Biological Invasions

, Volume 1, Issue 2–3, pp 223–236

Establishment and Control of Hay-scented Fern: A Native Invasive Species

  • Avril L. de la Cretaz
  • Matthew J. Kelty


Hay-scented fern (Dennstaedtia punctilobula (Michx.) Moore) is a native forest understory species that behaves as an invasive plant under certain conditions. Previous work has shown that both increased understory light intensity following overstory thinning and removal of competing plants by herbivores can lead to accelerated growth of hay-scented fern, allowing it to develop dense, nearly monospecific understories that inhibit tree seedling regeneration. To study the relationship between these two factors, we sampled 28 forest stands thinned at different times and subjected to different levels of browsing by white-tailed deer (Odocoileus virginianus), and concluded that more than 15 years of intensive browsing following thinning was necessary for ferns to form closed understory canopies with densities of >90 fronds/m2 and canopy heights of 60–80 cm; thinning alone or intensive browsing alone was not sufficient to cause this level of fern invasion. We applied three treatments to dense fern understories to determine the relative importance of the fern canopy and the dense mat of roots, rhizomes, and dead fronds in the inhibition of tree seedling establishment. Results after two years were: (1) complete removal of the organic mat produced a large germination response of woody and herbaceous species; (2) mixing the organic mat into the mineral soil produced an initial germination response but poor seedling survivorship, as the fern canopy regrew to near pretreatment density; (3) repeatedly clipping the ferns for two years without disturbing the organic mat resulted in a lower germination response than the removal treatment, but rapid growth of seedlings.

