Interference to Hardwood Regeneration in Northeastern North America: Assessing and Countering Ferns in Northern Hardwood Forests Heather M. Engelman and Ralph D. Nyland, Faculty of Forest and Natural Resources Management, SUNY College of Environmental Science and Forestry, Syracuse, NY 13210. ABSTRACT: The extremely dense shade cast by spreading ferns, particularly hayscented, New York, and bracken ferns, interferes with the survival and development of tree seedlings in northern hardwood forests. Excessive bracken frond litter and hayscented fern root mats can also prevent adequate germination and seedling development. In addition, the herbaceous cover may harbor detrimental small herbivores, while large ones often preferentially browse seedlings that grow through this layer. Increased understory light levels after an overstory disturbance, abundant soil moisture, fire, and herbivory promote ferns, whereas excessive and repeated cold or drought deter fern development and propagation. The most promising control methods repress ferns until seedlings cast adequate shade to inhibit further development of the fern layer. When ferns cover more than 30% of the understory, well-timed applications of either glyphosate or sulfometuron methyl have successfully controlled hayscented, New York, and bracken ferns. Two carefully timed mowings annually for at least 2 years have also provided long-lasting control on level, accessible sites. Deer populations must be reduced where browsing prevents development of desirable plants. North. J. Appl. For. 23(3):166 –175. Key Words: Pteridium aquilinum, Dennstaedtia punctilobula, Thelypteris noveboracensis, shade, herbicides, mowing. T he survival and development of tree seedlings and sprouts of northern hardwoods, and other desirable woody species around the world (Table 1), can be significantly inhibited by dense bracken (Pteridium aquilinum (L.) Kuhn), hayscented (Dennstaedtia punctilobula (Michx.) Moore) and New York (Thelypteris noveboracensis (L.) NOTE: 166 Heather M. Engelman can be reached at (315) 470-4877; Fax (315) 470-6956; engelman@syr.edu. Ralph D. Nyland can be reached at rnyland@mailbox.syr.edu. This paper is the third in a five-part series on Interference to Hardwood Regeneration in Northeastern North America. The first paper appeared in the March 2006 issue and the second paper appeared in the June 2006 issue. Drs. Ruth Yanai, Susan Stout, Robin Hoffman, Christopher Nowak, Tsutomu Nakatsugawa, and two anonymous reviewers provided comments on early drafts. This research was supported by the Cooperative Forest Research Unit (CRFU), University of Maine, and the NY Center for Forestry Research and Development of SUNY-ESF. CFRU published a companion annotated bibliography that presents detailed abstracts of publications reviewed here. For a copy contact Dr. Robert Wagner, Director, CFRU, 5755 Nutting Hall, University of Maine, Orono, ME 04469-5755; E-mail: bob㛭wagner@umnefa.maine.edu. Copyright © 2006 by the Society of American Foresters. NJAF 23(3) 2006 Nieuwland) ferns. In general, these plants become a problem because of their low palatability to insects and other animals (Dent 1971, Cooper-Driver et al. 1977, Jarrett 1982, Schreiner et al. 1984), their suspected allelopathic effects on competing species (Torkildson 1950, Gliessman and Muller 1972, Stewart 1975, Maquire and Forman 1983, Horsley 1977a, 1977b, 1989, 1993b, Drew 1990, Dolling 1996b), their shading of shorter tree seedlings (Horsley 1993b), their high propagule mobility (Page 1985), and their wide climatic and edaphic tolerance (Cox 1915, Page 1979). Further, spore banks and buried rhizomes survive fires and readily restore the population after surface burning (Dyer and Lindsay 1992). Widespread spore banks (Raynor et al. 1976, Schneller 1988, Milberg 1991, Dyer and Lindsay 1992, Penrod 1994, Penrod and McCormick 1996) spread ferns to new areas, but vegetative spread of their rhizomes increases the extent and density of these ferns where they already exist (Furuya 1978, Leck and Simpson 1987, Schneller 1988, Milberg 1991, Dyer and Lindsay 1992, Penrod and McCormick Table 1. Partial list of commercial tree species inhibited when overtopped by Pteridium aquilinum (P), Dennstaedtia punctilobula (D), or Thelypteris noveboracensis (T). Tree Fern Source Abies balsamea P Abies grandis Acer rubrum P D Betula allegheniensis Fraxinus america Liriodendron tulipifera D D D Picea engelmannii Picea glauca P P Picea rubens P Pinus contorta Pinus radiata Pinus strobus Pinus sylvestris Populus