1

A literature review of management practices to support 

increased biodiversity in intensively managed loblolly pine 

plantations 

Final Technical Report to the National Commission on Science for 

Sustainable Forestry (NCSSF) 

July 31, 2005 

Kevin W. Zobrist 

Research Scientist 

Rural Technology Initiative 

University of Washington 

College of Forest Resources 

Box 352100 

Seattle, WA 98195-2100 

(206) 543-0827 

kzobr@u.washington.edu 

Thomas M. Hinckley 

Professor 

University of Washington 

College of Forest Resources 

hinckley@u.washington.edu 

Michael G. Andreu 

Assistant Professor 

University of Florida - IFAS 

School of Forest Resources and Conservation 

mandreu@ifas.ufl.edu 

The National Commission on Science for Sustainable Forestry (NCSSF) sponsored the research 

described in this report. The National Council on Science and the Environment (NCSE) conducts the 

NCSSF program with support from the Doris Duke Charitable Foundation, the David and Lucile Packard 

Foundation, the Surdna Foundation, and the National Forest Foundation. 
 2

Contents 

I. Introduction ..........................................................................................................3 

II. Management practices to support increased biodiversity ..............................4 

III. Other considerations.........................................................................................8 

IV. Summary ............................................................................................................9 

Metric equivalents..................................................................................................10 

Literature cited.......................................................................................................10 
 3

I. Introduction 

When European settlers first arrived in North America, it is estimated that the South had 200 

million acres in pine, mixed oak, and other forest systems. Pine savannahs and open woodlands 

containing longleaf (Pinus palustris), loblolly (P. taeda), shortleaf (P. echinata), slash (P. 

elliottii) and pond (P. serotina) pine dominated. Of these species, it is estimated that longleaf 

pine ecosystems may have covered over 60 million acres1 (Bragg 2002, Wahlenberg 1946). 

Longleaf pine stands were often characterized by a single species overstory, a sparse midstory/shrub layer, and a well-developed and species-rich ground layer. Frequent, low intensity 

fires, natural or anthropogenic in origin, were the primary disturbance regime (Noss 1988, Van 

Lear et al. 2004). These stands were also known for the diversity of wildlife, particularly game 

species, they harbored. Trees in these stands were often very large (Table 1) and ranged as old as 

600 plus years. Today, there are only about 3 million acres of longleaf pine forest, of which only 

12,000 are regarded as old-growth (Bragg 2002). These old growth stands are now recognized 

for the habitat that they provide for dozens of threatened species (Bragg 2002, Noss 1988). 

Table 1. Heights and diameters of old growth pines in the south and southeast (Bragg 2002). 

Species Height (feet) Diameter at breast height (inches) 

Loblolly pine 145 -180 39 - 65 

Longleaf pine 85 - 130 27 - 45 

Shortleaf pine 85 - 140 35 - 50 

Nearly 90% of the forestland in the South today is in private ownership (Wicker 2002), and 

much of it is comprised of dense plantations of fast-growing loblolly pine. The management 

intensity of these plantations has been increasing in recent decades (Siry 2002). At the same 

time, private landowners are facing an increasing demand to provide for broad, non-timber 

values such as biodiversity on these lands, which can lead to conflict over forest management 

practices. 

Forest plantations have long been negatively characterized as biological “deserts” in which 

concern for wildlife is limited to key game species (Margolin 1970). While it is true that today’s 

dense loblolly pine plantations are different from the natural, open pine stands that were 

historically prevalent throughout the South, these intensively managed forests can still contribute 

to biodiversity on the landscape (Wigley et al. 2000). This contribution may fall short relative to 

natural forests, but it may compare favorably to other competing land uses such as agriculture or 

urbanization (Moore and Allen 1999). 

At any rate, because of their prevalence in the region, intensively managed plantations have a 

significant potential impact on the level of biodiversity. Stand-level management changes can be 

made that can readily support increased biodiversity in these plantations. Even minor changes 

 

1

 Estimates vary from a low of 30 to a high of 92 million acres. If one uses the low estimate, 

then today’s acrage of longleaf pine is only 10% of the original and of that only a very small 

fraction is in “old-growth.” Therefore, there is value in discussing mechanisms to increase oldgrowth structures in these remaining forests in the South. 
 4

can significantly improve biodiversity values (Hartley 2002, Johnson et al. 1975). Some of these 

changes may be complimentary with timber production and economic goals; others may involve 

some costs and trade-offs (Allen et al. 1996, Buongiorno et al. 2004, Hunter 1990). In this paper 

we review the literature to identify a spectrum of practices that support increased biodiversity in 

intensively managed loblolly pine plantations. 

