-level dataset containing temporal-change values for community composition metrics between the initial census in the 1970s and the most recent census in the 2020s. This file supports analyses of changes in frequency, stem density, aboveground biomass, and importance value at both the species level and after aggregation by functional trait category. Functional trait categories include mycorrhizal type, fire tolerance, and shade tolerance.
Variables
- Site: Study site.
- Cross Creek
- Camp Branch.
- Plot: Hectare identifier within each site.
- 1
- 2
- 3
- 4
- Species: Tree species.
- ACRU = Acer rubrum
- ACSA = Acer saccharum
- ALSE = Alnus serrulata
- AMAR = Amelanchier arborea
- CADE = Castanea dentata
- CARY = Carya spp.
- CECA = Cercis canadensis
- COFL = Cornus florida
- CRAT = Crataegus spp.
- DIVI = Diospyros virginiana
- FAGR = Fagus grandifolia
- HAVI = Hamamelis virginiana
- ILDE = Ilex decidua
- ILOP = Ilex opaca
- JUVI = Juniperus virginiana
- KALA = Kalmia latifolia
- LIST = Liquidambar styraciflua
- LITU = Liriodendron tulipifera
- MAAC = Magnolia acuminata
- MALU = Malus spp.
- NYSY = Nyssa sylvatica
- OSVI = Ostrya virginiana
- OXAR = Oxydendrum arboreum
- PIEC = Pinus echinata
- PIST = Pinus strobus
- PIVI = Pinus virginiana
- PRSE = Prunus serotina
- QUAL = Quercus alba
- QUCO = Quercus coccinea
- QUFA = Quercus falcata
- QUMA = Quercus marilandica
- QUMO = Quercus montana
- QUST = Quercus stellata
- QUVE = Quercus velutina
- RHAM = Rhamnus caroliniana
- RHCA = Rhododendron spp.
- RHCO = Rhus copallinum
- RHMA = Rhododendron maximum
- ROPS = Robinia pseudoacacia
- SAAL = Sassafras albidum
- TSCA = Tsuga canadensis
- ULAL = Ulmus alata
- ULRU = Ulmus rubra
- VACC = Vaccinium spp.
- Fire: Fire-tolerance category.
- Intolerant
- Intermediate
- Tolerant
- Shade: Shade-tolerance category.
- Intolerant
- Intermediate
- Tolerant
- Myco: Predominant mycorrhizal association.
- AM = arbuscular mycorrhizal
- EM = ectomycorrhizal
- ErM = ericoid mycorrhizal
- Freq: Change in relative frequency between the most recent census in the 2020s and the initial census in the 1970s. Relative frequency was calculated within each census year as the species’ frequency divided by the sum of frequencies across all species and multiplied by 100.
- Density: Change in stem density, expressed as stems ha⁻¹, between the most recent census in the 2020s and the initial census in the 1970s.
- AGB: Change in aboveground biomass, expressed as Mg ha⁻¹, between the most recent census in the 2020s and the initial census in the 1970s.
- IV: Importance value, calculated as the average of relative frequency, relative density, and relative aboveground biomass, multiplied by 100.
File: SizeClass_Density_Myco.csv
Description: Analysis-ready size-class dataset summarizing adult stem density by mycorrhizal association, census year, site, plot, and DBH size class. This file supports the size-class analysis comparing AM, EM, and ErM-associated species.
Variables
- Year: Census year.
- 1977
- 2023
- Site: Study site.
- FSF = Franklin State Forest (Cross Creek)
- FCF = Fall Creek Falls State Park (Camp Branch)
- SiteHa: Plot or hectare identifier within each site.
- FSF.1
- FSF.2
- FSF.3
- FSF.4
- FCF.1
- FCF.2
- FCF.5
- FCF.6
- Myco: Predominant mycorrhizal association.
- AM = arbuscular mycorrhizal
- EM = ectomycorrhizal
- ErM = ericoid mycorrhizal
- SizeClass: Diameter size class in cm.
- 5 = 3.0-4.9
- 10 = 5.0-9.9
- 15 = 10.0-14.9
- 20 = 15.0-24.9
- 25 = 25.0-29.0
- 30 = 30.0-34.9
- 35 = 35.0-39.9
- 40 = 40.0-44.9
- 45 = 45.0-49.9
- 50 = 50.0-54.9
- 55 = 55.0-59.9
- 60 = 60.0-64.9
- 65 = 65.0-69.9
- 70 = 70.0-74.9
- 75 = 75.0-79.9
- 80 = 80.0-84.9
- 85 = 85.0-89.9
- Density: Stem density within the DBH class, expressed as stems ha⁻¹.
File: SizeClass_Density_Fire.csv
Description:
Variables
- Year: Census year.
- 1977
- 2023
- Site: Study site.
- FSF = Franklin State Forest (Cross Creek)
- FCF = Fall Creek Falls State Park (Camp Branch)
- SiteHa: Plot or hectare identifier within each site.
- FSF.1
- FSF.2
- FSF.3
- FSF.4
- FCF.1
- FCF.2
- FCF.5
- FCF.6
- Fire: Fire tolerance category.
- Intolerant
- Intermediate
- Tolerant
- SizeClass: Diameter size class in cm.
