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Can’t see the flowers for the trees: factors driving floral abundance within early-successional forests in the central Appalachian Mountains

Publication: Canadian Journal of Forest Research
3 August 2022

Abstract

Silviculture can be a powerful tool for restoring and enhancing habitat for forest-dependent wildlife. In eastern North America, regenerating timber harvests support abundant wildflowers that provide essential forage for native pollinators. Factors driving floral resource availability within regenerating forests remain almost entirely unstudied. Recent efforts to increase the area of regenerating forests (<10 years old) through overstory removal harvest in the central Appalachian Mountains provide an opportunity to investigate the development of forest wildflower communities following canopy removal. We conducted 1208 surveys of blooming plants across 143 harvests, recording 1 525245 flowers representing 220 taxa spanning 47 families. The number of flowers within recently harvested stands was negatively associated with fern and sapling cover but positively associated with grass and bramble (Rubus spp.) cover. Early in the growing season, more flowers bloomed in older regenerating stands (e.g., >5 years old), but this pattern reversed by the end of the growing season. Ultimately, our study demonstrates that the abundance of flowers available to pollinators within regenerating hardwood stands varies with factors associated with advancing succession. Recognizing the potential trade-off between woody regeneration (i.e., saplings) and pollinator forage availability may benefit forest managers who intend to provide floral resources to flower-dependent wildlife like pollinators via silviculture.

Résumé

La sylviculture peut être un outil précieux pour restaurer et améliorer l'habitat des espèces sauvages dépendant de la forêt. Dans l'est de l'Amérique du Nord, les coupes de bois en régénération abritent d'abondantes fleurs sauvages qui fournissent un fourrage essentiel aux pollinisateurs naturels. Les facteurs qui déterminent la disponibilité des ressources florales dans les forêts en régénération n'ont pratiquement pas été étudiés. Les efforts récents visant à augmenter la superficie des forêts en régénération (< 10 ans) par l'enlèvement de l'étage dominant dans les Appalaches centrales offrent l'occasion d'étudier le développement des communautés de fleurs sauvages forestières après l'élimination du couvert. Nous avons effectué 1 208 relevés de plantes en fleurs dans 143 récoltes, enregistrant 1 525 245 fleurs représentant 220 taxons répartis dans 47 familles. Le nombre de fleurs dans les peuplements récemment récoltés était négativement associé à la couverture de fougères et de jeunes arbres, mais positivement associé à la couverture d'herbes et de ronces (Rubus spp.). Au début de la saison de croissance, davantage de fleurs ont fleuri dans les peuplements en régénération plus âgés (par exemple, > 5 ans), mais cette tendance s'est inversée à la fin de la saison de croissance. En somme, notre étude démontre que l'abondance des fleurs disponibles pour les pollinisateurs dans les peuplements de feuillus en régénération varie en fonction des facteurs associés à l'avancement de la succession. La reconnaissance du compromis potentiel entre la régénération ligneuse (c'est-à-dire les jeunes arbres) et la disponibilité du fourrage pour les pollinisateurs peut profiter aux gestionnaires forestiers qui ont l'intention de fournir des ressources florales à la faune dépendant des fleurs, comme les pollinisateurs, par le biais de la sylviculture. [Traduit par la Rédaction]