II. Management practices to support increased biodiversity 

Biodiversity has several definitions, the simplest being a variety of life. Many definitions further 

specify that the definition includes all types of organisms as well as genotypes and even 

ecological processes and their inter-relationships (Hunter 1990, Oliver 1992, Reid and Miller 

1989).1 There is no single forest type or structure that maximizes biodiversity. Different species 

have different habitat requirements such that structures that support some species may not 

support others (Dickson and Wigley 2001). Even individual species require habitat diversity 

(Johnson et al. 1975). Thus the overall key to supporting a variety and abundance of species is to 

provide a diversity of structure and vegetation (Allen et al. 1996, Harris et al. 1979, Marion et al. 

1986, Sharitz et al. 1992). This includes both within stand diversity and between stand diversity 

(Marion and Harris 1982, Thill 1990). 

An important way to increase within-stand structural diversity is to maintain a lower overstory 

density. A more open canopy allows a diverse understory to develop, which provides forage and 

habitat for wildlife. Even plantations established with intensive site preparation are often very 

diverse in the early years as long as the canopy is open, but as the canopy closes this diversity 

rapidly decreases (Baker and Hunter 2002). Once the canopy closes, the stand moves into the 

stem exclusion stage that shades out the understory vegetation and subsequently lacks wildlife 

(Oliver and Larson 1990). Minimizing this stage can allow a stand to support more biodiversity 

over a given rotation. Maintaining an open canopy with a productive understory also makes 

plantations more similar to the diverse, natural pine communities that existed historically in this 

region (Bragg 2002, Noss 1988, Van Lear et al. 2004). 

One way to maintain lower stand density for biodiversity is by planting at a lower density. A 

wider spacing, such as 12 feet, delays canopy closure, extending the more diverse earlysuccessional stages (Dickson 1982, Johnson et al. 1975, Melchiors 1991). In addition to delaying 

canopy closure, a wider spacing between rows can also allow disking or mowing to help 

maintain a productive understory (Allen et al. 1996). A wider planting spacing may be 

undesirable, though, because of the resulting decreased wood quality. In this case a closer 

spacing followed by thinning may be a more desirable approach (Van Lear et al. 2004). 

 

1

 “Biodiversity is defined by the UN as the variability among living organisms from all sources 

including terrestrial, marine, and other aquatic systems and the ecological complexes of which 

they are a part; this includes diversity within species, between species and of ecosystems. The 

Global Biodiversity Strategy (WRL, IUCN, and UNEP 1992) espouses a shorter definition where 

biodiversity is the totality of genes, species and ecosystems in a region.” (Patel-Weynand 2002) 
 5

Perhaps the most important way to establish and maintain an open, diverse structure is by 

thinning. Thinning has been found to benefit numerous individual wildlife species such as deer 

(Blair 1960, Halls 1973, Hurst and Warren 1980), quail (Dougherty 2004), small mammals 

(Mengak and Guynn 2003), turkeys (Mississippi State University Extension Service 2004), 

nuthatches (Wilson and Watts 1999), and other birds (Turner et al. 2002). Thinning early and 

often is widely recognized as an important component of an overall strategy to increase 

biodiversity (Hunter 1990, Marion et al. 1986). This minimizes the time in the stem exclusion 

stage and can maintain and further develop an open, diverse structure throughout the rotation. In 

addition to stimulating the herbaceous understory by allowing light to reach the forest floor 

(Harrington and Edwards 1999, Schultz 1997), thinning also facilitates additional understory 

management such as disking or burning (Johnson et al. 1975), and it increases the understory 

response to such treatments (Melchiors 1991, Tucker et al. 1998). 

Van Lear et al. (2004) suggest that a commercial thinning be done by the time a plantation 

reaches age 15. Hurst and Warren (1980) suggest that it be done as early as age 12 if no precommercial thinning was done. The recommended frequency of thinning to maintain an open 

stand structure is usually around five years (Blair and Enghardt 1976, Conroy et al. 1982, Halls 

1973, Hunter 1990, Schultz 1997). Maintaining an open stand structure requires heavier thinning 

than for timber production, with a target of 50-70 ft2

/acre of residual basal area (Blair and 

Enghardt 1976, Halls 1973, Van Lear et al. 2004). A residual basal area of 80 ft2/acre is usually 

considered a minimum for timber production and economic return (Siry 2002, Siry et al. 2001). 

A potential problem with thinning to open up the pine overstory is that it can allow understory 

hardwoods to develop into a dense midstory. Hardwoods produce heavy shade that inhibits 

understory vegetation (Blair and Enghardt 1976, Dickson and Wigley 2001, Schultz 1997, 

Tappe et al. 1993). Thinning can also increase vines and shrubs, which further shade out the 

herbaceous layer (Harrington and Edwards 1999). Thus, without controlling hardwoods and 

other woody vegetation, thinning can ultimately result in a less productive and less diverse 

understory (Blair and Feduccia 1977, Hunter 1990). A hardwood midstory can add vertical 

stratification and benefit some midstory-associated birds (Dickson 1982, Melchiors 1991, 

Turner et al. 2002). However, a hardwood midstory is generally undesirable for most wildlife, 

including deer, small mammals, and other birds (Dickson 1982, Lohr et al. 2002, Melchiors 

1991, Wilson and Watts 1999). 

