DOI: 10.5061/dryad.0gb5mkm8v
title: Multiple disturbances, multiple legacies: Fire, canopy gaps and deer jointly change the forest seed bank

description: Disturbance regimes, like low-intensity fire, canopy gaps, and ungulate
 browsing, play a critical role in determining ecological composition and
 structure in temperate forests around the world. Each disturbance (or lack
 thereof) can lead to unique plant communities, but we do not understand
 how combined disturbances change plant diversity and the resulting soil
 seed bank. Changes in the soil seed bank, which depend on the plants that
 survive post-disturbance, can then influence future biodiversity and
 succession. We used a long-term experiment in West Virginia, U.S.A. that
 factorially manipulated low-intensity fire, deer exclusion, and canopy
 gaps. Thirteen years after disturbance initiation, we sampled the seed
 bank from each disturbance treatment. We found that low-intensity fire led
 to increased seed bank density, with additional canopy gaps and deer
 exclusion each creating unique seed bank communities. Combined fire,
 canopy gaps, and deer presence led to high seed bank diversity and the
 most unique seed communities, while canopy gaps and deer had no effect on
 seed banks unless the area was previously burned. In contrast, combined
 fire, canopy gaps, and deer exclusion led to the lowest seed bank
 diversity of all treatments, reflecting the continued legacy of extant
 plants that grew immediately after disturbance. Seed communities were also
 distinct from extant understory species over thirteen years, regardless of
 disturbance treatment. Each reintroduced disturbance combination left a
 unique legacy in the seed bank that will likely influence future forest
 reorganization following disturbances, adding to our understanding of how
 multiple disturbances influence forest succession and organization.
 Synthesis: Forest disturbance regimes have changed around the world and
 are being restored or manipulated to support biodiversity. Reintroduction
 of disturbance leads to unique plant communities, but we do not understand
 how combined disturbances change the soil seed bank. Using an experiment
 that manipulates low-intensity fire, canopy gaps, and deer exclusion, we
 find that combinations of these experimental treatments leads to
 substantially different seed communities. These disturbance-altered seed
 banks will likely influence future biodiversity and successional patterns,
 highlighting how the restoration of disturbance can strongly and
 indirectly influence temperate forest community dynamics.
description: <strong>Site:</strong> We manipulated prescribed
 fire, deer presence, and canopy gap creation in four replicate Appalachian
 hardwood stands in central West Virginia, USA. We established this
 experiment in 2000 in the Western Allegheny Mountain ecological subsection
 using two stands in the Monongahela National Forest (39º06’ N, 79º43’ W)
 and two stands in the Fernow Experimental Forest (39º01’ N, 79º42’ W).
 Each stand was 60 to 90 years old and between 670 to 800 m in
 elevation.  <strong>Experimental Design:
 </strong>Our experimental design was a split-plot factorial (Fig.
 1), with each stand split in half and randomly assigned a burn treatment
 (burned or unburned). In each burned and unburned half stand, we
 established treatment plots (20 x 20 m, 400 m<sup>2</sup>)
 representing 8 disturbance treatments: 1) no burn + no deer + no canopy
 gap, 2) no burn + no deer + canopy gap, 3) no burn + deer + no canopy gap,
 4) no burn + deer + canopy gap, 5) burn + no deer + no gap, 6) burn + no
 deer + canopy gap, 7) burn + deer + no canopy gap, 8) burn + deer + canopy
 gap. There were 8 replicates per treatment. Treatment plots were 20 m from
 one another, stand edges, and burn lines to avoid nonindependence and edge
 effects. One fire and canopy gap treatment plot could not be found in
 2013, thus n = 63. We placed five permanent 1-m<sup>2
 </sup>sampling quadrats within each treatment plot. We collected
 seed bank samples at each corner of the five quadrats using a 5-cm long
 section of a 10-cm PVC pipe (Soil Volume: 392.5 cm<sup>3</sup>
 x 4 = 1570 cm<sup>3</sup> soil sampled per quadrat). All 20
 soil cores per treatment plot were then pooled, mixed, and subsampled for
 use in emergence trials. Three subsamples were taken from each of the 63
 treatment plot’s pooled soils and placed in separate 625
 cm<sup>2</sup> square trays in a greenhouse (625
 cm<sup>2</sup> x 3 = 1875 cm<sup>2</sup> soil per
 plot; 1563 cm<sup>3</sup> x 3 = 4689
 cm<sup>3</sup> soil per plot), with 2.5 cm of subsampled soil
 placed on top of 2 cm of sterile sand in each tray. We watered all 189
 trays (63 treatment plots x 3 subsamples) daily and occasionally rotated
 the trays to minimize any greenhouse positional effects (e.g., light,
 temperature). All germinants were identified to species or genera
 depending on life form, counted, and removed from the tray. After 5
 months, we subjected trays to a 90-day, 5º C cold stratification period,
 after which they were returned to the greenhouse for another germination
 phase. <strong>Analysis: </strong>For
 operational purposes we define seed species density as the total number of
 species found across the three trays representing a single treatment plot.