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References

Albright T.A., McWilliams W.H., Widmann R.H., Butler B.J., Crocker S.J., Kurtz C.M., 2017. Pennsylvania forests 2014. Resour. Bull. NRS-111. Newtown Square, PA: US Department of Agriculture, Forest Service, Northern Research Station.
Arnold T.W. 2010. Uninformative parameters and model selection using Akaike's information criterion. J. Wildl. Manage. 74: 1175–1178.
Askins R.A. 2001. Sustaining biological diversity in early successional communities: the challenge of managing unpopular habitats. Wildl. Soc. Bull. 29(2): 407–412.
Bates D., Maechler M., Bolker B., Walker S. 2015. Fitting linear mixed-effects models using lme4. J. Stat. Softw. 67(1), 1–48.
Bolker B.M., Brooks M.E., Clark C.J., Geange S.W., Poulsen J.R., Stevens M.H.H.,. 2009. Generalized linear mixed models: a practical guide for ecology and evolution. Trends Ecol. Evol. 24(3): 127–135.
Borcard D., Legendre P., Gillet F. 2011. Numerical Ecology in R. Springer, New York.
Burnham K.P., Anderson D.R. 2002. A practical information-theoretic approach. Model selection and multimodel inference. 2nd ed. Springer, New York.
Chips M.J., Yerger E.H., Hervanek A., Nuttle T., Royo A.A., Pruitt J.N., 2015. The indirect impact of long-term overbrowsing on insects in the Allegheny National Forest region of Pennsylvania. Northeastern Nat. 22(4): 782–797.
Clarke K.R., Ainsworth M. 1993. A method of linking multivariate community structure to environmental variables. Mar Ecol Progr Ser. 92: 205–205.
de Jesus Jatoba L., Varela R.M., Molinillo J.M. Gonzalez, Din Z. Ud, Gualtieri S.C. Juliano, Rodrigues-Filho E., Macías F.A. 2016. Allelopathy of bracken fern (Pteridium arachnoideum): new evidence from green fronds, litter, and soil. PLoS ONE, 11(8):.
deCalesta D.S. 1994. Effect of white-tailed deer on songbirds within managed forests in Pennsylvania. J. Wildl. Manag. 58: 711–718.
Department of Conservation & Natural Resources (DCNR). 2020. DCNR invasive plant list. Pennsylvania Department of Conservation & Natural Resources. Available from http://elibrary.dcnr.pa.gov/GetDocument?docId=2700788&DocName=dcnr_20033786 [accessed 29 December 2020].
Donnelly J.P., Tack J.D., Doherty K.E., Naugle D.E., Allred B.W., Dreitz V.J. 2017. Extending conifer removal and landscape protection strategies from sage-grouse to songbirds, a range-wide assessment. Rangel. Ecol. Manag. 70(1): 95–105.
Duguid M.C., Ashton M.S. 2013. A meta-analysis of the effect of forest management for timber on understory plant species diversity in temperate forests. For. Ecol. Manag. 303: 81–90.
Eisenhauer N., Yee K., Johnson E.A., Maraun M., Parkinson D., Straube D., 2011. Positive relationship between herbaceous layer diversity and the performance of soil biota in a temperate forest. Soil Biol. Biochem. 43(2): 462–465.
Elmore A.J., Nelson D.M., Craine J.M. 2016. Earlier springs are causing reduced nitrogen availability in North American eastern deciduous forests. Nat. Plants. 2(10): 1–5.
ESRI. 2011. Release 10. Documentation manual. Environmental Systems Research Institute. Redlands, CA.
Fredericksen T.S., Ross B.D., Hoffman W., Morrison M.L., Beyea J., Johnson B.N., 1999. Short-term understory plant community responses to timber-harvesting intensity on non-industrial private forestlands in Pennsylvania. For. Ecol. Manag. 116(1-3): 129–139.
George L.O., Bazzaz F.A. 2014. The herbaceous layer as a filter determining spatial pattern in forest tree regeneration. In The herbaceous layer in forests of Eastern North America. Edited by Gilliam Frank. Oxford University Press, Oxford UK. pp. 340–355.
Gilliam F.S. 2007. The ecological significance of the herbaceous layer in temperate forest ecosystems. Bioscience, 57(10): 845–858.
Gilliam F.S. 2016. Forest ecosystems of temperate climatic regions: from ancient use to climate change. New Phytol. 212(4): 871–887.
Gilliam F.S.(Ed.). 2014. The herbaceous layer in forests of eastern North America. Oxford University Press, Oxford, UK.
Govaert S., Vangansbeke P., Blondeel H., De Lombaerde E., Verheyen K., De Frenne P. 2021. Forest understorey plant responses to long-term experimental warming, light and nitrogen addition. Plant Biol. 23: 1051–-1062.
Hanula J.L., Ulyshen M.D., Horn S. 2016. Conserving pollinators in North American forests: a review. Nat. Areas J. 36(4): 427–439.
