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P.R. Kapeluck*, W.D. Carroll, D.H. Van Lear, E.A. Mikhailova, C. J. Post, M. A. Schlautman and W.M. Post
P.R. Kapeluck, W.D. Carroll, D.H. Van Lear, E.A. Mikhailova, C.J. Post, Department of Forestry and Natural Resources, Clemson University, 261 Lehotsky Hall, Clemson, SC 29634-0317; M.A.
Schlautman, Department of Environmental Toxicology, Clemson University, Clemson, SC
29634-0317; W.M. Post, Environmental Sciences Division, Oak Ridge National Lab., Oak Ridge,
TN 37831-6422.
*Corresponding author (pkplck@clemson.edu).

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17
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19
20 Rationale: The effects of the whole-tree and stem-only harvests on plant biomass, plant/soil
21 carbon (C) and nitrogen (N) stocks of naturally regenerated loblolly pine ( Pinus taeda L.) grown
22 in the Southeastern U.S. are poorly understood.
23 Objectives or hypothesis: This study was conducted to determine whether different tree-
24 harvesting methods could be manipulated to maximize plant biomass, plant/soil C and N
25 sequestration in naturally regenerated loblolly stands.
26 Methods: The effects of the whole-tree and stem-only harvests on the plant biomass, plant/soil C
27 and N stocks were investigated at the naturally regenerated loblolly pine stand at Clemson
28 Experimental Forest in Clemson, SC, on an abandoned agricultural field with eroded Pacolet soils
29 (Typic Kanhapludults).
30 Results:
31 A study at 5 yr found the S harvest had 29% more accumulated above-ground phytomass (live
32 and dead biomass) and 29 and 15% more C and N, respectively, than the W harvest. The trend
33 was similar at 16 yr with S harvest phytomass, C, and N levels 20, 20, and 103 % greater than the
34 whole-tree harvest. More growth and larger stocks of C and N for the S harvest may be related to
35 the release of elements contained in the logging debris during decomposition, N-fixation in
36 decomposing logging debris, and the environment it created. However, there were no statistically
37 significant differences in soil C and N between harvest methods.
38 Conclusions:

3
39 INTRODUCTION
40 Temperate forest ecosystems have been identified as potentially important for sequestering
41 carbon from the atmosphere for long-term storage (Tans et al. 1990) and harvesting effects on
42 forest productivity have been a concern for decades. As long ago as 1923, Wiedeman attributed
43 second rotation declines in productivity of spruce stands in Germany, Norway, and Switzerland to
44 soil degradation caused by clearcutting. However, solid evidence of reduced growth after
45 repeated harvests is limited to relatively few studies (Rennie 1955; Bednall 1968; Gholz et al.
46 1984).
47 More recently, concern has been expressed about effects on soil organic matter and forest
48 productivity when harvesting removes nearly all aboveground biomass (Johnson and Todd 1998).
49 Powers (1989) and Van Cleve and Powers (1995) suggested that productivity is ultimately related
50 to the amount and kind of soil organic matter. Johnson (1992) and Fisher (1995), while agreeing
51 that there is an intuitive relationship between soil organic matter and forest growth, noted that
52 there is seldom a strong statistical relationship.
53 Mann et al. (1988) described conventional vs. whole-tree harvesting impacts on nutrient
54 pools and regeneration at a variety of sites in the United States. This paper characterizes
55 productivity of loblolly pine ( Pinus taeda L.) of the Clemson, SC site 16 yr after treatment. We
56 hypothesize that, on eroded Piedmont sites, removal of logging debris during whole-tree
57 harvesting would reduce productivity and C and N sequestration.
58 MATERIALS AND METHODS
59 The study was located on the Clemson University Experimental Forest in the upper Piedmont of
60 South Carolina. The sites were within a 12.4 ha loblolly pine plantation established in 1939 on a
61 severely eroded nutrient-depleted agricultural field. Soil is an eroded phase of the Pacolet fine

4
62 sandy loam, a typic thermic Kanhapludult, and well drained. The coarse textured brown surface
63 layer ranged from sandy loam to sandy clay loam and was underlain by red, firm clay loam and
64 clay. Most of the surface soil had been lost during a century of farming (Trimble 1974) which
65 caused its depth to vary widely and frequently exposed the subsoil. Erosion essentially ceased
66 after plantation establishment.
67 The plantation occupied a hillside with an average slope of 13 % and S-SW aspect.
68 Average diameter breast height (DBH) was 25.5 cm and average height of dominant and
69 codominant trees was 23.2 m. Site index was 24.4 m at age 50 yr. Thinning from below at ages
70 22 and 30 yr had increased the original 2 x 2-m spacing to about 5 x 5 m, leaving an average of
71 437 stems ha
-1
and 18.5 m
2
ha
-1 basal area. Annual precipitation averages 130 cm and is well
72 distributed throughout the year.
73 Two watersheds (<2.0 ha/ea) within the plantation, separated by a 200-m-wide uncut area,
74 were clearcut in the winter of 1979 – 80, one using stem-only harvesting and the other whole-tree
75 harvesting. The methods of Van Lear et al. (1984) and pre-harvest DBH values generated
76 estimates of biomass and N content of logging debris remaining on the S harvest (20 Mg ha
-1
). All
77 debris was removed from the W harvest. Natural regeneration of both watersheds was from seed
78 produced by the surrounding stand on seedbeds prepared prior to harvest with prescribed fire.
79 Prior to the 1979 – 80 harvests, a 100% inventory of the 41-yr-old plantation was completed,
80 including the two study watersheds, for estimates of stem density, DBH, and basal area. Nutrient
81 content of the forest floor and soil just after harvest were reported in Van Lear et al. (1983).
82 Forest floor C content was estimated as 50 % (Taylor et al. 1991, Maier et al. 2004, Katul 2005)
83 of ash-free biomass. Five year post-harvest estimates of pine and other vegetation density,
84 biomass, and nutrient content were reported in Cox and Van Lear (1985). Residual pine root

