2007 SOUTHEASTERN NATURALIST 6(3):523–534
Influences on Oak and Pine Establishment with Time Since
Fire in Sandhills Pinus paulustris (Longleaf Pine) Forests
Jaclyn L. Schnurr1,* and Beverly S. Collins2
Abstract - We investigated biotic and abiotic mechanisms that affect Quercus
falcata (southern red oak) establishment from seeds in southeastern Pinus
paulustris (longleaf pine) forests along a time-since-fire chronosequence (unburned
since 1950, the mid-1990s, or winter 2001) and slope gradient. To determine if seed
availability limits southern red oak recruitment, we measured seed production,
populations of small mammals, and seed removal across sites. To determine if
seedling survival limits establishment, we outplanted greenhouse-raised southern
red oak seedlings and followed their survival throughout one growing season. We
also measured abiotic conditions such as canopy openness, soil moisture, and soil
temperature, and censused longleaf pine recruitment. Contrary to expectations
based on species’ fire-tolerances, longleaf pine recruitment was consistent over the
chronosequence, but there was little oak seedling establishment in the long-unburned
sites. Sites last burned in the mid-1990s had the lowest oak seed rain,
highest small-mammal populations, and highest seed -removal rates; yet, they had
the highest oak seedling establishment. Seedling survival was highest in the pre-
1950 sites, and was most influenced by intermediate light levels and high soil
moisture. We conclude that biotic factors, specifically, seed removal by small
mammals, are less important to seedling recruitment than abiotic factors in these
forests. Further, the assumption that longleaf pine forests will eventually undergo
to succession to hardwoods in the absence of fire is complicated by abiotic conditions
associated with landscape position.
In the Piedmont-Coastal Plain fall-line region of the southeastern US,
upland Pinus palustris P. Mill. (longleaf pine) forests grade from sandhills
longleaf-scrub oak communities on ridgetops through mixed pine-hardwood
forests on slopes (Christensen 1988, Collins et al. 2006, Jacqmain et al.
1999, Skeen et al. 1993, Ware et al. 1993). Upland oak species that codominate
with longleaf pine in these forests are distributed from sandhills
species, such as Quercus margaretta Ashe (sand post oak) and Quercus
laevis Walt. (turkey oak), which regenerate primarily by sprouting, through
more mesic species such as Quercus falcata Michx. (southern red oak), the
focus of this study, Quercus nigra L. (water oak), and Quercus alba L.
(white oak), which tend to regenerate from seeds (Collins et al. 2006).
Vegetation composition and regeneration of canopy species in upland
longleaf forests has been shown to be related to both topography and fire
regime. Decreased frequency of natural or prescribed fires has been linked to
1Wells College, Aurora, NY 13026. 2Western Carolina University, Cullowhee, NC
28723. *Corresponding author - firstname.lastname@example.org.
524 Southeastern Naturalist Vol. 6, No. 3
increased recruitment of hardwoods, including oaks (Brockway and Lewis
1997; Hartnett and Krofta 1989; Heyward 1939; Jones et al. 1981, 1984;
Monk 1965, 1968; Quarterman and Keever 1962; Shelton and Cain 1999;
Walters 1991). Lower light levels in infrequently burned forests can favor
regeneration of more shade tolerant hardwoods over less shade tolerant
longleaf pine (Gardiner and Hodges 1998, Larsen and Johnson 1998, Li and
Ma 2003, Phares 1971), and the lack of fire will allow the hardwood
seedlings to persist.
Although birds and small mammals can have large effects on tree seedling
recruitment (Elkinton et al. 1996; Hulme and Hunt 1999; Janzen 1970;
Jensen 1985; LoGiudice and Ostfeld 2002; McCracken et al.1999; Mosandl
and Kleinert 1998; Nilsson 1985; Ostfeld et al. 1996, 1997; Schnurr et al.
2002, 2004; Schupp 1988; Sork et al. 1993; Whelan et al. 1991; Wolff 1996),
the effects of animals on regeneration have been relatively unstudied in
longleaf pine forests of the fall-line sandhills region. Seed production and
input are the first biotic filters (Clark et al. 1998, Houle 1998, Schnurr et al.