spp. Prunus serotina P P D P P D Prunus serotina Prunus serotina Pseudotsuga menquisii P T P Quercus rubra D Place 1952; Cody and Crompton 1975 Ferguson and Adams 1994 Maquire and Forman 1983; George and Bazzaz 1999a George and Bazzaz 1999a Bowersox and McCormick 1987 Bowersox and McCormick 1987, McCormick and Bowersox 1997 Comeau et al. 1993 Place 1952; Cody and Crompton 1975 Place 1952; Cody and Crompton 1975 Ferguson and Adams 1994 Karjalainen and Boomsma 1989 George and Bazzaz 1999a Jones 1947, Dolling 1996b Dolling 1996a Horsley and Marquis 1983, Horsley 1977b, 1989, 1993a, Drew 1990 Horsley 1977a Horsley 1977b McCulloch 1942, Stewart 1975, Stewart et al. 1979 Hanson and Dixon 1985, 1987, Bowersox and McCormick 1987, McCormick and Bowersox 1997, Lyon and Sharpe 1996, George and Bazzaz 1999a, 1999b 1996). On forested sites, the most aggressive fern species may form into a dense continuous layer that shades seedlings (Horsley 1993a, 1993b). Bracken in particular may also intercept falling seed, preventing germination. Its senescing fronds, either with or without the additional weight of snow, may also smother or crush other plants (Place 1952). These species of fern quickly proliferate and develop after understory light levels and other resources increase after an overstory disturbance (Horsley 1984b, Daniels 1986, Hollinger 1987, Chazon and Pearcy 1991, Hughes and Fahey 1991, Brach et al. 1993, Hurd 1995, Demchik and Sharpe 2001, Hill and Silander 2001). The resultant dense fern cover has little impact on nutrient availability to desired plants (Horsley 1988b, 1989, 1993a, Hurd 1995, Demchik and Sharpe 1999). Rather, the extremely dense shade from overlapping fronds prevents tree seedling establishment after germination (Bowersox and McCormick 1987, Horsley 1993a, 1993b, Hippensteel and Bowersox 1995, George and Bazzaz 1999a) and encourages fern persistence (Hall 1955). An intact hayscented fern root mat beneath these dense beds may also inhibit regeneration of desirable species (Hippensteel and Bowersox 1995, de la Cretaz and Kelty 1999). Allelopathic interactions have been reported (Gliessman and Muller 1972, Horsley 1977a, 1979, Maquire and Forman 1983, Drew 1990, Dolling 1996a), but the impact on germination and seedling establishment is minor compared to that of shading by a dense bed of these ferns (Horsley 1993b). In addition, browsing by deer may prevent height growth of established seedlings that grow through the fern layer (MacCammon 1938, Marquis and Grisez 1978, Kelty 1979, Tilghman 1989, Stroymayer and Warren 1997, Horsley et al. 2003), further encouraging fern persistence. Regeneration and Growth of Ferns Ferns are often distinguished by their growth habits, described as either clumping or spreading. Clumping ferns such as cinnamon fern (Osmunda cinnamomea L.) and spinulose woodfern (Drypopteris spinulosa (O.F. Muell.) Watt, Dryopteris carthusiana (Vill.) H.P. Fuchs, and Dryopteris austriaca (Jacq.) Woynar ex Schinz & Thellung var. spinulosa (O.F. Muell.) Fisch.) generally grow in selfcontained clusters. Spreading species such as bracken, hayscented, and New York ferns proliferate as dense beds of individual stalks that arise from rapidly spreading rhizomes and pose a much greater concern for forest managers. They occur throughout the eastern deciduous forest (Montgomery and Fairbrothers 1992), and have a wide tolerance of site conditions (Page 1979). In fact, their presence indicates very little about site quality. Bracken Fern Bracken fern is the only species in its genus (Cody and Crompton 1975). Two subspecies and many varieties have been reported, including Austral bracken (P. aquilinum var. esculentum (formerly P. esculentum (Forst) Nakai)) (Tryon 1941). Bracken fern is a large aggressive fern of open habitats with an almost worldwide distribution (Cody and Crompton 1975, Montgomery and Fairbrothers 1992). It often dominates the understory where it grows beneath forest cover (Tryon 1941, McCulloch 1942, Watt 1976, Biggin 1982, MacLeod 1982, Montgomery and Fairbrothers 1992, McDonald et al. 1999). Common bracken fern has been found in uncultivated places and rocky hillsides (Underwood 1881), in woodlands, and on dry slopes (Poel 1961). Northeastern bracken (P. aquilinum var. latiusculum (Desv.) Underw.) grows in dry open woods, open fields, and borders of wetlands from New Foundland to Minnesota; south to the uplands of North Carolina, Tennessee, Missouri, and Oklahoma, as well as in Mexico, Europe, and eastern Asia (Montgomery and Fairbrothers 1992). In