While a dense midstory is undesirable, some hardwoods are necessary for supporting 

biodiversity. Mature hardwoods such as oaks provide hard mast that is important for many 

wildlife species (Dickson 1982, Dickson and Wigley 2001, Johnson 1987, Johnson et al. 1975). 

Maintaining a desirable component of hardwoods can improve wildlife habitat (Tappe et al. 

1993). When controlling hardwoods, individual fruit or mast producing trees can be selectively 

retained (Blair 1967, Blair and Feduccia 1977). Maintaining whole areas of hardwoods is also 

important. An interspersion of hardwood and pine forest types provides good wildlife habitat 

(Shultz 1997). Hardwoods should especially be maintained in sensitive areas such as 

bottomlands and drainages (Halls 1973, Johnson et al. 1975). Hardwood maintenance should 

generally stay focused on hardwood sites (Johnson 1987). 
 6

In historic, natural pine stands, frequent low-intensity fires helped to control hardwoods and 

maintain an open stand structure with a productive and diverse understory (Noss 1988, Van Lear 

et al. 2004). Frequent low-intensity fires tend to favor growth of herbaceous vegetation by 

suppressing hardwoods and other woody vegetation (Reed et al. 1994). Prescribed burning can 

be used in conjunction with thinning in intensively managed plantations to achieve desired 

conditions that support increased biodiversity. Many of the plants and animals associated with 

southern pine communities are adapted to or even dependent on fire, and wildlife mortality from 

fire is generally very low (Landers 1987, Means and Campbell 1981, Moorman 2002). Regular 

burning improves habitat for many species, including deer (Dickson 1982, Halls 1973, Hurst et 

al. 1980, Marion et al. 1986), quail (Dougherty 2004), turkey (Marion et al. 1986, Mississippi 

State University Extension Service 2004), amphibians and reptiles (Means and Campbell 1981), 

and Bachman’s sparrow (Tucker et al. 1998). To help provide for a broad suite of species in the 

short and long term, areas should not be burned evenly, but rather patches of unburned areas 

should be left to provide for nesting and cover (Landers 1987, Moorman 2002). 

Prescribed burning is recommended when the dominant pine trees are 15 feet tall (Halls 1973, 

Moorman 2002). Recommended burning intervals range from 3-6 years (Blair and Enghardt 

1976, Blair and Feduccia 1977, Halls 1973, Mississippi State Extension Service 2004, Schultz 

1997). Marion et al. (1986) suggest 3-5 years to allow enough time for browse and cover to 

develop, as well as enough fuel to carry the next fire. Historically, longleaf pine communities in 

Florida burned naturally every 2-5 years (Noss 1988). Prescribed burning should not be 

overdone, or the cumulative impacts could become negative in the long term (Melchiors 1991). 

For example, if burning is done too frequently, such as annually, it can eliminate hardwoods 

altogether, (Dickson 1982, Grano 1970, Schultz 1997). Complete loss of the hardwood 

component would have a negative impact on biodiversity. 

Many authors recommend burning in winter (Allen et al. 1986, Blair and Feduccia 1977, Halls 

1973, Mississippi State Extension Service 2004, Schultz 1997). However, Robbins and Myers 

(1992) note that varying both the season and the frequency of burning avoids favoring only one 

suite of species. Adding this element of variability can increase overall stand diversity. 

Coordinating burning with thinning is also important. Thinning increases the effectiveness of 

prescribed burning for wildlife (Hurst and Warren 1982, Melchiors 1991, Tucker et al. 1998). 

Burning before thinning can make thinning easier (Hurst et al. 1980), and it avoids the problem 

of the fire burning too intensely in the slash from the thinned trees (Van Lear et al. 2004). 

An alternative to prescribed burning for the control of nonpine woody vegetation is to use 

herbicides (Dickson and Wigley 2001, Harrington and Edwards 1999). Herbicides can be less 

costly than burning and may be especially desirable where burning opportunities are limited 

(Wigley et al. 2002). Normal applications of herbicides are generally not directly toxic to 

wildlife (McComb and Hurst 1987). Herbicides may have a longer residual effect on understory 

diversity than prescribed burning or mechanical vegetation control (Hunter 1990). Nonetheless, 

vegetation seems to recover quickly within 1-3 years (Keyser et al. 2003, Reed et al. 2004). A 

longer term study found no significant impact on floristic diversity 11 years after herbicide 

treatment (Miller et al. 1999). 
 7

The intensity and methods of site preparation for controlling vegetation at the beginning of the 

rotation should also be considered when managing pine plantations for increased biodiversity. 