Heil L.J., Burkle L.A. 2018. Recent post-wildfire salvage logging benefits local and landscape floral and bee communities. For. Ecol. Manag. 424: 267–275.
Hilty J., Merenlender A.M. 2003. Studying biodiversity on private lands. Conserv. Biol. 17(1): 132–137.
Hunter W.C., Buehler D.A., Canterbury R.A., Confer J.L., Hamel P.B., 2001. Conservation of disturbance-dependent birds in eastern North America. Wildl. Soc. Bull. 29: 440–455.
Hutchinson T.F. 2006. Fire and the herbaceous layer of eastern oak forests. Fire in Eastern Oak Forests: delivering science to land managers. USDA Forest Service, Northern Research Station, Columbus, OH, pp. 136–149.
Jackson M.M., Turner M.G., Pearson S.M. 2014. Logging legacies affect insect pollinator communities in southern Appalachian forests. Southeastern Nat. 13(2): 317–336.
James F.C., Shugart Jr H.H. 1970. A quantitative method of habitat description. Audubon Field Notes, 24(6): 727–736.
Joshi N.K., Otieno M., Rajotte E.G., Fleischer S.J., Biddinger D.J. 2016. Proximity to woodland and landscape structure drives pollinator visitation in apple orchard ecosystem. Front. Ecol. Evol. 4: 38.
Jost L. 2006. Entropy and diversity. Oikos, 113(2): 363–375.
King D.I., Schlossberg S. 2014. Synthesis of the conservation value of the early-successional stage in forests of eastern North America. For. Ecol. Manag. 324: 186–195.
Klaus F., Tscharntke T., Uhler J., Grass I. 2021. Calcareous grassland fragments as sources of bee pollinators for the surrounding agricultural landscape. Glob. Ecol. Conserv. 26: e01474.
Lake J.C., Leishman M.R. 2004. Invasion success of exotic plants in natural ecosystems: the role of disturbance, plant attributes and freedom from herbivores. Biol. Conserv. 117(2): 215–226.
Lee M.R., McNeil Jr D.J., Mathis C.L., Grozinger C.M., Larkin J.L. 2021. Microhabitats created by log landings support abundant flowers and insect pollinators within regenerating mixed-oak stands in the Central Appalachian Mountains. For. Ecol. Manag. 497: 119472.
Leong M., Ponisio L.C., Kremen C., Thorp R.W., Roderick G.K. 2016. Temporal dynamics influenced by global change: bee community phenology in urban, agricultural, and natural landscapes. Glob. Change Biol. 22(3): 1046–1053.
Litvaitis J.A. 1993. Response of early successional vertebrates to historic changes in land use. Conserv. Biol. 7(4): 866–873.
Litvaitis J.A., Larkin J.L., McNeil D.J., Keirstead D., Costanzo B. 2021. Addressing the early-successional habitat needs of at-risk species on privately owned lands in the Eastern United States. Land, 10(11): 1116.
Mathis C.L., McNeil Jr D.J., Lee M.R., Grozinger C.M., King D.I., Otto C.R., 2021. Pollinator communities vary with vegetation structure and time since management within regenerating timber harvests of the Central Appalachian Mountains. For. Ecol. Manag., 496: 119373.
McNeil D.J., Fiss C.J., Wood E.M., Duchamp J.E., Bakermans M.J., Larkin J.L. 2018. Using a natural reference system to evaluate songbird habitat restoration. Avian Conserv Ecol. 13(1): 22.
McNeil D.J., Rodewald A.D., Robinson O.J., Fiss C.J., Rosenberg K.V., Ruiz-Gutierrez V., 2020. Regional abundance and local breeding productivity explain occupancy of restored habitats in a migratory songbird. Biol. Conserv. 245: 108463.
Milam J., Cunningham-Minnick M., Roberts H.P., Buelow C., King D.I. 2022. The contribution of canopy samples to assessments of forestry effects on native bees. Conserv. Sci. Pract. 4(6): e12690.
Minogue P.J., Cantrell R.L., Griswold H.C. 1991. Vegetation management after plantation establishment. In Forest regeneration manual. Springer, Dordrecht. pp. 335–358.
Mola J.M., Hemberger J., Kochanski J., Richardson L.L., Pearse I.S. 2021. The importance of forests in bumble bee biology and conservation. Bioscience, 71(12): 1234–1248.
Muller R.N. 2014. Nutrient relations of the herbaceous layer in deciduous forest ecosystems. In The herbaceous layer in forests of Eastern North America. 2nd ed. Edited by F.S. Gilliam, M.R. Roberts. Oxford University Press, New York. pp. 13–34.
Naiman R.J., Johnston C.A., Kelley J.C. 1988. Alteration of North American streams by beaver. BioScience, 38(11): 753–762.
Oksanen J., Blanchet F.G., Kindt R., Legendre P., Minchin D., O'Hara R.B.,. 2013. vegan: community ecology package. Version 2(9), pp. 1–295
Padmanaba M., Corlett R.T. 2014. Minimizing risks of invasive alien plant species in tropical production forest management. Forests, 5(8): 1982–1998.