persistent openings within the Allegheny hardwood forest, it grows in association with wild oat grass (Danthonia compressa Aust.), rough goldenrod (Solidago rugosa Mill.), and flattop aster (Aster umbellatus Mill.) (Horsley 1981). Despite its exceedingly wide range and competitive nature, young plants are unusual in the field. Spores are commonly consumed by insects (Conway 1953) and sporelings succumb to low temperature or the fungal pathogen Botrytis (Conway and Stephens 1957). Where dominant, bracken fern spreads in a regular pattern that appears to maximize coverage and minimize competition between ramets. Its long-creeping rhizomes spread NJAF 23(3) 2006 167 forward, and die away behind as it advances; when competition is encountered, the direction of growth changes accordingly (Watt 1976). Although resources are allocated to existing buds, and then to new rhizomes that facilitate spreading (Dolling 1999), bracken may both readily spread in area and increase in density to thousands of plants per acre. Dormant buds on the rhizomes provide a reservoir of potential fronds that replace those lost to frost, fire, or other agents (Biggin 1982, Daniels 1985, Ferguson and Boyd 1988). Although bracken fern spreads in clearcut areas where it has been present before cutting, it generally does not increase in frond density or colonize new ground in closed forests (Watt 1976, Dolling 1999). In general, bracken requires both abundant moisture (Biggin 1982, Thomson et al. 1985) and adequate drainage (Poel 1961). In dry weather, emerging fronds die and shrivel. In wet weather, they become chlorotic and die. If persistent, these conditions kill short shoots and cause death of fronds (Poel 1961, Watt 1976, Thomson et al. 1985). Excessive cold is also a naturally occurring deterrent to bracken fern. Unless adequately insulated by snow cover, litter, or other plants, shallow rhizome apices may be killed at low temperatures (Watt 1950, 1976). In fact, bracken is rarely found in consistently frosty areas where weather extremes may cause the loss of dormant buds and the depletion of stored carbohydrates (MacCammon 1938, Watt 1950, Williams and Foley 1976, Smith 1990). Allelopathy has been reported for bracken fern—the inhibition of germination, growth, or metabolism of one plant because of the release of organic chemicals from another (Stewart 1975). Bracken’s phytotoxicity increases during and after frond senescence (Gliessman and Muller 1972). Allelopathic chemicals from bracken fern have limited the growth and survival of aspen (Dolling 1996a) and orchard-grown black cherry seedlings (Horsley 1977a, 1979). The impact is lessened as litter becomes incorporated into the soil (Stewart 1975). The litter layer beneath bracken fern can act as a mechanical barrier against seeds reaching the soil and for seedlings emerging from below the litter mat (George and Bazzazz 1999a). It takes several years for holocellulose and lignin in bracken fern fronds to decompose, and much of the dry matter may remain for 11 years or more, particularly when moisture or nitrogen is limiting (Frankland 1976). Once seedlings have emerged from the frond mat, light levels rather than litter depth, or other factors associated with excessive litter, limit growth and development (George and Bazzazz 1999b). Hayscented Fern Hayscented fern is a large terrestrial fern of forest or forest border, and has wide ecological amplitude. It is common throughout the Appalachian Mountains where it may remain the dominant herb indefinitely (Horsley and Marquis 1983, Montgomery and Fairbrothers 1992, Stromayer and Warren 1997). It develops best on well-drained stony or sandy soils, and often forms thick beds from perennial rhizomes (Conard 1908, Cody et al. 1977, Horsley 168 NJAF 23(3) 2006 1984b). Hayscented fern is a common understory component of oak-transition hardwoods-hemlock (Hill and Silander 2001), sugar maple-beech (Berglund 1980), black cherry-maple (Marquis 1980), hemlock-hardwood (Maquire and Forman 1983), and transition hardwood-white-pine-hemlock types (George and Bazzaz 1999a). It generally occurs in open woods, clearings, and on roadside banks (Conard 1908, Flaccus 1959, Siccama et al. 1970, Hughes and Fahey 1991, Montgomery and Fairbrothers 1992, Hill and Silander 2001). Beneath closed forests, hayscented fern fronds and clones generally are sparsely distributed and do not noticeably interfere with the growth of a diversity of understory plants. The formation of dense colonies of hayscented fern is facilitated where deer