More intensive site preparation favors grass and forbs, while less intensive site preparation 

favors vines and woody vegetation (Johnson 1975, Locascio et al. 1990). More intensive site 

preparation also reduces the availability of fruit for wildlife (Hunter 1990, Stransky and Roese 

1984). Thus, while intensive site preparation can benefit some game species like deer, less 

intensive site preparation is generally better for a diversity of wildlife (Marion and Harris 1982, 

Marion et al. 1986). Locascio et al. (1990) found that moderate intensity site preparation 

produced the greatest understory biomass, and moderate intensity treatments may be the most 

cost effective, especially for non-industrial landowners. In terms of site preparation methods, 

Locascio et al. (1991) observed that mechanical site preparation (shear, chop, disk, etc.) did not 

seem to diminish understory plant diversity. Mechanical methods may provide for greater 

understory diversity and food production compared to herbicides (Fredericksen et al. 1991, 

Keyser et al. 2003). Burning may also be a desirable option for stimulating stored seeds (Hunter 

1990). 

Other management activities like fertilization and pruning can also impact biodiversity. Use of 

fertilization in pine plantations has increased in recent decades, though it is mostly done on 

industry lands (Siry 2002). The impacts of fertilization on biodiversity are somewhat mixed. 

Fertilization can improve understory food production for wildlife, especially in stands that have 

been thinned (Hunter 1990, Hurst and Warren 1982, Melchiors 1991). Fertilization can also 

accelerate canopy closure, though, which can offset wildlife benefits (Dickson and Wigley 

2001). Thus fertilization treatments should be done in conjunction with thinning to maximize 

wildlife benefits. Pruning can benefit biodiversity by increasing understory vegetation (Baker 

and Hunter 2002, Hurst and Warren 1982) as well as creating more horizontal openings. 

Another way to support increased biodiversity in pine plantations is by retaining key structural 

features such as snags, coarse woody debris, and mature live trees. These elements add 

additional structural complexity that benefits a wide range of wildlife (Allen et al. 1996, Baker 

and Hunter 2002, Dickson and Wigley 2001, Lohr et al. 2002, Marion et al. 1986, Sharitz et al. 

1992). Maintaining riparian buffers, or streamside management zones, can provide for some of 

these elements (Dickson and Wigley 2001, Thill 1990). Riparian buffers further contribute to 

biodiversity by providing for aquatic species and water quality (Baker and Hunter 2002) and by 

providing habitat connectivity (Dickson and Wigley 2001, Johnson 1987). 

All of the management practices described above will be most effective if done in conjunction 

with longer rotations. Pulpwood rotations can be as short as 20 years (Biblis et al. 1998, 

Melchiors 1991). Short rotation management limits pine plantations to early successional 

structures and does not provide for species needing older seral stages (Johnson et al. 1975). 

Because of the dominance of short rotations, older seral stages are becoming rare in the region 

(Allen et al. 1996). Managing for longer rotations can increase diversity (Sharitz et al. 1992). 

Rotations of 40-100 years can provide for long-term wildlife forage as well as key habitat 

elements such as hardwood mast, snags, and cavities (Melchiors 1991). 

Longer rotations can impact economic returns. Because future revenues are discounted, longer 

rotations must produce significantly more revenue to be economically competitive with shorter 
 8

rotations. Dean and Chang (2002) found that economic performance decreased with increasing 

rotation length. In contrast, Biblis et al. (1998) noted that 50-year sawtimber rotations performed 

better economically than 20-year pulpwood rotations if the target rate of return was 7% or less. 

Ultimately it depends on the relative prices of pulpwood and sawtimber and the rate of return 

that is acceptable to the landowner. 

III. Other considerations 

In looking at management practices to increase biodiversity in intensively managed loblolly pine 

plantations, some additional considerations should be made. The management practices 

described in this paper are geared towards increasing stand-level biodiversity. Ultimately, 

though, a landscape approach is needed. A variety of different stand structures and age classes 

should be present on the landscape to support the full range of biodiversity (Marion et al. 1986, 

Moore and Allen 1999, Oliver 1992). The size, shape, and spatial arrangement of these 

structures are also important (Johnson 1987). For landowners with large areas of contiguous 

holdings, a landscape management approach to providing for biodiversity may be feasible. 

When the landscape is broken up among different ownerships, landscape management requires 

coordination between different landowners with different needs and goals. The issues involved 

with such coordination are beyond the scope of this review. In any case, maintaining 

biodiversity at the landscape level depends on a collection of stand-level decisions. If individual 

landowners employ practices to increase stand level biodiversity, their practices are likely to 

support significantly increased biodiversity across the landscape. 

Another important consideration when managing for biodiversity is land use history. Hedman et 

al. (2000) found that understory vegetation is driven more by previous land use than forest 

management practices within the past 35 years. Plantations established on old field sites do not 

have biological legacies such as seeds and rootstocks that are present in plantations established 

on cutover lands (Baker and Hunter 2002). Because of this, old field sites have low understory 

diversity regardless of management practices (Hedman et al. 2000, Marion and Harris 1982, 

Marion et al. 1986). On the other hand, old field sites have greater pine growth and yield and can 

produce more wood per area of land (Yin and Sedjo 2001). Thus, intensive timber management 

that maximizes wood production and economic return should be focused on old field sites where 

biodiversity is likely to be poor regardless of management practices. Likewise, practices to 

improve biodiversity should be targeted to cutover lands. 