5
85 system biomass and nutrient content just-after-harvest was determined as in Kapeluck and Van
86 Lear (1995). Carbon content of residual root systems was estimated as 50 % of biomass
87 (citation).
88
89 A comparison of preharvest data for both treatments is shown in Table 1. While the S
90 harvest has 26% more stems than the W harvest, both treatments have fewer stems than the
91 plantation average, and basal area for the W harvest exceeds the conventional harvest and the
92 plantation average. Higher stem density may explain the lower DBH in the S harvest. Greater
93 pre-harvest biomass (33%) on the W harvest suggests it was the better quality site.
94 Treatment effects were examined using estimates made just prior to harvest of the 41-yr-
95 old stand, just after harvest, 5 yr after harvest, and 16 yr after harvest. Estimates included 1),
96 density, biomass and nutrient content of the preharvest pine stands, 2), pine biomass and nutrient
97 content just after harvest, 3), forest floor biomass and nutrient content just after harvest, and 4),
98 nutrient content of the upper 60 cm of mineral soil just after harvest. Estimates at five years post-
99 harvest included pine density, biomass, and nutrient content and that of competing vegetation.
100 Sixteen years post-harvest estimates included 1), above-ground pine density, biomass, and C and
101 N content, 2), forest floor biomass and C and N content, and 3), C and N content of the upper 60
102 cm of mineral soil.
103 Carbon mass was estimated as 50% of biomass ( Wood Energy Hanbook, Maier et al.
104 2004, Katul 2005). Forest floor and soil nutrient content just after harvest were reported in Van
105 Lear et al. (1983). Residual pine root biomass and nutrient content just after harvest was
106 estimated as in Kapeluck and Van Lear (1995).

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107 Treatment effects on heights of the 16-yr-old regeneration were estimated from height-
108 pole measurements of 15 codominant and dominant trees from each treatment.
109 Aboveground biomass of 16-yr-old stands was estimated using equations developed from
110 16 trees representing the range of diameters present in the two treatments as in Van Lear et al.
111 (1984). These equations were applied to diameter-breast-high (1.4 m) of trees on three randomly
112 located 10 x 20 m plots/treatment. Each plot contained 20-30 pine trees. Necromass was
113 estimated the same way from 16 standing dead trees/treatment. Pine root biomass and nutrient
114 content of the 16-yr-old stand was estimated as in Kapeluck and Van Lear (1995). Foliage,
115 branch, bole, necromass, forest floor, and soil C and N of the 16-yr-old stands were determined on
116 a Perkin-Elmer 240C Analyzer.
117 Soil was sampled at 0-8, 8-30, and 30-60 cm depths in each plot from cores (2.5-cm dia)
118 taken at three points/plot/treatment. Soil mass excluded rock and pine root volume determined in
119 previous studies (Kapeluck and Van Lear 1995, Parker and Van Lear 1996). Concentrations were
120 multiplied by compartment mass to determine elemental content.
121
123
Height for the 16-yr-old regeneration were . Differences in phytomass components and
122 elemental concentrations were detected using t tests.
Equations were developed using SAS (1992).
124
125
126
RESULTS
An essential estimate needed to assess the effects of the two harvesting treatments was an
127 evaluation of comparative productivity of the two areas prior to treatment. Stem analysis of the
128 mature trees growing along the boundaries of each treatment provided one detail to answer this
129 question. Comparison of height/age growth patterns revealed that growth on the two treatments

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130 were different during the 41 yr preceding treatment (Fig. 1). Although the average height growth
131 of the conventional harvest treatment was only 1.6 m less than the whole-tree harvest treatment
132 after 41 yr, the difference between regressions was significant (p = 0.001). More importantly, this
133 difference corresponds with the other pretreatment estimates which indicate that productivity of
134 the conventional harvest treatment was the lesser of the two treatments.
135 Table 2 compares mass and elemental content of residual pine biomass (above and
136 belowground), forest floor, and mineral soil for both treatments just after harvest operations were
137 completed. The totals found at the bottom of the table provide some explanation for the greater
138 preharvest productivity of the whole-tree harvest treatment area. Only overall biomass of the
139 conventional harvest treatment, which includes 20 Mg ha
-1
of residual crown biomass, is higher
140 (11.7%) than that of the whole-tree harvest treatment. For element content, the whole-tree harvest
141 treatment ranges from about 5 to 128% greater than the conventional harvest treatment. Among
142 the three compartments, differences between treatments for C, N, and P were greatest in the forest
143 floor. The 22% greater forest floor biomass of the whole-tree harvest area implies its greater
144 productivity and reflects its greater pretreatment foliage biomass. For K and Ca, differences
145 between treatments were greatest in mineral soil where the whole-tree harvest treatment contained
146 about twice the amount of the conventional harvest treatment. Forest floor differences between
147 treatments for K and Ca were also large, at 23 and 61%, respectively.
148
149
After the first growing season the whole-tree harvested treatment had more than twice the regeneration of the conventional harvest treatment with 82,000 and 33,000 stems ha
-1
,
150 respectively. At the end of the fifth growing season, regeneration density of the conventional
151 harvest and whole-tree harvest treatments had declined to nearly equal numbers, 23,000 and
152 20,500 stems ha
-1
, respectively. However, at the end of the fifth growing, total biomass of