browsing prevents development of other understory plants (MacCammon 1938, Marquis and Grisez 1978, Kelty 1979, Tilghman 1989, Stroymayer and Warren 1997, Horsley et al. 2003), especially after fires (McCulloch 1942, Cody et al. 1977, McGee et al. 1995) and where other overstory disturbances (e.g., thinning) increase light to the understory (Marquis et al. 1975, Cody et al. 1977, Horsley and Marquis 1983, Horsley et al. 2003). As with bracken fern, the direction of growth of the main axis facilitates forward spread until competition is encountered. Yet because rhizome buds are located on the rear-facing side of the rachis, rhizomes may reverse direction of growth if an obstacle prevents additional forward growth (S.B. Horsley, personal communication). Stems and roots from all portions of the rhizomes buried 5 to 15 cm beneath the soil surface may spread rapidly when conditions are suitable (Conard 1908, Horsley 1984b). Even small viable segments separated from the main rhizome mass may regrow to form dense beds that once more dominate the understory in as little as 5 years after control measures reduced the density and coverage of this fern (Horsley 1988a). In addition, viable sporebanks readily germinate on moist mineral seedbeds exposed by disturbance (Penrod and McCormick 1996). Seedlings having a wide range of shade tolerances (Acer rubrum, Betula, Prunus, and Tsuga canadensis) often occur at only low densities beneath hayscented fern. This suggests allelopathic interactions (Maquire and Forman 1983). Allelopathy was also suspected when growth of previously germinated black cherry did not immediately increase after hayscented fern was removed, just as the seedlings did when released from overtopping asters (Drew 1990). Heavy shading by the ferns was eventually hypothesized as the cause after greenhouse results could not be duplicated in a forested environment (Horsley 1977b). The effect of shading was finally demonstrated in field trials in which black cherry seedlings failed on plots having a closed fern cover, but grew well on plots where low fences held back the fern fronds to expose the seedlings to better light (Horsley 1993b). New York Fern New York fern occurs in mesic to damp woods and in full to partial shade, from Newfoundland to Michigan, south to Georgia, Mississippi, and Arkansas (Montgomery and Fairbrothers 1992). It is very hardy and grows relatively rapidly at optimal temperature, light, and nutrients (Hoshizaki and Moran 2001). New York fern is mediumsized, with creeping rhizomes. It forms colonies (Montgomery and Fairbrothers 1992) and is a common understory component of black cherry-maple (Marquis 1980), beechmaple (Smith 1980), and transition hardwood-white pinehemlock types (George and Bazzaz 1999a). Because they are often found together (Horsley 1981), New York and hayscented ferns had been thought to have similar growth habits. However, New York fern spreads more slowly than hayscented fern, posing less of a problem in forest regeneration efforts (Hill and Silander 2001). Further, its abundance tends to be highest, although not significantly so, under canopies of red (Acer rubrum L.) and sugar (A. saccharum Marsh.) maples at sites with abundant soil moisture, and lower under stands of other species or in the open. Spinulose Woodfern Spinulose woodfern is a medium-sized fern common to moist woods and swamps. Its erect rhizome system facilitates clumping of fronds, and results in much slower spreading (Marquis et al. 1992). Like many of the less invasive fern species, it only becomes restrictive when the density of its fronds prevents light from reaching new seedlings (S.B. Horsley, personal communication). Yet hayscented and New York ferns and spinulose woodfern are all considered potential threats to forest regeneration. They are counted together in tallies of interfering plants, with the importance of spinulose woodfern counts weighted by one-half that of hayscented or New York fern (Marquis et al. 1992). Guidelines for Assessing Fern Interference In northern hardwood stands with ⱕ60% canopy cover, sufficient light reaches the understory to promote the establishment and development of raspberries and blackberries, American beech, pin cherry, striped maple, hobblebush, sedges, ferns, and forbs (Hannah 1991, Nyland et al. 2005a, 2005b). When tall and dense, at least some of these plants may interfere with the development of more desirable tree species (Leak 1988, Nyland et al. 2006a, 2006b). In fact, some guidelines suggest that where undesirable species