Weeds, worms, and deer: positive relationships among common forest understory stressors
Abstract
Biotic global change agents, such as non-native plants (‘weeds’), non-native earthworms (‘worms’), and overabundant herbivores (white-tailed ‘deer’), can be major stressors in the forest understory. The status and relationships among these global change stressors across large spatial extents and under naturally varying conditions are poorly understood. Here, through an observational study using a network of U.S. National Park Service forest health monitoring plots (n = 350) from eight parks in seven northeastern states, we modeled causal pathways among global change stressors through model selection in a structural equation (SEM) framework. Weeds, worms, and, deer were common across all parks in the study—46% of plots had non-native plants, 42% of plots had evidence of earthworms, and all parks had plots with high deer browse damage. All biotic global change stressors were significantly and positively correlated with one another (all Spearman rank correlations ≥ 0.44). Consequently, 28% of plots had a combination of earthworms absent, low deer browse, and no non-native plants, and 29% of plots included earthworms, non-native plants, and moderate or greater browse damage. Through SEM, we found strong support for pathways among global change stressors, e.g., deer browse positively influenced earthworm presence and both deer and earthworms promoted non-native plants. Warmer air temperatures and higher soil pH also facilitated non-natives. This research highlights the tremendous…
Integrity signals
Registry
Cites
Statistics
Claims the paper makes
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
Many of the parks experiencing the most severe impacts of chronic deer browse also have the highest and increasing invasive plant loads.
“Many of the parks experiencing the most severe impacts of chronic deer browse also have the highest and increasing invasive plant loads (Fisichelli & Miller, 2018;Miller et al., 2021).”
From Overabundant deer and invasive plants drive widespread regeneration debt in eastern United States national parks (2023) · cites this paper as Fisichelli & Miller, 2018
Cites
Fitting Linear Mixed-Effects Models Using <b>lme4</b>
Bates, D., Mächler, M., Bolker, B., & Walker, S. (2015) · Journal of Statistical Software · doi:10.18637/jss.v067.i01 · Cited by 86,835 · see citing works
Model selection and multimodel inference: a practical information-theoretic approach.
Guthery, F. S., Burnham, K. P., & Anderson, D. R. (2003) · The Journal of Wildlife Management · doi:10.2307/3802723 · Cited by 41,495 · see citing works
Biometry: The Principles and Practice of Statistics in Biological Research
Sokal, R. R., & Rohlf, F. J. (1969) · Cited by 20,963 · see citing works
Nakagawa, S., & Schielzeth, H. (2013) · Methods in Ecology and Evolution · doi:10.1111/j.2041-210x.2012.00261.x · Cited by 10,356 · see citing works
Biometry. The Principles and Practice of Statistics in Biological Research
Fienberg, S. E., Sokal, R. R., Rohlf, F. J., Rohlf, F. J., & Sokal, R. R. (1970) · Biometrics · doi:10.2307/2529087 · Cited by 9,363 · see citing works
BIOTIC INVASIONS: CAUSES, EPIDEMIOLOGY, GLOBAL CONSEQUENCES, AND CONTROL
Mack, R., Simberloff, D., Lonsdale, W., Evans, H., Clout, M., & Bazzaz, F. (2000) · Ecological Applications · doi:10.1890/1051-0761(2000)010[0689:bicegc]2.0.co;2 · Cited by 6,770 · see citing works
The diversity and biogeography of soil bacterial communities
Fierer, N., & Jackson, R. B. (2006) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.0507535103 · Cited by 5,711 · see citing works
piecewiseSEM : Piecewise structural equation modelling in r for ecology, evolution, and systematics
Lefcheck, J. S. (2015) · Methods in Ecology and Evolution · doi:10.1111/2041-210x.12512 · Cited by 4,893 · see citing works
Handbook of Structural Equation Modeling
Howard, A. L. (2013) · Structural Equation Modeling A Multidisciplinary Journal · doi:10.1080/10705511.2013.769397 · Cited by 3,465 · see citing works
Vilà, M., Espinar, J. L., Hejda, M., Hulme, P. E., Jarošík, V., Maron, J. L., Pergl, J., Schaffner, U., Sun, Y., & Pyšek, P. (2011) · Ecology Letters · doi:10.1111/j.1461-0248.2011.01628.x · Cited by 3,068 · see citing works
Structural Equation Modeling and Natural Systems
Grace, J. B. (2006) · doi:10.1017/cbo9780511617799 · Cited by 2,432 · see citing works
Positive Interactions of Nonindigenous Species: Invasional Meltdown?