2004, Shibata and Nakashizuka 1995) to regeneration, but even if a seed
arrives at a site, it must escape seed predators (Hulme and Hunt 1999;
LoGiudice and Ostfeld 2002; Ostfeld et al. 1997; Schnurr et al. 2002, 2004;
Whelan et al. 1991). Oaks, for example, are subject to small-mammal and
insect effects on seed viability, dispersal, and longevity, and these influences
may interact with environmental drivers and/or forest management
practices, including time since fire, to determine the proportion of oak
species in the regeneration pool and, ultimately, the canopy of sandhills
longleaf pine forests.
The objective of our research was to determine the relative effects of
seed predation and abiotic conditions (canopy openness, soil moisture,
soil temperature) on regeneration of oaks, with a focus on southern red
oak, in upland longleaf pine stands that differ in forest management and
fire history (time since fire). These study species are dominant oak species
in upland pine and pine-hardwood forests on slopes in the fall-line
sandhills region (e.g., Collins et al. 2006, Dilustro et al. 2002, Workman
and McLeod 1990). They grade upslope into longleaf pine-scrub oak
communities and downslope into upland hardwoods (Collins et al. 2006,
Workman and McLeod 1990). To compare regeneration over the slope
gradient, we included longleaf pine-scrub oak and upland hardwoods
stands. We asked, does seed production limit recruitment of oaks, either
directly or through an increase in small-mammal seed predators? We
predicted sites with greater acorn production would have greater densities
of seed predators, specifically small mammals, and that these mammals
would cause increased seed predation and decreased seedling recruitment.
We used southern red oak to examine seed removal and seedling survival
over the topographic gradient. We also predicted that, in the absence of
biotic effects on seed and seedling survival, oak recruitment would increase
down the topographic gradient from sandhills to mesic hardwood
2007 J.L. Schnurr and B.S. Collins 525
forest, and with time since fire in the upland pine stands. We sampled
longleaf pine regeneration to determine if patterns of oak and pine regeneration
Site description and sampling
The Savannah River Site (SRS) is a 78,000-ha facility of the United
States Department of Energy, located on the upper coastal plain physiographic
province along the South Carolina/Georgia border. When the SRS
was acquired in 1950, the majority of land was forested (67%), while the rest
was cropland (33%), although most of the forests had been logged (Workman
and McLeod 1990). Starting in 1952, the US Forest Service (USFS)
planted pine (primarily Pinus taeda L. (loblolly pine), but also P. palustris
P. Mill. (longleaf pine), P. elliotii Engelm. (slash pine), and some crosses
among these) seedlings on abandoned sites throughout the SRS. Today,
forest communities grade from swamp forests and bottomland hardwoods at
lower elevations, through sloped upland hardwood forests, to upland managed
pine forests, to more xeric fire-adapted sandhill longleaf-scrub oak
communities at 80–130 m elevation (Workman and McLeod 1990). Pine
forests are currently managed using even- aged techniques, such as prescribed
fires and thinning (Kilgo and Blake 2005).
In spring of 2001, we established ten 1-ha sites over a topographic
gradient from upland hardwoods (HDW, N = 2) upslope through upland
longleaf pine (N = 6) to sandhills longleaf pine-scrub oak stands (SND,
N = 2). Composition and structure of the selected stands were representative
to those along topographic gradients on the SRS generally (Workman and
McLeod 1990) and to previously sampled stands in other sites along the
Piedmont-Coastal Plain fall-line region (e.g., Collins et al. 2006).
The upland longleaf pine forests differed in fire management and time
since fire; they were unburned since 1950 (B50, N = 2), the mid-1990s (B90,
N = 2), or winter 2001 (B01, N = 2). Our sampling areas were located within
a large homogeneous patch of each stand, with at least 30 m separating them
from a different habitat type; however, there was a potential for seeds and
animals to disperse into the sites from the surrounding variable habitat
matrix. Due to the labor-intensive nature of our sampling regime, we were
only able to sample two 1-hectare sites in each habitat.
To characterize the sites, understory vegetation was surveyed along a
5-m line-intercept transect at 20 locations within each site in summer of
2001. We included all herbaceous and woody vegetation less than 2 m tall,
and individuals were identified to species when possible. Species richness
and the Shannon-Weiner diversity index were calculated for the herbaceous
layer in each site. We determined percent cover for each species in each site,
and then averaged the values for the 2 sites per treatment.