represent 50% or more of the advance regeneration, they should be controlled before shelterwood cutting (Hannah 1991, Sage et al. 2003). Those for Allegheny hardwoods prescribe site preparation when interfering plants occur on ⱖ30% of the stand area (Marquis et al. 1975). The control must remain effective for 3 to 4 years after an overstory cutting to ensure successful development of mixed-species regeneration (Horsley 1984b). Browsing pressure must also remain low (MacCammon 1938, Hough 1953, Marquis 1978, Horsley and Marquis 1983, Horsley et al. 1994, 2003). Management Strategies to Inhibit Fern Development Complete, long-term eradication of ferns is generally not necessary in forested communities. Rather, control is re- quired only until trees of desirable species grow taller than fern height. Also, for some species, little action may be required. For example, pin cherry (Prunus pennsylvanica L. f.) and blackberries (Rubus allegheniensis Porter) grow through dense fern cover (Horsley and Marquis 1983, Horsley 1984a), as will yellow (Betula alleghaniensis Britt.) and black (B. lenta L.) birch (Horsley and Marquis 1983, Horsley 1984a, de la Cretaz and Kelty 1999), eastern hemlock (Horsley et al. 2003), and white pine (de la Cretaz and Kelty 1999, Horsley et al. 2003). Substantial numbers of northern red oak (Quercus rubra L.) can grow through a fern canopy if not first consumed by insects and rodents (George and Bazzaz 1999b), but in reduced numbers (George and Bazzaz 1999b) and with slower height development (McCormick and Bowersox 1997). Other tree species of the northern hardwood forest will not survive beneath ferns nor develop through them. Managers may find fern cover excessive in a stand and elect to control it as part of their regeneration program strategy. Yet they must follow federal and state regulations regarding threatened and endangered species. In New York, for example, picking, plucking, severing, or otherwise killing “exploitably vulnerable” New York fern may be done only with the consent of a landowner (NY State Dept. of Environmental Conservation. 1989. New York State protected native plants. Title 6, New York Codes, Rules and Regulations, Section 193.3. State of NY, Albany). In other states, the intentional or negligent acts that result in the death of threatened or endangered plants may be generally prohibited. Ultimately, regulations such as these will determine the options available to a landowner. Manual and Mechanical Controls for Ferns In limited areas, pulling up shallow rhizomes or mowing the fronds close to the ground will control the vegetative spread of ferns (Brooks 1979, Biggin 1982, Hoshizaki and Moran 2001). For larger areas, other measures are required. Hayscented Fern Early studies showed that repeated mowing prevented mature fronds from developing sufficiently to maintain starch reserves necessary for production of new hayscented fronds (Cox 1915). More recently, de la Cretaz and Kelty (1999) compared scalping (root-raking), scarification, and mowing as control measures for hayscented fern in red and white pine (Pinus resinosa Soland and P. strobes L.) plantations. Clipping the fronds and leaving the litter in place provided nutrients to seedlings and also suppressed germination of seed-bank herbaceous species. Only a moderate number of seedlings become established, but there was less competition around them. Other evidence suggested that mowing the ferns for two growing seasons might substantially reduce fern abundance during the third growing season. Nevertheless, repeatedly clipping the fronds while also leaving existing tree seedlings unharmed could not easily be applied commercially. NJAF 23(3) 2006 169 Bracken Fern Bracken fern has been considered resistant to cutting, slashing, and bruising unless the treatment is continuous (Cody and Crompton 1975). Even so, a cutting during mid-June to early August, followed by a second mowing 5 to 6 weeks later, may provide adequate control (Preest 1975, Marrs et al. 1998). In fact, the levels of bracken in plots cut twice yearly for 6 years stayed low long after (i.e., 12 years) the treatment was stopped (Marrs et al. 1998). Historically, harvested bracken fronds were sold in Great Britain, providing a financial incentive for mechanical control. Fronds were mixed with grasses to feed livestock; used to cushion packed fruit and other items; and burned as a source of potash for glass (Tryon 1941) and soap making (Fletcher and Kirkwood 1979). They were valuable for stock-bedding, durable