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153 herbaceous vegetation, hardwood sprouts, and pine saplings on the conventional harvest treatment
154 comprised about 29% more than that on the whole-tree harvest treatment (Table 3). Although
155 other species continued to be present in the stand their fraction of total plant biomass had declined
156 sharply during the previous five years as pine regeneration had rapidly become the dominant
157 vegetation. At that time pine biomass when averaged for both treatments made up about 78% of
158 total vegetation biomass. Because of high regeneration density crown closure was apparently
159 already underway on both treatments.
160 During the next 11 yr live pine density of the whole-tree harvest treatment continued its
161 more precipitous decline until at age 16 yr it contained about 35% fewer live stems (Table 4).
162 Standing dead stems on the conventional harvest treatment were also more numerous (17%) than
163 those on the whole-tree harvest treatment. Combined live and dead stems on the conventional
164 harvest treatment were about 38% more abundant than on the whole-tree harvest treatment.
165 Further attrition of live stems through self thinning during the years from 5 to 16
166 eliminated about 50% of the regeneration on the whole-tree harvest treatment in contrast to the
167 30% loss on the conventional harvest treatment. These comparisons suggest that the environment
168 of the whole-tree harvest treatment was considerably less favorable for seedling and sapling
169 survival. Although not significant, and perhaps unexpected, mean dbh of both live and dead
170 stems on the whole-tree harvest treatment are slightly larger than those of the conventional
171 harvest treatment (Table 4). This trend may result from the distribution of site resources among
172 fewer trees.
173 By age16 yr loblolly pine regeneration completely occupied the growing space on both
174 treatments which created a deeply shaded understory nearly free of other plants. Phytomass
175 accumulation after 16 yr were similar to those observed after 5 yr (Fig. 2), which continued the

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176 trend of lower productivity on the whole-tree harvest treatment. Tree boles, the largest and only
177 component significantly different between treatments, were 35% greater on the conventional
178 harvest treatment. Boles comprised 60% and 68% of the total above-stump phytomass for the
179 whole-tree and conventional harvest treatments, respectively (Table 5).
180 As discussed, mortality rates were high during the 16 yr prior to these measurements
181 because of intense intraspecies competition as trees continued to differentiate into crown classes.
182 Standing necromass at age 16 yr was 10.9 Mg ha -1 for the whole-tree harvest treatment and 12.0
183 Mg ha
-1
for the conventional harvest treatment (Table 5). Most dead trees remained standing
184 because high stand densities on both treatment areas prevented their falling. When necromass
185
186 was combined with biomass, total aboveground loblolly pine phytomass was 19.6% more (141.4 vs. 118.2 Mg ha
-1
) for the conventional harvest treatment. In comparison, Wells et al.(1975)
187 reported 156.0 Mg ha -1 of aboveground biomass for a 16-yr-old loblolly pine plantation with
188 2,243 trees ha
-1
(approximately 1/7 and 1/5 the density of our stands) on an excellent site in the
189 North Carolina Piedmont.
190 Our stands are growing on eroded and nutrient deficient soil that often experiences
191 moisture deficits during the growing season (Myers et al. 1986). Extremely high seedling density
192 probably initiated intense competition for nutrients, water, light, and growing space during the
193 first growing season after the treatments were applied. Thus, treatment-induced stress may have
194 been reflected earlier in these dense stands than in more widely spaced plantations, especially on
195 poor quality sites. Since pine density remained extremely high throughout the study period the
196 magnitude of growth differences between treatments may have been amplified.
197 Although biomass productivity was about 20% greater on the conventional harvest
198 treatment, no differences between treatments could be detected in height/age growth patterns of

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199 the regeneration during their 16 yr of growth (Fig. 3). Nor were differences found in the mean
200 stand heights of the treatments (Table 6).
201 A partial explanation for similar height/age growth patterns and stand height may be the
202 more rapid self thinning that occurred on the whole-tree harvest treatment. As previously
203 discussed, high stem density intensified competition for finite resources on both treatments. And,
204 although stem density had declined dramatically, especially on the whole-tree harvest treatment, it
205 remained very high for both treatments. The continued high density exerted the competitive
206 stimulation for height growth on both treatments, but, for the whole-tree harvest treatment, its
207 apparently more limited resources could be concentrated in fewer surviving trees. Thus,
208 comparable height growth was maintained. Smith (1986) postulated that on poorer sites, seasonal
209 moisture deficiencies or other soil factors limit production more at high stand densities than at
210 intermediate levels consistent with full or nearly full occupancy.
211 It is also possible that the codominant and dominant crown classes on the whole-tree
212 harvest treatment are primarily composed of trees growing near the old decomposing root systems
213 of the harvested 41-yr-old stand. How trees growing in these locations benefit will be discussed
214 later. On the whole-tree harvest treatment these trees would have a distinct competitive advantage
215 over other trees because the other trees would be subjected to the evident detrimental treatment
216 effects without the mitigation offered by the old root systems.
217 The greater aboveground mass on the conventional harvest treatment would be expected to
218 have correspondingly larger mass of coarse roots to maintain allometric relations between tops
219 and roots (Waring and Schlesinger 1985). Van Lear and Kapeluck (1995) demonstrated such a
220 relationship between tops and roots for mature loblolly pine on this site. Van Lear et al. (2000)
221 found that decomposing root systems of harvested loblolly pines provide a favorable medium for

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222 fine root growth and support a high density of such roots. Sixteen-year-old loblolly pines tended
223 to be larger and grew at greater densities close to these decomposing root systems. We believe
224 that tops of trees on the conventionally harvested site, as they decomposed, provided similar
225 microsites of enhanced rooting potential, i.e., less resistance to root penetration, greater aeration,
226 higher nutrient concentrations, and higher moisture-holding capacity, than the surrounding
227 mineral soil. Better quality rooting habitat and increased rooting volume offered by the
228 decomposing wood of the tops on the conventionally harvested treatment may account for some
229 of the greater productivity.
230 Other potential benefits of logging slash included shading the 40 to 60% of the soil surface
231 that it covered. After harvesting much of the solar load formerly used for evapotranspiration
232 heats the air, forest floor, and soil (Geiger 1965). Evaporation from forest floor and soil increase,
233 but the most important effect is the increase of temperature extremes. High temperatures increase
234 the decomposition rate of the forest floor and on bare soil surface temperatures have been
235 recorded as high as 58 C and a 15-day average of 48 C at 0.6 cm depth (USDA 1951). High
236 temperatures like these inhibit both survival and growth of seedlings. High soil temperatures also
237 dessicate the surface soil and reduce it to a structureless mass.
238 Shading from the residual crowns would moderate surface temperatures, reduce
239 evapotranspiration, lower moisture stress, and help maintain the structural integrity of the friable
240 and nutrient rich surface soil. This would have benefitted the trees on the conventional harvest
241 treatment since soil moisture has a greater influence on root and top growth than any other soil
242 factor because it effects nutrient availability, uptake, and transport (Pritchett 1979). Slash would
243 also provide a mulching effect which aids root growth (Bilan 1968) while preventing increases in
244 soil strength because soil strength increases sharply as it dries which limits root growth (Zahner