Simberloff, D., & Von Holle, B. (1999) · Biological Invasions · doi:10.1023/a:1010086329619 · Cited by 2,231 · see citing works
Ecological Impacts of Deer Overabundance
Côté, S. D., Rooney, T. P., Tremblay, J., Dussault, C., & Waller, D. M. (2004) · Annual Review of Ecology Evolution and Systematics · doi:10.1146/annurev.ecolsys.35.021103.105725 · Cited by 2,076 · see citing works
Alien species in a warmer world: risks and opportunities
Walther, G.-R., Roques, A., Hulme, P. E., Sykes, M. T., Pyšek, P., Kühn, I., Zobel, M., Bacher, S., Botta-Dukát, Z., & Bugmann, H. (2009) · Trends in Ecology & Evolution · doi:10.1016/j.tree.2009.06.008 · Cited by 1,424 · see citing works
Harrison, X. A. (2014) · PeerJ · doi:10.7717/peerj.616 · Cited by 1,150 · see citing works
Invasiveness, invasibility and the role of environmental stress in the spread of non-native plants
Alpert, P., Bone, E., & Holzapfel, C. (2000) · Perspectives in Plant Ecology Evolution and Systematics · doi:10.1078/1433-8319-00004 · Cited by 919 · see citing works
Direct and indirect effects of white-tailed deer in forest ecosystems
Rooney, T. P., & Waller, D. M. (2003) · Forest Ecology and Management · doi:10.1016/s0378-1127(03)00130-0 · Cited by 885 · see citing works
Schoenholtz, S. H., Miegroet, H. V., & Burger, J. A. (2000) · Forest Ecology and Management · doi:10.1016/s0378-1127(00)00423-0 · Cited by 829 · see citing works
Guidelines for a graph‐theoretic implementation of structural equation modeling
Grace, J. B., Schoolmaster, D. R., Guntenspergen, G. R., Little, A., MITCHELL, B. R., Miller, K. M., & Schweiger, E. W. (2012) · Ecosphere · doi:10.1890/es12-00048.1 · Cited by 601 · see citing works
Self-citationManaging the whole landscape: historical, hybrid, and novel ecosystems
Hobbs, R. J., Higgs, E. S., Hall, C. M., Bridgewater, P. B., Chapin, F. S., Ellis, E. C., Ewel, J. J., Hallett, L. M., Harris, J., Hulvey, K. B., et al. (2014) · Frontiers in Ecology and the Environment · doi:10.1890/130300 · Cited by 506 · see citing works
The Causal Foundations of Structural Equation Modeling
Pearl, J. (2012) · doi:10.21236/ada557445 · Cited by 359 · see citing works
Earthworm invasion into previously earthworm-free temperate and boreal forests
Frelich, L. E., Hale, C. M., Scheu, S., Holdsworth, A. R., Heneghan, L., Bohlen, P. J., & Reich, P. B. (2006) · Biological Invasions · doi:10.1007/s10530-006-9019-3 · Cited by 336 · see citing works
Indicators of ecological change: new tools for managing populations of large herbivores
Morellet, N., Gaillard, J., Hewison, A. J. M., Ballon, P., Boscardin, Y., Duncan, P., Klein, F., & Maillard, D. (2007) · Journal of Applied Ecology · doi:10.1111/j.1365-2664.2007.01307.x · Cited by 290 · see citing works
Exotic Earthworm Invasions in North America: Ecological and Policy Implications
Hendrix, P. F., & Bohlen, P. J. (2002) · BioScience · doi:10.1641/0006-3568(2002)052[0801:eeiina]2.0.co;2 · Cited by 284 · see citing works
Factors Affecting the Abundance of Earthworms in Soils
Curry, J. (2004) · CRC Press eBooks · doi:10.1201/9781420039719.pt3 · Cited by 278 · see citing works
Ecological Subregions: Sections and Subsections for the conterminous United States
Cleland, D. T., Freeouf, J. A., Keys, J. E., Nowacki, G. J., Carpenter, C. A., & McNab, W. H. (2007) · doi:10.2737/wo-gtr-76d · Cited by 236 · see citing works
A general model to quantify ecological integrity for landscape assessments and US application