thatching, composting (Tryon 1941, Frankland 1976, Rymer 1976, Fletcher and Kirkwood 1979, MacLeod 1982), and as a fuel in brick making, brewing, and heating (Rymer 1976). As the demand for these uses fell, the impetus for harvesting waned, and bracken problems increased in the region. Nevertheless, bracken fronds could still be used as an opportunity energy crop if fronds were converted to pellets or anaerobically digested to methane, depending on the time of year of harvest (Callaghan et al. 1982, 1985, Lawson et al. 1984). Opening of such markets could provide incentives to renew bracken fern harvesting, particularly in open areas. Bracken fern is inherently unpalatable to most animals (Dent 1971, Cooper-Driver et al. 1977, Jarrett 1982, Schreiner et al. 1984), making grazing ineffective as a means of control. Young bracken fronds are popular as food in Japan and Korea, and many Japanese- and Korean-Americans harvest them for social or recreational reasons (Hirono 1986, Chavez and Gill 1999, Anderson et al. 2000). Connoisseurs prefer to harvest the fronds in their fiddlehead or crosier stage, and this occurs before the ferns shade shorter plants and before carcinogenic spores are released and inhaled. Even so, cutting of fronds as a food source seems unlikely to affect fern density over large areas. In addition, consumption of bracken fiddleheads is linked to high rates of digestive tract cancers (Hirono et al. 1972, 1987, Evans 1982a, 1982b, Soeder 1985, Hirono 1986), and that should discourage widespread consumption. Hayscented and New York ferns are not generally used for food. Chemical Control of Ferns Herbicide treatments limit regrowth of ferns by killing their rhizomes, but no selective herbicides are labeled for fern control in northern hardwood forests. Available broadspectrum herbicides can have the unwanted side effect of removing desirable tree seedlings as well as ferns (Horsley 1981, 1982), so formulations, rates, and timing must be carefully controlled. In addition, managers must ensure that pesticides in their prescriptions are registered for silvicultural site preparation in their locales. Table 2 summarizes current herbicide recommendations. Users should consult the source publications for details about formulations and application methods. The Herbicide Handbook (Vencill 2002) and the Environmental Protection Agency website (www.epa.gov/oppread201/international/ piclist.htm) provide extensive information about currently available compounds, including those approved for use in forestry operations and conditions of their registration. State and local regulations and herbicide labels should serve as a final reference with respect to any herbicide application described here. Hayscented and New York Ferns Correctly timed and applied Oust (E.I. du Pont de Nemours and Company, Wilmington, DE) herbicide (sulfometuron methyl) will control interfering hayscented and New York ferns, grasses, and sedges for 3 (Horsley 1991) to 8 years (S.B. Horsley, personal communication), including those that develop in the vehicle tracks after heavy machinery severs the rhizomes (Horsley 1988a, 1991). Late season (early August to early October) application of 2 oz product/ac (46.3 ml/ha) of Oust without a surfactant has provided nearly complete control of hayscented and New York ferns, while minimizing damage to valuable regeneration of Allegheny hardwoods and oaks in northwestern and central Pennsylvania (Horsley et al. 1992). Applications before frond maturance or after frond senescence resulted in significantly less control than applications from July through September (Horsley1988a). Glyphosate will also control ferns. The time of its application affects the rate needed for effective control of hayscented and New York fern. July or August application at 1 lb ai/ac (1.1 kg ai/ha) can economically reduce their populations, but an earlier application may require up to twice the dosage for a similar level of control (Horsley 1981). When bracken fern is also present, the treatment should be Table 2. Suggested herbicides for control of Dennstaedtia punctilobula (D), Thelypteris noveboracensis (T), and Pteridium aquilinum (P) in northern hardwoods. Chemical Target Sulfometuron methyl D, T Glyphosate D, T P 170 NJAF 23(3) 2006 Timing, rates, and comments ⫺1 Late season with 2 oz product ac (46.3 ml ha⫺1) using a low-pressure sprayer July 1 to September 1 with 1 lb ai ac⫺1 (1.1 ai kg ha⫺1) using a small compressed-air garden sprayer August to September 1 with 1 lb ai ac⫺1 (1.1 ai kg ha⫺1) using a