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245 1968). Also, the scattered crowns formed a tall (3m) barrier to horizontal air movement near the
246 ground which increases after harvesting (Hursh 1948). This barrier effectively shifts the
247 temperature, humidity, and wind movement boundary layer above the soil surface (May 1977)
248 which provides a diffusion barrier for moisture and shades the surface from direct insolation. The
249 result is improved conditions for survival and growth of trees and greater protection of the
250 desirable properties of forest soils such as low density, high porosity, and high infiltration rates
251 (Nutter and Douglass ______?).
252 The S - SW aspect of both treatments caused daily exposure to long periods of direct
253 insolation. Without logging slash the whole-tree harvest treatment’s environment was not
254 moderated by the benefits discussed above and moisture loss/stress was undoubtably greater. Tree
255 and root growth slows or ceases with water deficits through interference with photosynthesis,
256 nitrogen metabolism, salt absorption, translocation, and cell enlargement (Kramer 1969, Pritchett
257 1979). The absence of these moderating factors may also have contributed to the productivity
258 differences.
259
260
261 Based on differences in nutrient accumulation in phytomass, the growth differences
262 between the two treatments appears to be primarily nutritional. At age 5 yr, the regenerating plant
263 community on the conventional harvest treatment had accumulated 13-15% more N, P, K, and Ca
264 in aboveground biomass (Table 7), indicating that larger quantities of nutrients were absorbed
265
266 early. Nutrient accumulation in aboveground phytomass at age 5 yr was very high on both treatments, e.g., 156 and 136 kg ha
-1
N for the conventional and whole-tree harvest treatments,
267 respectively. The extremely high densities of pine saplings in these stands may have allowed

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268 more complete capture of available elements and more rapid element accumulation in pine
269 phytomass in contrast to the far lower pine densities found in plantations. In addition, the woody,
270 more decay resistant qualities of pine tissue compared to the rapid turnover rate of herbaceous
271 plant tissue may have slowed the nutrient turnover rate in these natural stands compared to
272 plantations. This would follow from the greater dominance and early canopy closure by pine in
273 these natural stands compared to plantations. Nemeth (1973) reported a decrease in turnover rate
274 of ground litter to soil and a net decline of the net primary production of the lesser vegetation of
275 89% associated with canopy closure in pine plantations. As competition gradually reduced pine
276 density, the woody tissues of dead pines would have decomposed and released nutrients relatively
277 slowly.
278 The lower turnover rate of woody pine tissue may have also contributed to a tighter
279 nutrient cycle because pine root occupancy of the soil would have been consistently at or near
280 maximum because of constant high stand density. High stem density is concomitant with equal
281 numbers of root systems which would have produced a densely populated subsurface mat of
282 interwoven pine roots. This root mat would have been present almost continuously from the first
283 through the sixteenth growing season. Therefore, in these natural stands, pine roots would have
284 been more likely to absorb available elements and those released during detrital decay rather than
285 their being lost to leaching. However, element conservation may have been less effective on the
286 whole-tree harvest treatment because of its more rapid decline in stem density/root systems,
287 perhaps allowing greater leaching losses.
288 Aboveground phytomass on the conventional harvest treatment at age 16 yr contained
289 markedly greater quantities of nutrients. Nitrogen, P, K, and Ca were 2.0, 1.7, 1.2, and 1.4 times,
290 respectively, more than the whole-tree harvest treatment. Nitrogen content of the natural stand on

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291 the conventional harvest treatment was much larger than values reported for similarly aged
292 plantations of southern pines, e.g., more than twice that of a 16-yr-old loblolly plantation on an
293 excellent site in the Piedmont of North Carolina (Wells et al. 1975) and 50% greater than that of a
294 fertilized 15-yr-old slash pine plantation in Florida (Pritchett and Smith 1974).
295 The large differences in nutrient accumulation by stands on the two treatments relates to
296 land-use history, as well as to effects of treatment on nutrient availability and uptake. Upper
297 Piedmont soils were depleted of nutrients and organic matter by a century of row cropping for
298 cotton, tobacco, and other crops prior to reforestation in the 1930s (Giddens 1957; Trimble 1974;
299 Van Lear and Kapeluck 1995). The absence of logging debris following whole-tree harvesting
300 may jeopardize productivity on sites with low levels of available N (Weetman and Webber 1972;
301 van den Driessche 1991). Nitrogen often limits growth on Piedmont sites, but responses to N
302 fertilization by Piedmont plantations are generally short lived (Van Lear 1980; Wells and Allen
303 1985; Jokela et al. 1991). Therefore, the slowly released N in logging debris may be important to
304 the productivity of these soils.
305
306
Estimates of elemental content of logging debris (Van Lear et al. 1984) were approximately 53, 5, 19, and 37 kg ha
-1
of N, P, K, and Ca, respectively (Table 8). About 50% of
307 the N and P in this debris is in relatively nutrient-rich and readily decomposable foliage. Some of
308 these nutrients would have been available for plant uptake during the regeneration period and
309 probably enhanced early growth on the conventional harvest treatment. In addition, the mulching
310 effect of the debris, which covered about 40 - 60% of the area (based on aerial photographs and
311 tree crown calculations), probably conserved soil moisture creating a more favorable rooting
312 environment during the regeneration period. As the woody portions of logging slash decomposed,
313 it probably became an enhanced medium for fine root growth much in the same manner that