Theobald, D. M. (2013) · Landscape Ecology · doi:10.1007/s10980-013-9941-6 · Cited by 201 · see citing works
Nuzzo, V., Maerz, J. C., & Blossey, B. (2009) · Conservation Biology · doi:10.1111/j.1523-1739.2009.01168.x · Cited by 194 · see citing works
Kalisz, S., Spigler, R. B., & Horvitz, C. C. (2014) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1310121111 · Cited by 153 · see citing works
Ecology and management of white‐tailed deer in a changing world
McShea, W. J. (2012) · Annals of the New York Academy of Sciences · doi:10.1111/j.1749-6632.2011.06376.x · Cited by 152 · see citing works
Craven, D., Thakur, M. P., Cameron, E. K., Frelich, L. E., Beauséjour, R., Blair, R. B., Blossey, B., Burtis, J. C., Choi, A., Dávalos, A., et al. (2016) · Global Change Biology · doi:10.1111/gcb.13446 · Cited by 146 · see citing works
Self-citationKourtev, P. S., Huang, W. Z., & Ehrenfeld, J. G. (1999) · Biological Invasions · doi:10.1023/a:1010048909563 · Cited by 143 · see citing works
Patterns Of Litter Disappearance In A Northern Hardwood Forest Invaded By Exotic Earthworms
Suárez, E., Fahey, T. J., Yavitt, J. B., Groffman, P. M., & Bohlen, P. J. (2006) · Ecological Applications · doi:10.1890/04-0788 · Cited by 107 · see citing works
A subcontinental view of forest plant invasions
Oswalt, C. M., Fei, S., Guo, Q., Iannone, B. V., Oswalt, S. N., Pijanowski, B. C., & Potter, K. M. (2015) · NeoBiota · doi:10.3897/neobiota.24.4526 · Cited by 99 · see citing works
Sapling growth responses to warmer temperatures ‘cooled’ by browse pressure
Fisichelli, N. A., Frelich, L. E., & Reich, P. B. (2012) · Global Change Biology · doi:10.1111/j.1365-2486.2012.02785.x · Cited by 97 · see citing works
Self-citationWarren, R. J., Bahn, V., & Bradford, M. A. (2011) · Oikos · doi:10.1111/j.1600-0706.2011.20174.x · Cited by 86 · see citing works
Climate Exposure of US National Parks in a New Era of Change
Monahan, W. B., & Fisichelli, N. A. (2014) · PLoS ONE · doi:10.1371/journal.pone.0101302 · Cited by 82 · see citing works
Self-citationFisichelli, N. A., Frelich, L. E., Reich, P. B., & Eisenhauer, N. (2012) · Biological Invasions · doi:10.1007/s10530-012-0350-6 · Cited by 78 · see citing works
Self-citationRemoval of invasive shrubs reduces exotic earthworm populations
Madritch, M. D., & Lindroth, R. L. (2008) · Biological Invasions · doi:10.1007/s10530-008-9281-7 · Cited by 77 · see citing works
Karberg, N. J., & Lilleskov, E. A. (2008) · Biological Invasions · doi:10.1007/s10530-008-9290-6 · Cited by 71 · see citing works
Dobson, A. M., & Blossey, B. (2014) · Journal of Ecology · doi:10.1111/1365-2745.12350 · Cited by 70 · see citing works
Revisiting Leopold: Resource Stewardship in the National Parks
Colwell, R., Avery, S., Berger, J., Davis, G. E., Hamilton, H., Lovejoy, T. Ε., Malcom, S. M., McMullen, A., Novacek, M. J., Roberts, R. J., et al. (2014) · PARKS · doi:10.2305/iucn.ch.2014.parks-20-2.drc.en · Cited by 63 · see citing works
Single and interactive effects of deer and earthworms on non-native plants
Dávalos, A., Nuzzo, V., & Blossey, B. (2015) · Forest Ecology and Management · doi:10.1016/j.foreco.2015.04.026 · Cited by 57 · see citing works
Waller, D. M., & Maas, L. I. (2013) · Forest Ecology and Management · doi:10.1016/j.foreco.2013.05.011 · Cited by 49 · see citing works
Roth, A. M., Whitfeld, T. J. S., Lodge, A. G., Eisenhauer, N., Frelich, L. E., & Reich, P. B. (2014) · Oecologia · doi:10.1007/s00442-014-3175-4 · Cited by 44 · see citing works