 Similarly, seed abundance is defined as the total number of germinants
 found across the three trays per plot. Classic Shannon diversity is
 calculated via the vegan R package (Oksanen et al. 2022).
description: # Multiple disturbances and multiple legacies: Fire, canopy gaps and deer
 jointly change the forest seed bank
 [https://doi.org/10.5061/dryad.0gb5mkm8v](https://doi.org/10.5061/dryad.0gb5mkm8v) ## Description of the data and file structure ### Files and variables | Datasheet(s) | Column Name | Variable Description | Variable Type | Values | | :----------------------------------------------------- | :-------------------------- | :----------------------------------------------------------------------------------------------------------------------------- | :------------------- | :------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------ | | | | | | | | bank.plot.wide\_named.var | MP | Site (4 total) | Categorical, numeric | Site 1 = 10, Site 2 = 20, Site 3 = 30, Site 4 = 40 | | bank.wide.forb | SP | Sampling Plot (20 x 20 m disturbed plots with a total of 16 per site) | Categorical, numeric | Small Plots 1-12 | | bank.wide.graminoid | Fire | Burned or unburned plots | Categorical | Fire; No Fire | | bank.wide.shrub | Fence | Deer are fenced out (NoDeer) or deer are present (Deer) | Categorical | Deer; No Deer | | bank.wide.tree | Gap | Plots with or without canopy gaps | Categorical | Gap; No Gap | | bank.wide.vine | SpeciesDensity | The density of seed species per plot | Numeric | | | | Abund | The abundance of seeds per plot | Numeric | | | | H | The classic Shannon diversity per plot | Numeric | | | | Columns F:BM or Column 6:65 | Species abbreviations and their associated counts. Refer to USDA Plants for Greater Detail | Numeric | Acer pensylvanicum = ACPE; Acer rubrum = ACRU; Ageratina altissima = AGAL5; Agrostis perennans = AGPE; Andropogon spp. = ANDRO2; Apocynum spp. = APOCY; Aralia spinosa = ARSP2; Berberis thunbergii = BETH; Betula spp. = BETUL; Boehmeria cylindrica = BOCY; Carex spp. = CAREX; Dalibarda repens = DARE; Dichanthelium clandestinum = DICL; Erechtites hieraciifolius = ERHI2; Eupatorium serotinum = EUSE2; Eurybia divaricata = EUDI16l Galium spp. = GALIU; Gaultheria procumbens = GAPR2; Hypericum punctatum = HYPS; Juncus effusus = JEUF; Liriodendron tulipifera = LITU; Lobelia spicata = LOSP; Luzula multiflora = LUMU2; Medeola virginiana = MEVI; Mitchella repens = MIRE; Packera aurea = PAAU3; Phytolacca americana = PHAM4; Pilea pumila = PIPU2; Plantago lanceolata = PLLA; Polygonum persicaria = POPE3; Potentilla canadensis = POCA; Prenanthes trifoliolata = PRTR; Rhus typhina = RHTY; Robinia pseudoacacia = ROPS; Rosa multiflora = ROMU; Rubus spp. = RUBUS; Sambucus spp. = SAMBU; Sassafras albidum = SAAL5; Saxifraga spp. = SAXIF; Sisyrinchium