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314 decomposing root systems do (Van Lear et al. 2000). Enhanced root growth would have allowed
315 trees on the conventionally harvested treatment to absorb greater quantities of nutrients. Nitrogen,
316 P, and Ca in regrowth on the conventional harvest treatment diverged sharply from that of the
317 whole-tree harvest treatment between ages 5 to 16 yr (Fig. 4).
318 Phytomass nutrient differences between the two treatments (Table 9) were larger than
319 absolute quantities of nutrients in logging debris. Bole and branch components in the 16-yr-old
320 stand accounted for most of the unexpectedly large quantities of nutrients on the conventional
321 harvest treatment. Boles there contained 3.8, 2.5, 2.0, and 1.6 times more N, P, K, and Ca,
322 respectively, than those on the whole-tree harvest treatment, although bole mass was only 35%
323 greater on the conventional harvest treatment.
324 Differences in nutrient content of boles were a result of large differences in nutrient
325 concentrations, rather than in phytomass (Table 10). Nitrogen concentrations of boles from the
326 conventional harvest treatment were almost three times higher than those of the whole-tree
327 harvest treatment, while P concentrations were twice as high. There were significantly greater
328 concentrations of N, P, and K in branches (live and dead) of trees on the conventional harvest
329 treatment. Nitrogen concentrations in foliage from the two treatments did not differ. Bole
330 concentrations of N reported here for the conventional harvest treatment are higher than others
331 reported in the literature for pine plantations (Wells et al. 1975, Tew et al. 1984). We have no
332 reason to doubt the accuracy of our reported concentrations. As seen in Table 6, concentrations of
333 all elements in boles of trees in the conventional harvest treatment were higher and had small
334 standard errors. Other support for unbiased concentrations is that the elemental analysis was
335 performed at an independent lab and the sample vials were only identified by sequential numbers.

16
336 Reasons for higher concentrations of nutrients in phytomass on the conventional harvest
337 treatment have not been determined, although the literature suggests several processes may be
338 involved. The gradual decay of logging debris, acting on a burned seedbed, may have enhanced
339 nutrient supply and availability, which acted as a starter dose of fertilizer while stimulating N
2
340 fixation. Reported rates of N fixation by free-living diazotrophs and N-fixing plants (Richards
341 1964; Roskoski 1980; Stevenson 1986; Jurgensen et al. 1987; Perry et al. 1987) range from
342 relatively insignificant amounts to well over 150 kg ha -1 yr -1 . Nitrogen supply and availability
343 also may have been enhanced by termites (Lee and Wood 1971; Behnke 1977; Brian 1978),
344 which were observed in logging debris.
345 Large N accumulations in pines have been attributed to associative N
2
fixation in the
346 rhizospheres (Richards 1973; Bormann et al. 1993). Although evidence that associative N
2
347 fixation exists in pines is convincing, organisms and processes involved are not well understood.
348 However, the moderated environment under and around decomposing logging debris may favor
349 microflora, including beneficial rhizosphere organisms. In addition to associative fixation,
350 mycorrhizal fungi have the ability to obtain N from organic sources and pass it on to conifers.
351 They also increase P availability (Harley and Lewis 1969; van den Driesssche 1991). It has also
352 been postulated that the pine rhizosphere mineralizes some fraction of soil N and P (Voight 1965;
353 Fisher and Stone 1969), providing another source of nutrients. The greater root mass assumed the
354 conventional treatment would have more rhizosphere, thus greater nutrient uptake would be
355 expected.
356 Some or all of these soil processes were operating in both treatments but those in the
357 conventional harvest treatment were evidently enhanced by debris remaining on the site.
358 However, we suggest that the enhanced rooting value of the decomposing tops of harvested trees

17
359 is the most likely reason for the increased growth on the conventional harvest site. We believe
360 that decomposing tops provide a medium for enhanced fine root growth and root viability,
361 allowing greater nutrient uptake and accumulation in boles of trees on the conventional harvest
362 site. The network of decomposing tops scattered over the site essentially improved site fertility
363 (the ability of the site to supply nutrients) and thereby increased its productivity.
364 A summary of the status of the three ecosystem compartments 16 yr after the application
365 of treatments is shown in Table 11 in the same format as Table 2 for ease of comparison. Pine
366 phytomass on the conventional harvest treatment has exceeded pretreatment mass by 41% while
367 the whole-tree harvest treatment is 11% below pretreatment mass. Elemental accretion in pine
368 phytomass reflect the different rates of stand aggradation between treatments resulting in a
369 reversal in ranking of pine elemental capital. Swaps in forest floor mass ranking correspond to
370 the trends exhibited by the regenerating stands. Forest floor mass increased by16% for the
371 conventional harvest treatment during the past 16 yr while the whole-tree harvest treatment lost
372 14%. Similarly the ranking of elemental content of the forest floors also reversed except for
373 Ca.Ranking changes in mineral soil elemental content occurred only for C and P. The 29% loss in
374 C content for the whole-tree harvest treatment during the 16 yr since treatment may be related to
375 its lost productivity and therefore lower C inputs while detrital catabolic rates remained constant.
376 Nitrogen levels have remained about the same and any soil losses could have moved into pine
377 biomass or forest floor. Soil P on the conventional harvest treatment has increased dramatically
378 (296%) while the whole-tree harvest treatment increased a comparatively modest 37%. Potassium
379 in soil declined 69 and 62% for the conventional and whole-tree harvest treatments, respectively,
380 perhaps relocated in pine biomass. Calcium levels in soil have increased (55%) for the

18
381 conventional harvest treatment and decreased (-9%) for the whole-tree harvest treatment.
382 Calcium depletion citations/discussion.
383 Sums of elemental capital of the three ecosystem compartments estimated at the time of
384 treatment and16 yr later are compared in Table 12. This summary is perhaps the most dramatic
385 indication of the long-term treatment effects. During the 16 post-treatment years the conventional
386 harvest treatment has increased its combined elemental capital in all five elements from 16 to
387 197%. If C is included, the average elemental gain is 92%. For elements N, P, K, and Ca alone
388 the average increase is 34%. As a C sink, the conventional harvest treatment has nearly doubled
389 the quantity present in logging slash, forest floor, roots, and mineral soil on the site just after
390 harvest.
391 In marked contrast, changes in elemental capital for the whole-tree harvest treatment
392 consisted of more modest gains of 2 to 53 % combined with a 20% loss of K and a 0.2% loss of
393 Ca. If C is included, the average elemental gain is 50% and for N, P, K, and Ca alone the average
394 increase is only 2%. As a C sink, the whole-tree harvest treatment has increased its content by
395 53% since treatment. This quantity represents about 60% of the rate of C sequestration of the
396 conventional harvest treatment.
397 Although statistics are not possible for comparisons of the summary data in Table 12, the
398 magnitude of the differences in treatments strongly implies the detrimental effects of whole-tree
399 harvesting. Sixteen years after the treatments were applied the ranking of the original qualities of
400 the two sites has essentially been reversed.
401 If organic matter and associated nutrients from decomposing logging slash are stored
402 preferentially in old root channels and new roots disproportionately use them, should we not look
403 there to detect relationships between soil C and productivity? Old root channels are dynamic