Fisichelli, N. A., Abella, S. R., Peterson, D. L., & Krist, F. J. (2014) · Forest Ecology and Management · doi:10.1016/j.foreco.2014.04.033 · Cited by 44 · see citing works
Self-citationNon-consumptive Effects of Native Deer on Introduced Earthworm Abundance
Dávalos, A., Simpson, E. G., Nuzzo, V., & Blossey, B. (2015) · Ecosystems · doi:10.1007/s10021-015-9881-x · Cited by 43 · see citing works
Interactive effects of deer, earthworms and non-native plants on rare forest plant recruitment
Dávalos, A., Nuzzo, V., & Blossey, B. (2015) · Biological Conservation · doi:10.1016/j.biocon.2015.04.025 · Cited by 43 · see citing works
Bartuszevige, A. M., Hrenko, R. L., & Gorchov, D. L. (2007) · The American Midland Naturalist · doi:10.1674/0003-0031(2007)158[472:eolloe]2.0.co;2 · Cited by 42 · see citing works
Effectiveness of Exotic Plant Treatments on National Park Service Lands in the United States
Abella, S. R. (2014) · Invasive Plant Science and Management · doi:10.1614/ipsm-d-13-00058.1 · Cited by 41 · see citing works
Earthworm Invasions in Northern Hardwood Forests: a Rapid Assessment Method
Loss, S. R., Hueffmeier, R. M., Hale, C. M., Host, G. E., Sjerven, G., & Frelich, L. E. (2013) · Natural Areas Journal · doi:10.3375/043.033.0103 · Cited by 37 · see citing works
Miller, K. M., Dieffenbach, F. W., Campbell, J. P., Cass, W. B., Comiskey, J. A., Matthews, E. R., McGill, B. J., Mitchell, B. R., Perles, S. J., Sanders, S., et al. (2016) · Ecosphere · doi:10.1002/ecs2.1404 · Cited by 29 · see citing works
Self-citationRearick, D. C., Kintz, L., Burke, K. L., & Ransom, T. S. (2011) · Pedobiologia · doi:10.1016/j.pedobi.2011.09.001 · Cited by 18 · see citing works
Flagged first, then by citations
Cited by
Sorted by citations
Gorchov, D. L., Blossey, B., Averill, K. M., Dávalos, A., Heberling, J. M., Jenkins, M. A., Kalisz, S., McShea, W. J., Morrison, J. A., Nuzzo, V., et al. (2021) · Biological Invasions · doi:10.1007/s10530-021-02551-2 · Cited by 56 · see citing works
Miller, K. M., Perles, S. J., Schmit, J. P., Matthews, E. R., Weed, A. S., Comiskey, J. A., Marshall, M. R., Nelson, P., & Fisichelli, N. A. (2023) · Ecological Applications · doi:10.1002/eap.2837 · Cited by 45 · see citing works
Self-citationLand use change and forest management effects on soil carbon stocks in the Northeast U.S.
Nave, L. E., DeLyser, K., Domke, G. M., Holub, S. M., Janowiak, M. K., Keller, A. B., Peterson, D. L., Solarik, K. A., Walters, B. F., & Swanston, C. W. (2024) · Carbon Balance and Management · doi:10.1186/s13021-024-00251-7 · Cited by 32 · see citing works
FLORISTIC CHANGE IN NEW ENGLAND AND NEW YORK: REGIONAL PATTERNS OF PLANT SPECIES LOSS AND DECLINE
MacKenzie, C. M., Mittelhauser, G. H., Miller‐Rushing, A. J., & Primack, R. B. (2019) · Rhodora · doi:10.3119/18-04 · Cited by 25 · see citing works
Abas, A. (2023) · Frontiers in Environmental Science · doi:10.3389/fenvs.2023.970730 · Cited by 18 · see citing works
The effects of deer and an invasive shrub, Lonicera maackii, on forest understory plant composition
Haffey, C. M., & Gorchov, D. L. (2019) · Ecoscience · doi:10.1080/11956860.2019.1582195 · Cited by 15 · see citing works
Linked disturbance in the temperate forest: Earthworms, deer, and canopy gaps
Reed, S. P., Bronson, D. R., Forrester, J. A., Prudent, L. M., Yang, A. M., Yantes, A. M., Reich, P. B., & Frelich, L. E. (2023) · Ecology · doi:10.1002/ecy.4040 · Cited by 12 · see citing works
Two Centuries of Change in the Native Flora of Franklin County, Massachusetts, U.S.A.