deer browse restoration ecology understory regeneration 


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Barten PK, Snowman TK, Lyons PJ, Mahlstedt T, O'Connor R and Spencer BA (1998) Managing a watershed protection forest. Journal of Forestry 96(8): 10–15Google Scholar
  2. Bohm BA and Tryon RM (1967) Phenolic compounds in ferns. 1. A survey of some ferns for cinnamic acid and benzoic acid derivatives. Canadian Journal of Botany 45: 585–593Google Scholar
  3. Cody WJ, Hall IV and Crompton CW (1977) The biology of Canadian weeds. 26. Dennstaedtia punctilobula (Michx.) Moore. Canadian Journal of Plant Science 57: 1159–1168Google Scholar
  4. Cohen AL, Singhakumara BMP and Ashton PMS (1995) Releasing rain forest succession: a case study in the Dicranopteris linearis fernlands of Sri Lanka. Restoration Ecology 3(4): 261–270Google Scholar
  5. Conard HS (1908) The Structure and Life History of the Hay-scented Fern. Carnegie Institution of Washington, Washington, DC, 56 ppGoogle Scholar
  6. Drew AP (1988) Interference of black cherry by ground flora of the Allegheny uplands. Canadian Journal of Forest Research 18: 652–656Google Scholar
  7. Fenner M (1985) Seed Ecology. Chapman and Hall, London, 151 ppGoogle Scholar
  8. George LO and FA Bazzaz (1999) The fern understory as an ecological filter: emergence and establishment of canopy-tree seedlings. Ecology 80: 833–845Google Scholar
  9. Groninger JW and McCormick LH (1991) Effects of sulfometuron on hay-scented fern spore emergence. Canadian Journal of Forest Research 21: 942–943Google Scholar
  10. Groninger JW and McCormick LH (1992) Effects of soil disturbance on hay-scented fern establishment. Northern Journal of Applied Forestry 9(1): 29–31Google Scholar
  11. Hammen SCL (1993) Density-dependent phenotypic variation in the hay-scented fern, Dennstaedtia punctilobula. Bulletin of the Torrey Botanical Club 120(4): 392–396Google Scholar
  12. Hart DH and Sharpe WE (1997) Response of potted northern red oak and hay-scented fern to additions of calcium, magnesium, potassium, and phosphorus. In: Proceedings of the 11th Central Hardwood Conference, Columbia, Missouri, USA, 23–26 March 1997, USDA Forest Service General Technical Report NC-188, pp 304–312Google Scholar
  13. Horsley SB (1984) Hay-scented fern rhizome development in uncut and thinned Allegheny hardwood stands. American Journal of Botany 71 (5, part 2): 80–81Google Scholar
  14. Horsley SB (1988) Control of understory vegetation with Oust. Northern Journal of Applied Forestry 5: 261–262Google Scholar
  15. Horsley SB (1993a) Role of allelopathy in hay-scented fern interference with black cherry regeneration. Journal of Chemical Ecology 19(11): 2737–2755Google Scholar
  16. Horsley SB (1993b) Mechanisms of interference between hay-scented fern and black cherry. Canadian Journal of Forestry Research 23: 2059–2069Google Scholar
  17. Horsley SB and Marquis DA (1983) Interference by weeds and deer with Allegheny hardwood reproduction. Canadian Journal of Forestry Research 13: 61–69Google Scholar
  18. Hughes JW and Fahey TJ (1991) Colonization dynamics of herbs and shrubs in a disturbed northern hardwood forest. Journal of Ecology 79: 605–616Google Scholar
  19. Jones WR (1951) The deer herd of the Quabbin Reservation and its ecology. MS Thesis, Department of Forestry and Wildlife Management, University of Massachusetts, Amherst, Massachusetts, 126 ppGoogle Scholar
  20. Kolb TE, Bowersox TW, McCormick LH and Steiner KC (1989) Effects of shade and herbaceous vegetation on first-year germination and growth of direct-seeded northern red oak, white ash, white pine, and yellow-poplar. In: Proceedings of the 7th Central Hardwood Conference, Carbondale, Illinois, USA, 5–8 March 1989, USDA Forest Service General Technical Report NC-132, pp 156–161Google Scholar
  21. Kyker-Snowman T (1989) Quabbin forest regeneration study. Metropolitan District Commission, Division of Watershed Management, Quabbin Section, Belchertown, Massachusetts, 34 ppGoogle Scholar
  22. Leck MA and Simpson RL (1987) Spore bank of a Delaware river freshwater tidal wetland. Bulletin of the Torrey Botanical Club 114(1): 1–7Google Scholar
  23. Lyon J and Sharpe WE (1996) Hay-scented fern (Dennstaedtia punctilobula (Michx.) Moore) interference with growth of northern red oak (Quercus rubra L.) seedlings. Tree Physiology 16: 923–932Google Scholar
  24. Marquis DA, Grisez TJ, Bjorkbom JC and Roach BA (1975) Interim guide to regeneration of Allegheny hardwoods. USDA Forest Service General Technical Report NE-19, 14 ppGoogle Scholar
  25. Marrs RH, Johnson SW and Le Duc MG (1998) Control of bracken and restoration of heathland. VIII. The regeneration of the heathland community after 18 years of continued bracken control or 6 years of control followed by recovery. Journal of Applied Ecology 35: 857–870Google Scholar
  26. Mayer AM and Poljakoff-Mayber A (1982) The Germination of Seeds (Third Edition). Pergamon Press, Oxford, UK, 211 ppGoogle Scholar
  27. McCormick LH and Bowersox TW (1997) Grass or fern competition reduce growth and survival of planted tree seedlings. In: Proceedings of 11th Central Hardwood Conference, Columbia, Missouri, 23–26 March 1997, USDA Forest Service General Technical Report NC-188, pp 286–293Google Scholar
  28. Messier C and Puttonen P (1995) Spatial and temporal variation in the light environment of developing scots pine stands: the basis for a quick and efficient method of characterizing light. Canadian Journal of Forestry Research 25: 343–354Google Scholar
  29. O'Connor RC, Kyker-Snowman T, Lyons P and Spencer B (1995) Quabbin watershed: MDC land management plan, 1995–2004. Commonwealth of Massachusetts, Metropolitan District Commission Publication, 183 ppGoogle Scholar
  30. Penrod KA and McCormick LH (1996) Abundance of viable hay-scented fern spores germinated from hardwood forest soils at various distances from a source. American Fern Journal 86(3): 69–79Google Scholar
  31. Penrod KA and McCormick LH (1997) Hay-scented fern spore production following clearcutting. Poster abstract. In: Proceedings of the 11th Central Hardwood Conference. Columbia, Missouri, USA, 23–26 March 1997, USDA Forest Service General Technical Report NC-188, p 401Google Scholar
  32. Potvin C (1993) ANOVA: Experiments in controlled environments. In: Scheiner SM and Gurevitch J (eds) Design and Analysis of Ecological Experiments, pp 46–68, Chapman & Hall, New YorkGoogle Scholar
  33. SAS Institute Inc (1996) SAS/STAT Release 6.12 Edition, SAS Institute Inc, Cary, North CarolinaGoogle Scholar
  34. Seiger LA and Merchant HC (1997) Mechanical control of Japanese knotweed (Fallopia japonica (Houtt.) Ronse Decraene): Effects of cutting regime on rhizomatous reserves. Natural Areas Journal 17: 341–345Google Scholar
  35. Shaw SP and McLaughlin CL (1951) The management of white-tailed deer in Massachusetts. Research Bulletin No. 13. Federal Aid in Wildlife Restoration — Project No. 7-R, 59 ppGoogle Scholar
  36. Stromayer KAK and Warren RJ (1997) Are overabundant deer herds in the eastern United States creating alternate stable states in forest plant communities? Wildlife Society Bulletin 25: 227–234Google Scholar
  37. Tilghman NG (1989) Impacts of white-tailed deer on forest regeneration in northwestern Pennsylvania. Journal of Wildlife Management 53(3): 524–532Google Scholar
  38. Waller DM and Alverson WS (1997) The white-tailed deer: a keystone herbivore. Wildlife Society Bulletin 25: 217–226Google Scholar

Copyright information

© Kluwer Academic Publishers 1999

Authors and Affiliations

  • Avril L. de la Cretaz
    • 1
  • Matthew J. Kelty
    • 1
  1. 1.Department of Forestry and Wildlife ManagementUniversity of MassachusettsAmherstUSA (e-mail

Personalised recommendations