Finally, there should be economic considerations when examining ways to increase biodiversity. 

Intensively managed plantations are business enterprises for which landowners will expect some 

level of economic return. There are various costs associated with managing for increased 

biodiversity which create trade-offs between biodiversity and economic returns (Allen et al. 

1996, Hunter 1990). If management practices are too costly, they are unlikely to be implemented 

on private lands. Management strategies that balance both biodiversity and economic objectives 

should be identified (Buongiorno et al. 2004). 

One thing that may help offset the costs of managing for increased biodiversity is the potential 

for increased hunting lease revenue (McKee 1987, Melchiors 1991). Hunting leases can provide 
 9

significant revenue, especially if there is quality wildlife habitat (Baker and Hunter 2002, 

Johnson 1995, Jones et al. 2001). Ownership size may limit these opportunities, though. 

IV. Summary 

There is concern about the maintenance of biodiversity in the intensively managed loblolly pine 

plantations that are increasingly prevalent in the southeastern United States. There are a number 

of stand-level management practices that can support increased biodiversity in these plantations. 

The overall key to providing for biodiversity is to provide structural diversity. An open stand 

structure with a diverse, productive grass-herb understory is more similar to the natural, firemaintained pine communities that were historically present and can support a broad suite of 

plants and wildlife. 

Maintaining an open canopy with a diverse understory can be achieved with heavy thinnings 

early and often in the rotation. This may allow a dense hardwood midstory to develop, though, 

which would shade out the understory and negate the benefits of thinning. Consequently, 

hardwood control will be necessary either by prescribed burning or with mid-rotation herbicide 

applications. Hardwoods should not be eliminated entirely. A mast producing component should 

be maintained to provide wildlife food and structural diversity. 

Light to moderate site preparation is best for biodiversity, and mechanical methods may perform 

better in this respect than herbicides. Fertilization can benefit wildlife by increasing understory 

growth, but it should be done in conjunction with thinning to maximize benefits. Key structural 

features such as snags, coarse woody debris, and mature trees should be maintained, along with 

riparian buffers to protect aquatic areas and provide for habitat connectivity. Long rotations are 

necessary to provide a broader range of age classes, though the economic impacts may be a 

consideration. 

Biodiversity is ultimately achieved at the landscape level, but stand-level changes can go a long 

way towards making improvements and can be implemented regardless of ownership pattern. 

Land use history is an important consideration, as old field sites are unlikely to support a diverse 

stand structure regardless of management practices. Economics should also be considered, as 

management practices to increase biodiversity need to be economically viable if they are to be 

successful on private lands. Opportunities for hunting lease revenue may offset some of the costs 

of managing for biodiversity. 
 10

Metric equivalents 

When you know: Multiply by: To find: 

Acres 0.4047 Hectares 

Feet 0.3048 Meters 

Inches 2.54 Centimeters 

Square feet per acre (ft2/acre) 0.229 Square meters per hectare 

Literature cited 

Allen, A.W., Bernal, Y.K., and R.J. Moulton. 1996. Pine plantations and wildlife in the 

Southeastern United States: an assessment of impacts and opportunities. U. S. Dept. of the 

Interior, National Biological Service, Information and Technology Report 3. 32 p. Available 

online at http://www.nwrc.usgs.gov/wdb/pub/others/1996_03.pdf; last accessed June 2005. 

Baker, J.C. and W.C. Hunter. 2002. Effects of forest management on terrestrial ecosystems. 

Pages 91-112 in D.N. Wear and J.G. Greis, eds. Southern Forest Resource Assessment. General 

Technical Report SRS-53. USDA Forest Service, Southern Research Station, Ashville, NC. 

Biblis, E.J., H. Carino, and L. Teeter. 1998. Comparative economic analysis of two management 

options for loblolly pine timber plantations. Forest Products Journal 48(4):29-33. 

Blair, R.M. 1967. Deer forage in a loblolly pine plantation. Journal of Wildlife Management

31(3):432-437. 

Blair, R.M. 1960. Deer forage increased by thinnings in a Louisiana loblolly pine plantation. 

Journal of Wildlife Management 24(4):401-405. 

Blair, R.M. and H.G. Enghardt. 1976. Deer forage and overstory dynamics in a loblolly pine 

plantation. Journal of Range Management 29(2):104-108. 

Blair, R.M. and D.P. Feduccia. 1977. Midstory hardwoods inhibit deer forage in loblolly pine 

plantations. Journal of Wildlife Management 41:677-684. 

Bragg, D.C. 2002. Reference conditions for old-growth pine forests in upper west Gulf Coastal 

Plain. Journal of the Torrey Botanical Society 29(4):261-288. 

Buongiorno, J., B. Schulte, and K.E. Skog. 2004. Quantifying trade-offs between economic an 

ecological objectives in uneven-aged mixed-species forests in the southern United States. 