Bertin, R., Searcy, K., Motzkin, G., Hickler, M., & Grima, P. P. (2022) · Rhodora · doi:10.3119/21-18 · Cited by 7 · see citing works
Multiple stressors prevent gains in native plant diversity following invasive species removal
Collings, J. A., Endriss, S. B., & Dávalos, A. (2023) · Ecosphere · doi:10.1002/ecs2.4458 · Cited by 6 · see citing works
Donoso, M. U., Leonard, H., & Gorchov, D. L. (2024) · Invasive Plant Science and Management · doi:10.1017/inp.2024.2 · Cited by 6 · see citing works
Abella, S. R., Schetter, T. A., & Gallaher, T. D. (2022) · Wildlife Society Bulletin · doi:10.1002/wsb.1377 · Cited by 4 · see citing works
Non-Native Earthworms Invade Forest Soils in Northern Maine, USA
Puhlick, J. J., Fernandez, I. J., & Wason, J. (2021) · Forests · doi:10.3390/f12010080 · Cited by 4 · see citing works
Morrison, J. A., Roche, B. M., & Veatch‐Blohm, M. E. (2022) · Ecology and Evolution · doi:10.1002/ece3.8814 · Cited by 4 · see citing works
Lima, K. A., Wheeler, J. S., Connery, J. H., Miller‐Rushing, A. J., & Fisichelli, N. A. (2024) · Journal of Applied Ecology · doi:10.1111/1365-2664.14835 · Cited by 3 · see citing works
Self-citationNagele, E., Ashton, M. S., Harris, T. B., Forrest, T. A., DeLuca, E. A., Martello, M. E., & Duguid, M. C. (2024) · Journal of Forestry · doi:10.1093/jofore/fvad057 · Cited by 3 · see citing works
Heimburger, B., Klein, A., Roth, A. M., Scheu, S., Eisenhauer, N., & Schaefer, I. (2023) · Biological Invasions · doi:10.1007/s10530-023-03048-w · Cited by 2 · see citing works
Patterns and changes in the Nonnative Flora of Franklin County, Massachusetts
Bertin, R. I., Searcy, K. B., Motzkin, G., & Hickler, M. G. (2024) · Rhodora · doi:10.3119/23-01 · Cited by 2 · see citing works
Hawcroft, A., Bellingham, P. J., Jo, I., Richardson, S. J., & Wright, E. F. (2024) · Biological Conservation · doi:10.1016/j.biocon.2024.110637 · Cited by 2 · see citing works
Truchon, F. (2021) · eScholarship@McGill (McGill) · doi:10.82308/45733
Nave, L. E., DeLyser, K., Domke, G. M., Holub, S. M., Janowiak, M. K., Keller, A. B., Peters, M. P., Solarik, K. A., Walters, B. F., & Swanston, C. W. (2024) · Figshare · doi:10.6084/m9.figshare.26677547
Nave, L. E., DeLyser, K., Domke, G. M., Holub, S. M., Janowiak, M. K., Keller, A. B., Peters, M. P., Solarik, K. A., Walters, B. F., & Swanston, C. W. (2024) · Figshare · doi:10.6084/m9.figshare.26677544
Native earthworms and roe deer in temperate forests: Friends in the sun, foes in the shade
Lettenmaier, L., Eisenhauer, N., Cesarz, S., Ammer, C., Bucharová, A., Dittrich, S., Nußer, R., Kraus, D., Kölbl, A., Mallick, S., et al. (2026) · Ecology · doi:10.1002/ecy.70520
- Authors
- Fisichelli, N. A., & Miller, K. M.
- Affiliations
- Schoodic Institute (US)
- National Park Service (US)
- Corresponding author
- Fisichelli, Nicholas A.
- Venue
- Biological Invasions
- Published
- 2017-11-17
- Volume
- 20
- Issue
- 5
- Pages
- 1337-1348
- ISSN
- 1387-3547 (ISSN-L)
- 1573-1464
- OpenAlex
- W2770409279
- Cited by
- 21
- References
- 54
- Citations by year
- 2026: 1 · 2024: 7 · 2023: 5 · 2022: 3 · 2021: 3 · 2019: 2
- FWCI
- 0.45
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- Biological Invasions (publishedVersion)
- Funders
- Maine Timberlands Charitable Trust
- Topics
- Physical Sciences › Environmental Science › Nature and Landscape Conservation › Ecology and Vegetation Dynamics Studies
- Physical Sciences › Environmental Science › Ecology › Animal Ecology and Behavior Studies
- Physical Sciences › Environmental Science › Ecology › Forest Insect Ecology and Management
- Keywords
- understory vegetation, non-native plants, earthworms, white-tailed deer, invasive species, global warming, soil pH, forest health, causal pathways, national parks, northeastern United States
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
Are you an author, or do you see a problem with this page? Request a review