Pastor J., Naiman R.J. 1992. Selective foraging and ecosystem processes in boreal forests. Am. Nat. 139(4): 690–705.
Pollinator Health Task Force (PHTF). May 2015. National strategy to promote the health of honey bees and other pollinators. The White House, Washington, D.C. Available from https://obamawhitehouse.archives.gov/sites/default/files/microsites/ostp/Pollinator%20Health%20Strategy%202015.pdf [accessed January 2021].
Potts S.G., Biesmeijer J.C., Kremen C., Neumann P., Schweiger O., Kunin W.E. 2010. Global pollinator declines: trends, impacts and drivers. Trends Ecol. Evol., 25(6): 345–353.
R Core Team. 2020. R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available from https://www.R-project.org/
Ristau T.E., Stoleson S.H., Horsley S.B., deCalesta D.S. 2011. Ten-year response of the herbaceous layer to an operational herbicide-shelterwood treatment in a northern hardwood forest. For. Ecol. Manag., 262(6): 970–979.
Rivers J.W., Betts M.G. 2021. Postharvest bee diversity is high but declines rapidly with stand age in regenerating douglas-fir forest. For. Sci. 67(3): 275–285.
Rivers J.W., Mathis C.L., Moldenke A.R., Betts M.G. 2018. Wild bee diversity is enhanced by experimental removal of timber harvest residue within intensively managed conifer forest. GCB Bioenergy, 10(10): 766–781.
Roberts M.R. 2004. Response of the herbaceous layer to natural disturbance in North American forests. Can. J. Bot., 82(9): 1273–1283.
Roberts M.R., Gilliam F.S. 2014. Response of the herbaceous layer to disturbance in eastern forests. The herbaceous layer in forests of Eastern North America. 2nd ed. Edited by Gilliam F.S., and Roberts M.R. Oxford University Press, Oxford, UK. pp. 321–339.
Rooney T.P., Waller D.M. 2003. Direct and indirect effects of white-tailed deer in forest ecosystems. For. Ecol. Manag. 181(1-2): 165–176.
Roulston T.A.H., Goodell K. 2011. The role of resources and risks in regulating wild bee populations. Annu. Rev. Entomol. 56: 293–312.
Royo A.A., Collins R., Adams M.B., Kirschbaum C., Carson W.P. 2010. Pervasive interactions between ungulate browsers and disturbance regimes promote temperate forest herbaceous diversity. Ecology, 91(1): 93–105.
Saunders M.E. 2018. Insect pollinators collect pollen from wind-pollinated plants: implications for pollination ecology and sustainable agriculture. Insect Conserv. Divers. 11(1): 13–31.
Sevon W.D. 2000. Physiographic provinces of Pennsylvania, Map 13. Pennsylvania Geologic Survey, Harrisburg, PA.
Shifley S.R., Moser W.K., Nowak D.J., Miles P.D., Butler B.J., Aguilar F.X., 2014. Five anthropogenic factors that will radically alter forest conditions and management needs in the northern United States. For. Sci. 60(5): 914–925.
Smith D.M., Larson B.C., Kelty M.J., Ashton P.M.S. 1997. The practice of silviculture: applied forest ecology (No. Ed. 9). John Wiley and Sons, New York.
Sokal R.R., Rohlf F.J. 1981. Biometry: the principles and practice of statistics in biological research. W.H. Freeman & Company, San Francisco.
Stapanian M.A., Sundberg S.D., Baumgardner G.A., Liston A. 1998. Alien plant species composition and association with anthropogenic disturbance in North American forests. Plant Ecol. 139: 49–62.
Swanson M.E., Franklin J.F., Beschta R.L., Crisafulli C.M., DellaSala D.A., Hutto R.L., 2011. The forgotten stage of forest succession: early-successional ecosystems on forest sites. Front. Ecol. Environ. 9(2): 117–125.
Trillo A., Montero-Castaño A., Vilà M. 2020. Seasonality of bumblebee spillover between strawberry crops and adjacent pinewoods. Apidologie, 51(6): 1051–1061.
Trumbore S., Brando P., Hartmann H. 2015. Forest health and global change. Science, 349(6250): 814–818.
Van Lear D.H., Harlow R.F. 2002. Fire in the eastern United States: influence on wildlife habitat. In Proceedings: the role of fire for nongame wildlife management and community restoration: traditional uses and new directions. Edited by W. Mark Ford, Kevin R. Russell, Christopher E. Moorman. Gen. Tech. Rep. NE-288. U.S. Department of Agriculture, Newtown Square, PA. Forest Service, Northeastern Research Station. 2-10., 288.
Willems F.M., Scheepens J.F., Ammer C., Block S., Bucharova A., Schall P., 2019. Effects of forest management on the phenology of early-flowering understory herbs. bioRxiv, 733907.
Willmer P. 2011. Pollination and floral ecology. Princeton University Press, Princeton, NJ.