small compressed-air garden sprayer Source Horsley et al. 1992 Horsley 1981 Horsley 1981, Karjalainen and Boomsma 1989 restricted to August (Horsley 1981). New York and hayscented ferns have not been adequately controlled with bromacil or hexaxinone (Horsley 1981). Bracken Fern Glyphosate at 1 lb ai/ac (1.1 kg ai/ha) is recommended for controlling bracken fern in northern hardwoods, with application during August. Higher rates may be necessary if applied earlier or later in the season (Horsley 1981). The effectiveness may be reduced when glyphosate becomes bonded to metal ions found in hard water (Sandberg et al. 1978, Stahlman and Phillips 1979, Shea and Tupy 1984), but can be restored with ethylenediaminetetraacetic acid (Shea and Tupy 1984) or by decreasing diluent volume (Sandberg et al. 1978, Stahlman and Phillips 1979). Asulam has also successfully controlled bracken fern in Europe and Australia (Martin 1976, Blatchford 1979, Heywood 1982). A foliar spray translocates to the rhizomes and prevents buds from developing (Veerasekaran et al. 1978, Heywood 1982, Kirkwood et al. 1982). The effectiveness has been increased through the use of wetters and/or emulsifiable oils (Heywood 1982), and by windrowing the ferns 1 year before treatment (Karjalainen and Boomsma 1989). Some additives increased translocation of asulam, at great cost: using low concentrations (50 ppm) of 2,4-D or CEPA (3-chloroethyl phosphoric acid) antagonized the activity of asulam (Kirkwood et al. 1982), whereas adding diesel fuel damaged desirable pine seedlings (Preest 1975). The recommended dosage is 1.18 gallons ac⫺1 (11 L ha⫺1) or 4.0 lb ac⫺1 (4.5 kg ha⫺1) in July or the beginning of August (Williams 1977, Heywood 1982); or before planting at 2.5 lb ai ac⫺1 (2.8 kg ai ha⫺1) using an incremental drift applicator ULV before frond tips unfurl or after planting 1.8 lb ai ac⫺1 (2 kg ai ha⫺1) using a mistblower (Blatchford 1979). Although it provides an effective level of control, its carbamate-based formula has limited asulam labeling in North America, and led to discontinued use in Canada (Pest Management Regulatory Agency 2002, 2003). It is labeled for the use in the United States to control western bracken fern on noncropland areas and in Christmas tree plantations (Asulox herbicide product label; Bayer CropScience LP, Research Triangle Park, NC.). Aminotriazole (3-amino-1,2,4-triazole) was also investigated for bracken control because of its ease of use in comparison to asulam. It gave effective control when applied with ammonium thiocyanate at 1:1 or 1:0.05 formulations (Cook et al. 1982). However, aminotriazole’s registration was cancelled in 1988 at the manufacturer’s request (S. Orme and S. Kegley. 2004. PAN pesticide database. Pesticide Action Network, San Francisco, CA; available at www.pesticideinfo.org). Other herbicides have proven ineffective for long-term control of bracken. These include dicamba that provided only 31% control of bracken fern (Radosevich 1979). Bromacil control was short-lived, and high rates of hexaxinone and picloram proved less effective than other compounds (Horsley 1981). Garlon 4 (Dow AgroSciences LLC Indianapolis, IN), Escort (E.I. duPont de Nemours and Company, Wilmington, DE), and Velpar L (E.I. duPont de Nemours and Company, Wilmington, DE) treatments actually promoted bracken fern to various degrees, rather than releasing desirable vegetation (McDonald et al. 1999). Reducing Browsing Also Essential In the first year or two after overstory removal in Allegheny hardwoods, ferns share dominance with Rubus, grasses, and sedges (Horsley et al. 2003). Once seedlings emerge from beneath this layer, deer and insects begin to feed on the trees (Horsley and Marquis 1983, George and Bazzaz 1996a, 1996b) and Rubus (Hough 1953, Tilghman 1989). At some sites, intensive browsing by white-tailed deer (Odocoileius virginianus) (MacCammon 1938, Marquis 1978, Tilghman 1989, Horsley et al. 2003), coupled with overstory thinning that increased light near the ground (Marquis et al. 1975, Cody et al. 1977, Horsley and Marquis 1983), led to the formation of dense hayscented fern and an eventual failure of desirable tree regeneration. After shelterwood seed cutting, only the least palatable species developed. These included striped maple (Acer pennsylvanicum L.), black cherry (Tilghman 1989), or American beech (Curtis and Rushmore 1958). Yet at low levels of deer browsing or no overstory cutting, only moderate fern cover develops (de la Cretaz and Kelty 1999). These findings suggest a management