spp. = SISYR; Solanum carolinense = SOCA3; Solanum nigrum = SONI; Stellaria media = STME2; Trillium spp. = TRILL; Uvularia sessilifolia = UVSE; Verbascum thapsus = VETH; Verbena urticifolia = VEUR; Veronica officinalis = VEOF2; Viola blanda = VIBL; Viola canadensis = VICA4; Viola macloskeyi = VIMA2; Viola pubescens = VIPU3; Viola rotundifolia = VIRO2; Viola sororia = VISO; Viola spp. = VIOLA; Vitis aestivalis = VIAE | | | | | | | | | | | | | | allyrherb\_seed\_wide\_firenofencegap\_combgenera | MP | Site (4 total) | Categorical, numeric | Site 1 = 10, Site 2 = 20, Site 3 = 30, Site 4 = 40 | | allyrherb\_seed\_wide\_firenofencegap\_combgenera\_trt | SP | Sampling Plot (20 x 20 m disturbed plots with a total of 16 per site - Only plots with fire, no fence, and a gap in this data) | Categorical, numeric | Plots 1, 4, 7, 6 | | | FireFenceGap | Treatment combo | Categorical | Fire\_Gap\_Fence, Fire\_Gap\_NoFence, Fire\_NoGap\_Fence, Fire\_NoGap\_NoFence, NoFire\_Gap\_Fence, NoFire\_Gap\_NoFence, NoFire\_NoGap\_Fence, NoFire\_NoGap\_NoFence | | | Year | Year | Categorical, numeric | 2000 (pre-treatment), 2001, 2002, 2006, 2013 | | | Layer | Whether seed bank or extant understory | Categorical | Herb (Extant) or Seed  | | | Column B:BY or Columns 2:77 | Species abbreviation and their associated counts. Refer to USDA Plants for greater detail | Numeric | Actaea racemosa = ACRAR; Amphicarpaea bracteata = AMBR2; Aristolochia = ARIST2; Aralia spinosa = ARSP2; Arisaema triphyllum = ARTR; Asarum canadense = ASCA; Aster = ASTER; Berberis thunbergii = BETH; Boehmeria cylindrica = BOCY; Brachyelytrum erectum = BRER2; Cardamine diphylla = CADI10; Caulophyllum thalictroides = CATH2; Collinsonia canadensis = COCA4; Cryptotaenia canadensis = CRCA9; Cuscuta = CUSCU; Dalibarda repens = DARE; Desmodium = DESMO; Dichanthelium clandestinum = DICL; Dioscorea quaternata = DIQU; Dioscorea villosa = DIVI4; Erechtites hieraciifolius = ERHI2; Eurybia divaricata = EUDI16; Eurybia macrophylla = EUMA27; Eupatorium = EUPAT; Festuca = FESTU; Geum canadense = GECA7; Gnaphalium = GNAPH; Grass = GRASS; Hepatica nobilis = HENO2; Hypericum pseudomaculatum = HYPS; Laportea canadensis = LACA3; Luzula multiflora = LUMU2; Lysimachia quadrifolia = LYQU2; Medeola virginiana = MEVI; Monarda = MONAR; Osmorhiza claytonii = OSCL; Osmorhiza longistylis = OSLO; Packera aurea = PAAU3; Panicum = PANIC; Phytolacca americana = PHAM4; Pilea pumila = PIPU2; Polygonatum biflorum = POBI2; Potentilla canadensis = POCA17; Polygonum = POLYG4; Polygonum persicaria = POPE3; Potentilla simplex = POSI2; Poa sylvestris = POSY; Prosartes lanuginosa = PRLA9; Prenanthes trifoliolata = PRTR; Sanicula canadensis = SACA15; Sambucus = SAMBU; Saxifraga = SAXIF; Smilax = SMILA2; Smilax rotundifolia = SMRO; Solanum carolinense = SOCA3; Stellaria = STELL; Stellaria media = STME2; Thalictrum dioicum = THDI; Thalictrum thalictroides = THTH2; Thaspium trifoliatum = THTR; Unknown = UNK; Unknown Graminoid = UNK\_GRAMINOID; Urtica dioica = URDI; Uvularia perfoliata = UVPE; Uvularia = UVULA; Veronica officinalis = VEOF2; Verbascum thapsus = VETH; Verbena urticifolia = VEUR; Viburnum acerifolium = VIAC; Ageratina altissima = AGAL5\_true; Carex = CAREX\_comb; Galium = GALIU\_comb; Rubus = RUBUS\_comb; Solidago = SOLID\_comb; Viola = VIOLA\_comb; Vitis = VITIS\_comb | | | | | | | | | | | | | | bank.long.form\_namedvar | MP | Site (4 total) | Categorical, numeric | Site 1 = 10, Site 2 = 20, Site 3 = 30, Site 4 = 40 | | | SP | Sampling Plot (20 x 20 m disturbed plots with a total of 16 per site) | Categorical, numeric | Plots 1-12 | | | USDA\_CODE2 | USDA species codes. Refer to USDA Plants for greater detail | Categorical | Acer pensylvanicum = ACPE; Acer rubrum = ACRU; Ageratina altissima = AGAL5; Agrostis perennans = AGPE; Andropogon spp. = ANDRO2; Apocynum spp. = APOCY; Aralia spinosa = ARSP2; Berberis thunbergii = BETH; Betula spp. = BETUL; Boehmeria cylindrica = BOCY; Carex spp. = CAREX; Dalibarda repens = DARE; Dichanthelium clandestinum = DICL; Erechtites hieraciifolius = ERHI2; Eupatorium serotinum = EUSE2; Eurybia divaricata = EUDI16l Galium spp. = GALIU; Gaultheria procumbens = GAPR2; Hypericum punctatum = HYPS; Juncus effusus = JEUF; Liriodendron tulipifera = LITU; Lobelia spicata = LOSP; Luzula multiflora = LUMU2; Medeola virginiana = MEVI; Mitchella repens = MIRE; Packera aurea = PAAU3; Phytolacca americana = PHAM4; Pilea pumila = PIPU2; Plantago lanceolata = PLLA; Polygonum persicaria = POPE3; Potentilla canadensis = POCA; Prenanthes trifoliolata = PRTR; Rhus typhina = RHTY; Robinia pseudoacacia = ROPS; Rosa multiflora = ROMU; Rubus spp. = RUBUS; Sambucus spp. = SAMBU; Sassafras albidum = SAAL5; Saxifraga spp. = SAXIF; Sisyrinchium spp. = SISYR; Solanum carolinense = SOCA3; Solanum nigrum = SONI; Stellaria media = STME2; Trillium spp. = TRILL; Uvularia sessilifolia = UVSE; Verbascum thapsus = VETH; Verbena urticifolia = VEUR; Veronica officinalis = VEOF2; Viola blanda = VIBL; Viola canadensis = VICA4; Viola macloskeyi = VIMA2; Viola pubescens = VIPU3; Viola rotundifolia = VIRO2; Viola sororia = VISO; Viola spp. = VIOLA; Vitis aestivalis = VIAE | | | Fire | Burned or unburned plots | Categorical | Fire; No Fire | | | Fence | Deer are fenced out (NoDeer) or deer are present (Deer) | Categorical | Deer; No Deer | | | Gap | Plots with or without canopy gaps | Categorical | Gap; No Gap | | | Sample | Soil sample. Almost always added together in analysis.  | Categorical, numeric | 1, 2, 3 | | | Form | Plant Life Form | Categorical | Forb, Graminoid, Shrub, Tree, Vine | | | Count | Number of germinants | Numeric | |