General Technical Report FPL-CTR-145. USDA Forest Service, Forest Products Laboratory, 

Madison, WI. 5 p. 

Conroy, M.J., R.G. Oderwald, and T.L. Sharik. 1982. Forage production and nutrient 

concentrations in thinned loblolly pine plantations. Journal of Wildlife Management 46(3):719-

727. 
 11

Dean, T.J. and S.J. Chang. 2002. Using simple marginal analysis and density management 

diagrams for prescribing density management. Southern Journal of Applied Forestry 26(2):85-

92. 

Dickson, J.G. 1982 Impact of forestry practices on wildlife in southern pine forests. Pages 224-

230 in Proceedings of the 1981 Convention of the Society of American Foresters. 

Dickson, J.G. and T.B. Wigley. 2001. Managing forests for wildlife. Pages 83-94 in Dickson, 

J.G., ed. Wildlife of Southern Forests: Habitat and Management. Hancock House Publishers, 

Blaine, WA. 

Dougherty, D. 2004. Think habitat: Creating “usable space” for quail. Forest Landowner

63(3):5-8. 

Fredericksen, T.S., H.L. Allen, and T.R. Wentworth. 1991. Competing vegetation and pine 

growth response to silvicultural treatments in a six-year-old Piedmont loblolly pine plantation. 

Southern Journal of Applied Forestry 15(3):138-144. 

Grano, C.X. 1970. Eradicating understory hardwoods by repeated prescribed burning. Resource 

Paper SO-56. USDA Forest Service, Southern Forest Experiment Station, New Orleans, LA. 

Halls, L.K. 1973. Managing deer habitat in Loblolly-Shortleaf pine forests. Journal of Forestry

71(12):752-757. 

Harrington, T.B. and M.B. Edwards. 1999. Understory vegetation, resource availability, and 

litterfall responses to pine thinning and woody vegetation control in longleaf pine plantations. 

Canadian Journal of Forest Research 29:1055-1064. 

Harris, L.D., D.H. Hirth, and W.R. Marion. 1979. The development of silvicultural systems for 

wildlife. Pages 65-80 in C.L. Shilling and J.R. Toliver, eds. Recreation in the South’s third 

forest. Twenty-eighth Annual Forestry symposium, Louisiana State University, Baton Rouge, 

LA. 

Hartley, M.J. 2002. Rationale and methods for conserving biodiversity in plantation forests. 

Forest Ecology and Management 155:81-95. 

Hedman, C.W., S.L. Grace, and S.E. King. 2000. Vegetation composition and structure of 

southern coastal plain pine forests: an ecological comparison. Forest Ecology and Management

134:233-247. 

Hunter, M.L., Jr. 1990. Wildlife, forests, and forestry: Principles of managing forests for 

biological diversity. Prentice Hall, Englewood Cliffs, NJ. 370 p. 
 12

Hurst, G.A., J.J. Campo, and M.B. Brooks. 1980. Deer forage in a burned and burned-thinned 

pine plantation. Proceedings of the Annual Conference of the Southeastern Association of Fish 

and Wildlife Agencies 34:476-481. 

Hurst, G.A. and R.C. Warren. 1980. Intensive pine plantation management and white-tailed deer 

habitat. Pages 90-102 in R.H. Chabreck and R.H. Mills, eds. Integrating timber and wildlife 

management in southern forests. Twenty-ninth Annual Forestry symposium, Louisiana State 

University, Baton Rouge, LA. 

Hurst, G.A. and R.C. Warren. 1982. Impacts of silvicultural practices in loblolly pine plantations 

on white-tailed deer habitat. Pages 484-487 in E.P. Jones, ed. Proceedings of the second biennial 

Southern Silvicultural Research Conference. General Technical Report SE-24. USDA Forest 

Service, Southeastern Forest Experiment Station, Asheville, NC. 

Johnson, R. 1995. Supplemental sources of income for southern timberland owners. Journal of 

Forestry 93(3):22-24. 

Johnson, A.S. 1987. Pine plantations as wildlife habitat: a perspective. Pages 12-18 in J.G. 

Dickson and O.E. Maughan, eds. Managing southern forests for wildlife and fish, a proceedings. 

General Technical Report SO-65. USDA Forest Service, Southern Forest Experiment Station, 

New Orleans, LA. 

Johnson, A.S., J.L. Landers, and T.D. Atkeson. 1975. Wildlife in young pine plantations. Pages 

147-159 in Proceedings of the symposium on management of young pines. USDA Forest Service, 

Southeast Area, State and Private Forestry, Atlanta, GA. 

Jones, W.D., U.A. Munn, S.C. Grado, and J.C. Jones. 2001. Fee hunting—An income source for 

Mississippi’s non-industrial, private landowners. Resource Bulletin #FO 164, Forestry and 

Wildlife Resource Center, Mississippi State University, Mississippi State, MS. 15 p. 