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Information & Authors

Information

Published In

cover image Canadian Journal of Forest Research
Canadian Journal of Forest Research
Volume 52Number 7July 2022
Pages: 1002 - 1013

History

Received: 20 January 2022
Accepted: 11 May 2022
Accepted manuscript online: 7 June 2022
Version of record online: 3 August 2022

Data Availability Statement

Following requirements from our funding source, our data are not made available to protect the privacy of the landowners on whose land our research was conducted. Public lands data set can be made available upon request (see corresponding author).

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Key Words

  1. flower communities
  2. blooming
  3. regenerating forests
  4. forest management
  5. timber harvest
  6. overstory removal
  7. Pennsylvania
  8. Appalachian
  9. pollinator

Mots-clés

  1. communautés florales
  2. floraison
  3. forêts en régénération
  4. gestion forestière
  5. récolte de bois
  6. suppression de l'étage dominant
  7. Pennsylvanie
  8. Appalaches
  9. pollinisateur

Authors

Affiliations

Department of Biology, Indiana University of Pennsylvania, 1011 South Drive, Indiana, PA 15705, USA
Author Contributions: Conceptualization, Methodology, Formal analysis, Investigation, Data curation, Writing – original draft, Writing – review & editing, and Visualization.
Present address for Codey L. Mathis is Department of Entomology, Pennsylvania State University, University Park, PA, 16801, USA.
Darin J. McNeil Jr.
Department of Environmental Sciences, University of North Carolina Wilmington, 601 College Road, Wilmington, NC 28403, USA
Author Contributions: Conceptualization, Methodology, Formal analysis, Investigation, Writing – review & editing, Visualization, and Supervision.
Monica R. Lee
Department of Biology, Indiana University of Pennsylvania, 1011 South Drive, Indiana, PA 15705, USA
Author Contributions: Investigation, Data curation, and Writing – review & editing.
Christina M. Grozinger
Department of Entomology, Center for Pollinator Research, Huck Institutes of the Life Sciences, Pennsylvania State University, University Park, PA 16801, USA
Author Contributions: Writing – review & editing and Supervision.
Clint R.V. Otto
Northern Prairie Wildlife Research Center, US Geological Survey, 8711 37th Street SE, Jamestown, ND 58401, USA
Author Contributions: Conceptualization, Methodology, and Writing – review & editing.
Jeffery L. Larkin
Department of Biology, Indiana University of Pennsylvania, 1011 South Drive, Indiana, PA 15705, USA
Author Contributions: Conceptualization, Methodology, Writing – review & editing, Supervision, Project administration, and Funding acquisition.

Author Contributions

Codey L. Mathis: Conceptualization, Methodology, Formal Analysis, Investigation, Data Curation, Writing — Original Draft, Writing — Review and Editing, Visualization. Darin J. McNeil, Jr.: Conceptualization, Methodology, Formal Analysis, Investigation, Writing — Review and Editing, Visualization, Supervision. Monica R. Lee: Investigation, Data Curation, Writing — Review and Editing. Christina M. Grozinger: Writing — Review and Editing, Supervision. Clint R.V. Otto: Conceptualization, Methodology, Writing — Review and Editing. Jeffrey L. Larkin: Conceptualization, Methodology, Writing — Review and Editing, Supervision, Project Administration, Funding Acquisition.

Competing Interests

The authors declare there are no competing interests.

Funding Information

This research was funded through USDA Natural Resource Conservation Service's Conservation Effects Assessment Project (No. 68-7482-12-502) and Indiana University of Pennsylvania's School of Graduate Studies and Research. Our funding sponsors had no role in the study design, collection, analysis, and interpretation of data, report writing, nor the decision to submit the article for publication.

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