- 5 = 3.0-4.9
- 10 = 5.0-9.9
- 15 = 10.0-14.9
- 20 = 15.0-24.9
- 25 = 25.0-29.0
- 30 = 30.0-34.9
- 35 = 35.0-39.9
- 40 = 40.0-44.9
- 45 = 45.0-49.9
- 50 = 50.0-54.9
- 55 = 55.0-59.9
- 60 = 60.0-64.9
- 65 = 65.0-69.9
- 70 = 70.0-74.9
- 75 = 75.0-79.9
- 80 = 80.0-84.9
- 85 = 85.0-89.9
- Density: Stem density within the DBH class, expressed as stems ha -1.
File: SizeClass_Density_Shade.csv
Description:
Variables
- Year: Census year.
- 1977
- 2023
- Site: Study site.
- FSF = Franklin State Forest (Cross Creek)
- FCF = Fall Creek Falls State Park (Camp Branch)
- SiteHa: Plot or hectare identifier within each site.
- FSF.1
- FSF.2
- FSF.3
- FSF.4
- FCF.1
- FCF.2
- FCF.5
- FCF.6
- Shade: Shade tolerance category.
- Intolerant
- Intermediate
- Tolerant
- SizeClass: Diameter size class in cm.
- 5 = 3.0-4.9
- 10 = 5.0-9.9
- 15 = 10.0-14.9
- 20 = 15.0-24.9
- 25 = 25.0-29.0
- 30 = 30.0-34.9
- 35 = 35.0-39.9
- 40 = 40.0-44.9
- 45 = 45.0-49.9
- 50 = 50.0-54.9
- 55 = 55.0-59.9
- 60 = 60.0-64.9
- 65 = 65.0-69.9
- 70 = 70.0-74.9
- 75 = 75.0-79.9
- 80 = 80.0-84.9
- 85 = 85.0-89.9
- Density: Stem density within the DBH class, expressed as stems ha -1.
File: Small_DBH_Current.csv
Description: Current-census dataset summarizing stems 1–2.9 cm DBH by functional trait category. These stems were analyzed separately in the manuscript because comparable historical data were unavailable from the earliest census. This file supports the supplemental analysis of the current small-diameter stem layer.
Variables
- Site: Study site.
- Cross Creek
- Camp Branch.
- Plot: Hectare identifier within each site.
- 1
- 2
- 3
- 4
- Species: Tree species.
- ACRU = Acer rubrum
- ACSA = Acer saccharum
- ALSE = Alnus serrulata
- AMAR = Amelanchier arborea
- CADE = Castanea dentata
- CARY = Carya spp.
- CECA = Cercis canadensis
- COFL = Cornus florida
- CRAT = Crataegus spp.
- DIVI = Diospyros virginiana
- FAGR = Fagus grandifolia
- HAVI = Hamamelis virginiana
- ILDE = Ilex decidua
- ILOP = Ilex opaca
- JUVI = Juniperus virginiana
- KALA = Kalmia latifolia
- LIST = Liquidambar styraciflua
- LITU = Liriodendron tulipifera
- MAAC = Magnolia acuminata
- MALU = Malus spp.
- NYSY = Nyssa sylvatica
- OSVI = Ostrya virginiana
- OXAR = Oxydendrum arboreum
- PIEC = Pinus echinata
- PIST = Pinus strobus
- PIVI = Pinus virginiana
- PRSE = Prunus serotina
- QUAL = Quercus alba
- QUCO = Quercus coccinea
- QUFA = Quercus falcata
- QUMA = Quercus marilandica
- QUMO = Quercus montana
- QUST = Quercus stellata
- QUVE = Quercus velutina
- RHAM = Rhamnus caroliniana
- RHCA = Rhododendron spp.
- RHCO = Rhus copallinum
- RHMA = Rhododendron maximum
- ROPS = Robinia pseudoacacia
- SAAL = Sassafras albidum
- TSCA = Tsuga canadensis
- ULAL = Ulmus alata
- ULRU = Ulmus rubra
- VACC = Vaccinium spp.
- Fire: Fire-tolerance category.
- Intolerant
- Intermediate
- Tolerant
- Shade: Shade-tolerance category.
- Intolerant
- Intermediate
- Tolerant
- Myco: Predominant mycorrhizal association.
- AM = arbuscular mycorrhizal
- EM = ectomycorrhizal
- ErM = ericoid mycorrhizal
- Freq: Relative frequency of 1.0-2.9 cm DBH individuals in the most recent census in the 2020s. Relative frequency was calculated as the species’ frequency divided by the sum of frequencies across all species and multiplied by 100.
- Density: Stem density, expressed as stems ha -1, in the most recent census in the 2020s.
Notes on data processing
Adult stems were grouped into 5-cm DBH classes beginning with stems ≥ 3 cm DBH. Because historical and recent censuses differed in measurement precision for the smallest adult size class, the first size class included stems 3–5 cm DBH in the 1970s censuses and 3.0–5.4 cm DBH in recent censuses. Larger stems were measured to the nearest 0.1 cm across census periods and grouped in 5-cm increments.
Aboveground biomass was estimated from published allometric equations and aggregated to the stand level as Mg ha -1. Importance value was calculated as the average of relative frequency, relative density, and relative AGB, multiplied by 100.
Hickories were treated as a single Carya group because species-level identification was not available in the earliest census.
Code/software
Analyses were conducted in R v4.5.1. Size-class distributions were analyzed using linear models of log-transformed density, log(Density + 1), with DBH size class, census year, functional trait category, and their interactions as fixed effects. Community composition metrics were analyzed using general linear models with HC3 robust standard errors, estimated marginal means, and Tukey-adjusted pairwise contrasts. Species-level changes were analyzed using mixed-effects models with Site and Plot evaluated as candidate random effects and simplified using likelihood-ratio tests.