Keyser, P.D., V.L. Ford, and D.C. Guynn, Jr. 2003. Effects of herbaceous competition control on 

wildlife habitat quality in Piedmont pine plantations. Southern Journal of Applied Forestry

27(1):55-60. 

Landers, J.L. 1987. Prescribed burning for managing wildlife in Southeastern pine forests. Pages 

19-27 in J.G. Dickson and O.E. Maughan, eds. Managing southern forests for wildlife and fish, a 

proceedings. General Technical Report SO-65. USDA Forest Service, Southern Forest 

Experiment Station, New Orleans, LA. 

Locascio, C.G., B.G. Lockaby, J.P. Caulfield, M.G. Edwards, and M.K. Causey. 1990. Influence 

of mechanical site preparation on deer forage in the Georgia Piedmont. Southern Journal of 

Applied Forestry 14(2):77-80. 

Locasio, C.G., B.G. Lockaby, J.P. Caufield, M.B. Edwards, and M.K. Causey. 1991. Mechanical 

site preparation effects on understory plant diversity in the Piedmont of the southern USA. New 

Forests 4:261-269. 
 13

Lohr, S.M., S.A. Gauthreaux, and J.C. Kilgo. 2002. Importance of coarse woody debris to avian 

communities in loblolly pine forests. Conservations Biology 16(3):767-777. 

Margolin, M. 1970. Desert under the trees. National Parks and Conservation Magazine 44(279): 

8-13. 

Marion, W.R. and L.D. Harris. 1982. Relationships between increasing forest productivity and 

fauna in the flatwoods of the southeastern coastal plain. Pages 215-222 in Proceedings of the 

1981 Convention of the Society of American Foresters. 

Marion, W.R., M. Werner, and G.W. Tanner. 1986. Management of pine forests for selected 

wildlife in Florida. Circular 706. Florida Cooperative Extension Service, Institute of Food and 

Agricultural Sciences, University of Florida. Available online at 

http://wfrec.ifas.ufl.edu/range/pdf_docs/tanner/cir-706.pdf; last accessed June 2005. 

McComb, W.C. and G.A. Hurst. 1987. Herbicides and wildlife in southern forests. Pages 28-37 

in J.G. Dickson and O.E. Maughan, eds. Managing southern forests for wildlife and fish, a 

proceedings. General Technical Report SO-65. USDA Forest Service, Southern Forest 

Experiment Station, New Orleans, LA. 

McKee, C.W. 1987. Economics of accommodating wildlife. Pages 133-137 in Proceedings of 

the 1986 Society of American Foresters National Convention. 

Means, D.B. and W.H. Campbell. 1981. Effects of prescribed burning on amphibians and 

reptiles. Pages 89-97 in G.W. Wood, ed. Prescribed fires and wildlife in southern forests: 

Proceedings of a symposium. Belle W. Baruch Forest Science Institute of Clemson University, 

Georgetown, SC. 

Melchiors, M.A. 1991. Wildlife management in southern pine regeneration systems. Pages 391-

420 in M.L. Duryea and P.M. Dougherty, eds. Forest Regeneration Manual. Kulwer Academic 

Publishers, The Netherlands. 

Mengak, M.T. and D.C. Guynn Jr. 2003. Small mammal microhabitat use on young loblolly pine 

regeneration areas. Forest Ecology and Management 173:309-317. 

Miller, J.H., R.S. Boyd, and M.B. Edwards. 1999. Floristic diversity, stand structure, and 

composition 11 years after herbicide site preparation. Canadian Journal of Forest Research

29(7):1073-1083. 

Mississippi State University Extension Service. 2004. Forest management for wild turkeys. 

Publication 2033. Available online at http://msucares.com/pubs/publications/p2033.htm; last 

accessed June 2005. 

Moore, S.E. and H.L. Allen. 1999. Plantation forestry. Pages 400-433 in M.L. Hunter Jr. (ed). 

Maintaining biodiversity in forest ecosystems. Cambridge University Press, New York. 
 14

Moorman, C. 2002. Burnin’ for wildlife. Forest Landowner 61(3):5-7. 

Noss, R.F. 1988. The longleaf pine landscape of the Southeast: Almost gone and almost 

forgotten. Endangered Species Update 5(5):1-8. 

Oliver, C.D. 1992. A landscape approach: Achieving and maintaining biodiversity and economic 

productivity. Journal of Forestry 90(9):20-25. 

Oliver, C.D. and B.C. Larson. 1990. Forest stand dynamics. McGraw Hill, New York. 419 p. 

Patel-Weynand, T. 2002. Biodiversity and sustainable forestry: State of the science review. 

Report for the National Commission on Science for Sustainable Forestry, Washington, DC. 54 

pp. 

Reed, D.P., R.E. Noble, and T.R. Clason. 1994. Effects of timber management activities on 

understory plant succession in loblolly pine plantations. Pages 109-113 in Proceedings of the 

47th Annual Meeting of the Southern Weed Science Society. 