19
404 elements of forest soils, beginning as dying roots and proceeding through various stages of decay.
405 They are hollowed from the inside by termites and other organisms, leaving the outside of the
406 opening intact. Gradually, the hollow core of the large taproot and major laterals are filled with
407 mineral and organic matter by soil fauna activity, as well as with roots from the developing plant
408 community. In time, perhaps decades or centuries, the old root channels blend into the soil
409 matrix.
410 The root distribution of young loblolly pine trees suggests that we sample more intensively
411 in or near decomposing root systems of harvested trees if we wish to detect relationships between
412 organic matter and productivity. Deposition of decaying logging debris in old root channels by
413 soil fauna and heavy utilization of these root channels by roots of the developing stand may
414 account for the relatively long-term positive growth response on the conventionally harvested
415 area.
416
417
Although soil carbon levels were not significantly different based on our random sampling design it is possible that some of the 10 Mg ha
-1
C contained in the logging slash could be
418 detected if the points where the slash decomposed were sampled specifically. The greatest
419 quantity of carbon in logging slash is found in the larger woody portions. The woody parts of
420 each crown occupy only a small fraction of the soil surface and it is unlikely that their residual
421 carbon would diffuse very far in surrounding soil. Therefore, to detect carbon from decomposed
422 logging slash, requires a stratified scheme that samples points where logging slash decomposed as
423 well as the rest of the stand. Otherwise, the probability of sampling these points would be low
424 and the greater carbon content of these samples would be masked by the predominantly lower
425 content of samples taken elsewhere in the stand.
426

20
427 DISCUSSION
428
429
430
SUMMARY AND CONCLUSIONS
Whole-tree harvesting should be carefully scrutinized before applying the practice on
431 marginal sites in the Piedmont. The naturally regenerated loblolly pine on the conventional
432 treatment grew significantly faster (20% more phytomass after 16 yr) than the that on the whole-
433 tree treatment. The major portion of this growth differential is attributed to greater nutrient
434 availability and uptake on the conventional treatment.
435 Although not found in this study, perhaps differences in height/age growth patterns will
436 emerge in later years as the stands mature and internal transfer and recycling elements become the
437 more dominant metabolic strategy. This may result as the more limited nutrient capital is diluted
438 in accumulating biomass and concentrations fall to levels which limit metabolic functions and
439 height growth.
440 This study indicates that nutrients released gradually during natural decomposition of
441 logging debris, in combination with other debris-mediated mechanisms which allow additional
442 nutrients to become available, produce a long-term sustained growth response in dense natural
443 pine stands on poor sites. This long-term increase in nutrient supply and availability, as indexed
444 by nutrient uptake, is critical to loblolly pine growth on poor sites when growth rates are
445 accelerating. Results from this study suggest that leaving logging debris in place is extremely
446 important to sustaining productivity on marginal sites.
447 The marked deterioration of the whole-tree harvested treatment from pretreatment levels
448 may be linked to the initial degraded quality of the old field site. With only 41 yr under loblolly
449 pine forest, the reclamation of pre-agricultural site quality would have only just begun. The hard

21
450 won accumulation of elements and organic matter appear to have been in a fragile and tenuous
451 state. The conservative conventional harvest used here, with crowns left where they fell (not
452 piled), allowed better retention and continuation of the capacity added to these poor quality sites
453 by the 41-yr-old loblolly pine plantation. The continued improvement of the conventional harvest
454 treatment further suggests that, with conservative management that duplicates the method used
455 here, successive rotations of loblolly pine will continue to improve productivity. Thus, site index
456 in the future may rise incrementally with each new stand.
457 As was expected the relationship between soil carbon and productivity was poor in this
458 study. Perhaps this is because we do not completely understand the complex pathways of detritus
459 decomposition and because it is difficult to separate effects of C on productivity from those of
460 associated nutrients. However, relationships between forest productivity and soil carbon may
461 become clearer if soils are examined in those areas where biological activity is greatest, i.e., in
462 and adjacent to old root channels and other points where organic matter is concentrated.
463
464
ACKNOWLEDGMENTS
This study was funded by the National Council of the Paper Industry on Air and Stream
465 Improvement, Inc. and McIntire-Stennis. Funding secured by Dale W. Johnson of the Desert
466 Research Institute was instrumental to the initial and latter phases of this study.
467

22
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560 Taylor, et al. 1991. Four Rocky Mountain Coniferous Forests. Canadian Journal of Botany
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573 practices and the nutrient status of a loblolly pine plantation. p. 252-258. IUFRO
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577 30 :395-404.
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579
581 content of a mature loblolly pine plantation. Can. J. For. Res. 25 :361-367.
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27
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584 affected by decomposing root systems. Forest Ecology and Management __:1-9.
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590 two spruce stands. Can. J. For. Res. 2 :351-369.
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592
594 unteren Hohenlagen der Sachsischen Staatsforsten. Tharandt. (Transl. 302) USDA.
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595 Southeast. For. Exp. Stn., Asheville, NC.
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597
598 management guide. USDA Forest Serv., Southeast. Forest Exp. Sta., Ashevillle, N.C.
Gen. Tech. Rep. SE-36.
599 Zahner, R. 1968. Water deficits and growth of trees. In Water Deficits and Plant Growth. Vol. 2.
Academic Press, Inc., New York. pp. 191-254. 600
601
602