strategy that includes control of deer and ferns, and judicious cutting in the overstory (Marquis 1981, Walters and Nyland 1989, Horsley 1984a, Fredericksen et al. 1998, Horsley et al. 2003), particularly for thinning in even-aged stands (e.g., Marquis et al. 1992). Fencing could limit deer access to the regeneration area (Marquis and Grisez 1978). Alternatively, hunting of antlerless deer in the area surrounding a regeneration site can reduce deer pressure long enough for sufficient numbers of trees to regenerate and grow to heights beyond the reach of deer (Sage et al. 2003). In northwestern Pennsylvania, deer densities of ⱕ21 deer mi⫺2 (ⱕ8 deer km⫺2) will allow adequate regeneration of many tree species in large blocks of contiguous forest (Horsley et al. 2003). Kelty and Nyland (1981) observed repeated success with shelterwood seed cutting in New York’s Adirondacks after deer density had been reduced from 27 to 14 deer mi⫺2 (10 to 5 deer km⫺2). The maximum deer density depends on habitat quality. Management Implications Timber harvesting and other disturbances open the canopy, brightening the understory, and freeing moisture and nutrients to residual plants. Heavy machinery used during logging may disturb shallow rhizomes and promote vegetative proliferation of at least bracken, hayscented, and New York ferns across the brightened understory. As a countermeasure, some benefit accrues from limiting harvesting to dry periods (Groninger and McCormick 1992), or when the soil is frozen and covered with snow. Keeping overstory density sufficient to appropriately shade the ground also helps to limit the spread of understory ferns after thinning, NJAF 23(3) 2006 171 or after reproduction method cuttings that involve partial overstory removal. Because existing bracken, hayscented, and New York ferns readily spread if an upper canopy disturbance increases light levels near the ground, landowners should evaluate the status of ferns before any overstory treatment. Seventy percent of the area beneath a stand must be free of ferns for 3 to 4 years to insure adequate seedling development after a shelterwood seed cutting (Marquis et al. 1992), or any other reproduction method. Site preparation is essential before any of these treatments if ferns cover more of the area. Repeated mowing will control ferns, but is impractical for large areas, on uneven terrain, or in stands with uneven tree spacing. Also, where landowners prefer this option, the ferns must be cut twice per year for 2 years, with the mowing done after full frond development and again 5 to 6 weeks later. Because of these constraints, herbicide treatments (Table 2) have proven more practical for site preparation in forests. Either 1 lb ai lb ac⫺1 of Oust in July or August, or an early August to early October application of 2 oz/ac (46.3 ml/ha), is recommended for controlling hayscented and New York ferns. A single application of glyphosate in August will control bracken, hayscented, and New York ferns. Because the ferns will grow back after 3 to 4 years, landowners should couple the site preparation in susceptible stands with shelterwood seed cutting or other appropriate reproduction method to insure prompt establishment (Horsley 1982, Dolling 1996a). Research has not identified the exact proportion of basal area that can be safely removed during a seed cutting or thinning without promoting the spread of competing ferns and grasses in even-aged stands (Marquis et al. 1975, Horsley 1977b, Horsley et al. 2003). Management guides commonly recommend leaving a residual stand relative density of 60% for thinning (e.g., Marquis et al. 1992), or 50 to 60% stocking of upper canopy residual trees (21 to 25 m2/ha) for shelterwood seed cutting in northern hardwood stands having a pre-existing fern layer (Kelty 1986). The selection system may provide adequate shading to inhibit fern development, particularly if managers accept a shift to shade-tolerant tree species. The intermingled multilayered canopy (Kenefic and Nyland 1999, 2002) and the crowns of residual trees widen and shade the understory (Smith et al. 1997), especially if cutting is infrequent. Where browsing might prevent seedling development, deer density control must precede the site preparation and shelterwood cutting (Marquis 1981, Horsley 1984a, Walters and Nyland 1989, Frederickson et al. 1998, Horsley et al. 2003, Sage et al. 2003), and remain in effect until the trees have grown tall enough to exceed the reach of deer. 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