Reid, W.V. and K.R. Miller. 1989. Keeping options alive: the scientific basis for conserving 

biodiversity. World Resources Institute, Washington, DC. 128 p. 

Robbins, L.E. and R.L. Myers. 1992. Seasonal effects of prescribed burning in Florida: A 

review. Miscellaneous Publication No. 8. Tall Timbers Research, Inc, Tallahassee, FL. 96 p. 

Schultz, R.P. 1997. Multiple-use management of loblolly pine forest resources. Pages 9-3 – 9-14 

in Loblolly Pine: The Ecology and Culture of Loblolly Pine (Pinus taeda L.). Agriculture 

Handbook 713. USDA Forest Service, Washington, DC. 

Sharitz, R.R., L.R. Boring, D.H. Van Lear, and J.E. Pinder, III. 1992. Integrating ecological 

concepts with natural resource management of southern forests. Ecological Applications

2(3):226-237. 

Siry, J.P. 2002. Intensive timber management practices. Pages 327-340 in D.N. Wear and J.G. 

Greis, eds. Southern Forest Resource Assessment. General Technical Report SRS-53. USDA 

Forest Service, Southern Research Station, Ashville, NC. 

Siry, J., F. Cubbage, and A. Malmquist. 2001. Potential impact of increased management 

intensities on planted pine growth and yield and timber supply in the South. Forest Products 

Journal 51(3):42-48. 

Stransky, J.J. and J.H. Roese. 1984. Promoting soft mast for wildlife in intensively managed 

forests. Wildlife Society Bulletin 12(3): 234-240. 
 15

Tappe, P.A., M.G. Shelton, and T.B. Wigley. 1993. Overstory-understory relationships in natural 

loblolly pine-hardwood stands: implications for wildlife habitat. Pages 613-619 in J.C. Brissette, 

ed. Proceedings of the Seventh Biennial Southern Silvicultural Research Conference. General 

Technical Report SO-93. USDA Forest Service, Southern Forest Experiment Station, New 

Orleans, LA. 

Thill, R.E. 1990. Managing southern pine plantations for wildlife. Pages 58-68 in Proceedings of 

the 19th IUFRO World Congress. Canadian International Union of Forestry Research 

Organizations, Montreal, Canada. 

Tucker, J.W., Jr., G.E. Hill, and N.R. Holler. 1998. Managing mid-rotation pine plantations to 

enhance Bachman’s sparrow habitat. Wildlife Society Bulletin 26:342-348. 

Turner, J.C., J.A. Gerwin, and R.A. Lancia. 2002. Influences of hardwood stand area and 

adjacency on breeding birds in an intensively managed pine landscape. Forest Science

48(2):323-330. 

Van Lear, D.H., R.A. Harper, P.R. Kapeluck, and W.D. Carroll. 2004. History of Piedmont 

forests: implications for current pine management. Pages 127-131 in K.F. Connor, ed. 

Proceedings of the 12th biennial southern silvicultural research conference. General Technical 

Report SRS-71. USDA Forest Service, Southern Research Station, Asheville, NC. 

Wahlenberg, W.G. 1946. Longleaf Pine: Its use, ecology, regeneration , protection, growth, and 

management. Charles Lathrop Pack Forestry Foundation, Washington, DC. 

Wicker, G. 2002. Motivation for private landowners. Pages 225-237 in D.N. Wear and J.G. 

Greis, eds. Southern Forest Resource Assessment. General Technical Report SRS-53. USDA 

Forest Service, Southern Research Station, Asheville, NC. 

Wigley, T.B., W.M. Baughman, M.E. Dorcas, J.A. Gerwin, J.W. Gibbons, D.C. Guynn Jr., R.A. 

Lancia, Y.A. Leiden, M.S. Mitchell, and K.R. Russell. 2000. Contributions of intensively 

managed forests to the sustainability of wildlife communities in the South. In Sustaining 

southern forests: The science of forest assessment. Southern Forest Resource Assessment. 

Available online at http://www.srs.fs.usda.gov/sustain/conf/ppr/wigley-ssf2000.pdf; last accessed 

June 2005. 

Wigley, T.B., K.V. Miller, D.S. deCalesta, and M.W. Thomas. 2002. Herbicides as an alternative 

to prescribed burning for achieving wildlife management objectives. Pages 124-138 in M.W. 

Ford, K.R. Russell, and C.E. Moorman, eds. The role of fire for nongame wildlife management 

and community restoration: traditional uses and new directions: Proceedings. General Technical 

Report NE-288. USDA Forest Service, Northeastern Research Station, Newton Square, PA. 

Wilson, M.D. and B.D. Watts. 1999. Response of brown-headed nuthatches to thinning of pine 

plantations. Wilson Bulletin 111(1):56-60. 
 16

Yin, R. and R.A. Sedjo. 2001. Is this the age of intensive management? A study of loblolly pine 

on Georgia’s Piedmont. Journal of Forestry 99(12):10-17.