28
603 Table 1. Preharvest characteristics of the 41-yr-old loblolly pine for the conventional harvest
604 treatment and whole-tree harvest treatment and plantation.
605
606
607
608
609
610
611
612
613
614
617
618
Treatment Stem Frequency DBH Basal Area Aboveground Biomass
(stems ha
-1
) (cm) (m
2
ha
-1
) (Mg ha
-1
)
Conventional 399 21.7 (0.407)

17.1 100.0
Whole-tree 316 29.0 (0.331) 21.7 133.2
Plantation 437 25.5 18.5 109.6

Standard errors.
619

29
629
630
631
632
633
634
635
636
637
638
639
640
641
642
643
644
620
621
622
623
624
625
626
627
628
Table 2. Comparison of ecosystem components of the conventional (C) and the whole- tree harvest treatment (W) just after harvest of the 41-year-old loblolly pine stand.
________________________________________________________________________
Treatment Component Biomass C N P K Ca
(Mg ha -1 )
(kg ha
-1
)
C Pine 44.4

22,420 95.4 10.4 47.2 71.4
W Pine 33.3

16,651 55.7 6.9 38.1 45.2
C Forest Floor 16.9 8,441 132.3 12.7 17.5 102.5
W Forest Floor 21.6 10,246 176.1 17.2 21.6 164.8
C Mineral Soil — 42,338  2,115.3 12.6 109.0 501.0
W Mineral Soil — 49,916 2,333.0 16.7 213.0 1332.0
________________________________________________________________________
Totals C 61.3 73,199 2,343.0 35.7 173.7 674.9
W 54.9 76,813 2,564.8 40.8 272.7 1,542.0
________________________________________________________________________

Includes residual crowns and root systems.

Includes only residual root systems.
 Soil values represent the upper 60 cm.

30
653
654
655
656
657
658
645
646
647
648
649
650
651
652
Table 3. Biomass and nutrient accretion in regrowth 5 years after stem-only (S) and whole-tree (W) harvest of mature loblolly pine in the Piedmont of South Carolina.
________________________________________________________________________
Treatment Biomass C N P K Ca
(Mg ha
-1
) (kg ha
-1
)
S 20.8 10,388

156.3 14.3 76.6 74.6
W 16.1 8,033 136.1 12.4 67.9 82.8
________________________________________________________________________

Carbon quantities are 50% of biomass values.

31
671
672
673
674
675
676
659
660
661
662
663
664
665
666
667
668
669
670
Table 4. Frequency and dbh for live and dead loblolly pine natural regeneration on stem-only and whole-tree harvest treatments in the Piedmont of South Carolina.
_______________________________________________________________________
Live Stem Dead Stem
Treatment Frequency DBH Frequency DBH
(No. ha
-1
) (cm) (No. ha
-1
) (cm)
________________________________________________________________________
Stem-only 15,954 5.21 (0.083)

10,451 1.80 (0.031)
Whole-tree 10,380 6.34 (0.105) 8,928 2.03 (0.034)
________________________________________________________________________

Numbers in parentheses are standard errors.

32
677
678
679
680
681
682
683
684
685
686
687
688
689
690
691
692
693
694
695
696
697
698
Table 5. Phytomass (biomass plus necromass) of naturally regenerated 16-year-old loblolly pine stands in the Piedmont of South Carolina following whole-tree and stem-only harvesting.
________________________________________________________________________
Tree
Component
Stem-only _
(Mg ha
-1
) (%)
Whole-Tree __
(Mg ha
-1
) (%)
________________________________________________________________________
Foliage
Dead branches
9.9a
*
10.8a
7.0
7.7
10.7a
11.4a
9.0
9.6
Live branches
Boles
Necromass
12.4a
96.3a
12.0a
8.8
68.1
8.4
13.9a
71.3b
10.9a
11.8
60.3
9.2
________________________________________________________________________
Total 141.4a 100.0 118.2b 100.0
________________________________________________________________________
Note:
*
Means within a row followed by different letters are significantly different at the 95% confidence level.

33
708
709
710
711
712
713
714
699
700
701
702
703
704
705
706
707
Table 6. Mean heights of 16-yr-old natural regeneration following conventional and whole-tree harvests in the Piedmont of South Carolina.
________________________________________________________________________
Treatment n Height
(m)
________________________________________________________________________
Conventional 15 9.82 (1.35)

a
Whole-tree 15 9.65 (1.40) a
________________________________________________________________________

Numbers in parentheses are standard errors.
Heights followed by the same letter are not significantly different at the 95% confidence level.

34
715
716
717
718
719
720
721
722
723
724
725
726
727
728
729
730
731
732
733
734
735
736
737
Table 7. Phytomass and nutrient content of aboveground regrowth 5 and 16 yr after whole-tree and conventional harvest of a mature loblolly pine plantation.
______________________________________________________________________________
Treatment Phytomass Nutrient
N P K Ca
____________(Mg ha
-1
)_______________________(kg ha
-1
)_____________________________
5th Year
136.1 12.4 68.0 82.8 Whole-tree 16.1
Conventional 20.8 156.3 14.3 76.6 74.6
Whole-tree 118.2
16th-Year
255.6 20.0 98.9 195.4
Conventional 141.4 519.2 33.7 118.1 267.8
______________________________________________________________________________

35
746
747
748
749
750
751
752
753
754
755
756
738
739
740
741
742
743
744
745
Table 8. Phytomass and nutrient content of logging debris following conventional harvest of mature loblolly pine plantations in the Upper Piedmont of South Carolina.
______________________________________________________________________________
Component Phytomass Nutrient________________
N P K Ca
___________________(Mg ha
-1
)__________________________ (kg ha
-1
)__________________
Stemwood
Stembark
4.2
0.4
2.7
0.7
0.2
0.1
1.8
0.2
3.3
0.7
Branches 12.7 23.8 2.3 10.8 28.3
Foliage 2.5 26.4 2.6 5.8 5.2
______________________________________________________________________________
Total 19.8 53.6 5.2 18.6 37.5
______________________________________________________________________________

36
776
777
778
779
780
781
782
783
784
785
786
787
766
767
768
769
770
771
772
773
774
775
757
758
759
760
761
762
763
764
765
788
789
790
Table 9. Phytomass and associated nutrient content for 16-year-old naturally regenerated loblolly pine in the Piedmont of South Carolina following conventional and whole-tree harvests.
______________________________________________________________________________
Component Phytomass Conventional
(Mg ha
-1
) C N P K Ca__
_____________________________________________(kg ha
-1
)_________________________
Foliage 9.9
Dead Branch 10.8
Live Branch 12.4
5.0

90.3 9.9 37.7 26.4
5.4 51.6 2.2 4.3 30.3
6.2 54.6 4.6 19.9 27.9
Boles 96.3 48.2 300.2 14.5 53.9 153.6
Necromass 12.0 6.0 22.5 2.5 2.3 29.6
Roots 35.4 17.7

59.1 7.4 40.4 48.0
Total 176.8 88.5 578.3 41.1 158.5 315.8
------------------------------------------------------------------------------------------------------------------
Component Phytomass
(Mg ha
-1
) C N
Whole-Tree
P K Ca___
_____________________________________________(kg ha
-1
)__________________________
Foliage 10.7
Dead Branch 11.4
Live Branch 13.9
Boles 71.3
5.4 98.0
5.7 24.8
7.0 32.1
35.7 79.5
9.0
1.1
2.8
5.8
52.8
0.8
15.8
27.4
28.4
25.0
26.9
93.5
Necromass 10.9 5.5 21.2 1.3 2.1 21.6
Roots 29.6 14.8 49.1 6.1 33.8 40.1
Total 147.8 74.1 304.7 26.1 132.7 235.5
______________________________________________________________________________

Carbon values for all components represent 50% of phytomass.

Element content of roots from Van Lear and Kapeluck (1995).

37
812
813
814
815
816
817
818
819
820
805
806
807
808
809
810
811
791
792
793
794
795
796
797
798
799
800
801
802
803
804
Table 10. Concentration means (n=16) of tree components from naturally regenerated 16-yr-old loblolly pine in the Upper Piedmont of South Carolina following conventional (C) and whole- tree (W) harvests.
______________________________________________________________________________
Tree
Component Trt. N P K Ca
______________________________________________________________________________
Bole
Bole
C .312a
*
(.026)
#
.015a (.002) .056a (.003)
W .112b (.010) .008b (.001) .038b (.006)
.16a (.032)
.13a (.015)
Live Brch.
Live Brch.
C
W
.440a (.034) .037a (.003) .160a (.014)
.230b (.017) .020b (.002) .114a (.015)
.23a (.026)
.19a (.009)
Dead Brch.
Dead Brch.
Foliage
C
W
.478a (.037) .021a (.002) .040a (.004)
.218b (.024) .010b (.001) .007b (.003)
.28a (.021)
.22b (.017)
C .912a (.014) .100a (.002) .380b (.026) .27a (.025)
Foliage W .916a (.021) .084b (.003) .494a (.029) .27a (.014)
______________________________________________________________________________
Note:
*
Means of each component followed by different letters are significantly different at the
95% confidence level.
#
Standard errors.

38
829
830
831
832
833
834
835
836
837
838
839
840
841
821
822
823
824
825
826
827
828
842
843
844
Table 11. Comparison of ecosystem components of the conventional (C) and whole-tree (W) harvest treatments 16 years after harvest of the 41-year-old stand.
______________________________________________________________________________
Treatment Component Biomass C N P K Ca
(Mg ha
-1
) ________________(kg ha
-1
)______________________
C Pine 176.8

88,375 578.3 41.1 158.5 315.8
W Pine 147.8 73,875 305.0 26.1 132.7 235.5
C Forest Floor 19.7 10,835 250.6 15.1 9.7 69.0
W Forest Floor 18.6 8,234 187.7 9.4 5.5 84.7
C Mineral Soil — 43,192

2,028.1 50.0 33.4 774.4
W Mineral Soil — 35,299 2,121.8 22.9 80.4 1,218.7
______________________________________________________________________________
Totals C 196.5 142,402 2,857.0 106.2 201.6 1,159.2
W 166.4 117,408 2,614.5 58.4 218.6 1,538.9
______________________________________________________________________________

Pine values include above and belowground biomass.

Soil values represent the upper 60 cm.

39
853
854
855
856
857
858
859
860
861
862
863
864
845
846
847
848
849
850
851
852
Table 12. Consolidated elemental capital of the conventional (C) and whole-tree (W) harvest treatment just after harvest of the 41-year-old stand and 16 years later.
______________________________________________________________________________
Treatment Sampling Date C N P K Ca
___________________(kg ha
-1
)_______________________
C 1980 73,199 2,343.0 35.7 173.7 674.9
C 1995 142,402 2,857.0 106.2 201.6 1,159.2
______________________________________________________________________________
Difference 69,203 514.0 70.5 27.9 484.3
W 1980 76,813 2,564.8 40.8 272.7 1,542.0
W 1995 117,408 2,614.5 58.4 218.6 1,538.9
______________________________________________________________________________
Difference 40,595 49.7 17.6 - 54.1 - 3.1

40
873
874
875
876
877
878
879
880
881
882
883
884
865
866
867
868
869
870
871
872
Table 13. Carbon content of the mineral soil in a 16-year-old naturally regenerated loblolly pine stand in the Piedmont of South Carolina following conventional and whole-tree harvesting.
______________________________________________________________________________
Depth
(cm)
Conventional Harvest
(Mg ha
-1
) (%)
Whole-tree Harvest
(Mg ha
-1
) (%)
______________________________________________________________________________
0-8
8-34
10.9a
*
20.0a
25
46
9.0a
16.4a
26
46
34-60 12.3a 29 9.8a 28
______________________________________________________________________________
Total 43.2 100 35.2 100
______________________________________________________________________________
Note:
*
Means within rows followed by different letters are significantly different at the 95% confidence level.