526 Southeastern Naturalist Vol. 6, No. 3
Seed input into the sites was measured using 8 litter traps per site (0.37 m
x 0.5125 m x 0.70 m high) that were emptied monthly from August 2001
through August 2002. Although the seed traps were accessible to small
mammals, we found no seed fragments or small-mammal feces that usually
indicate that seeds had been removed. All collected seeds were sorted by
species, and live seeds (those with green embryos) were counted to estimate
total seed fall. Since seed fall of oak species was low, all Quercus sp. acorns
were summed for the analyses.
To determine the relationship between seed inputs and small-mammal
populations, trapping was conducted using large, folding Sherman live traps
located at 12.5-m intervals throughout each hectare. We trapped each grid
for 3 days in August and November 2001, and February and May 2002. After
a 3-day prebaiting period, each trap was set using black-oil sunflower seeds
and was checked each morning at 0700 h and each evening at 1700 h.
Captured small mammals were identified to species, sexed, weighed, eartagged,
and released at the point of capture. Due to extremely low capture
rates, traps remained open for the entire 3-day trapping period.
Southern red oak seed and seedling survival
We quantified seed removal rates experimentally by placing southern red
oak acorns directly on the ground at 15 locations randomly chosen throughout
each site in December 2001. Acorns were collected in November 2001
from sites throughout the SRS, and only acorns that had no indication of
insect infestation were used. We used 2 acorns per location: locations were
marked using fiberglass flags and the acorns were located 50 cm on either
side of the flag and marked with a stick found in the area. Acorn removal
was monitored daily for 3 days, then monthly for a total of 60 days of
exposure. Gloves were worn when handling acorns and twigs to decrease
olfactory cues to potential seed predators.
To assess seedling survival, the southern red oak acorns collected in fall
2001 were stored at 5 °C in a coldroom until early February 2002, when
seeds were germinated in the greenhouse. Successful germinants were
planted in homogenized soil collected from all sites. Seedlings were kept in
the greenhouse until danger of frost passed, and planted in the field in April
2002. Five seedlings were planted in a 1 x 1 m quadrat at six locations
randomly chosen throughout each site, for a total of 30 seedlings per site.
To assess the environment, soil gravimetric water content was measured
at each seedling transplant quadrat in April, May, July, and October 2002.
Canopy photos were taken 1 m above each quadrat in July 2002 and analyzed
using GLA software (Frazer and Canham 1999). Soil temperatures
(± 1 °C) were recorded at 2 locations within each site using iButtons®
(Dallas Semiconductor Thermochron, Sunnyvale, CA) buried at 10 cm. Data
collection began in October 2001 and continued until July 2002. Soil temperature
was logged every 1.5 hours, but data were averaged monthly.
Seedling survival was monitored until October 2002.
2007 J.L. Schnurr and B.S. Collins 527
Data were analyzed using SYSTAT v. 10 and SAS v. 9. Analyses of
variance ( = 0.05) were used to determine if mean seedling cover, seed
inputs, or seed removal differed among sites, and Bonferroni-adjusted t-tests
were used to examine pairwise differences between sites. Logistic analysis
(SAS, Proc Catmod) was used to examine the relative effect of site on
southern red oak seedling survival, i.e., to test the hypothesis that survival in
a site did not differ from mean survival averaged over all sites.
The sites that were most recently burned in the winter of 2001 (B01) had
the highest species richness of the herbaceous layer, while the sites burned in
the mid-1990s (B90) had the highest Shannon-Weiner diversity indices
(Table 1). The dominant species in the herbaceous layer varied among sites,
and usually made up about 30% of the total species composition (Table 1).
The dominant species among the burned sites varied from Andropogon spp.
(bluestem grass) in the most recently burned sites through the lichen
Cladonia leporina in the sites burned pre-1950 (Table 1). The sites most
recently burned had the largest portion of the ground covered by plants, with
the B90 sites having 55% coverage and B01 having 52%. The hardwood
sites (HDW) (14%) and sites burned pre-1950 (B50) (19%) had the least
ground covered, although the B50 sites also had a thick duff layer, and the
sandhills (SND) sites had an intermediate level (40%).
Pooled over species, oak seedling percent cover was greatest in the B90
sites compared with all other sites (F = 8.856, P < 0.001; Fig. 1A). Longleaf
pine seedling cover did not differ significantly among sites, but the B50 sites
had the greatest longleaf recruitment, while the HWD sites had none (F =
1.818, P = 0.16; Fig. 1A).
Maximum seed rain occurred in November 2001, when 29 of the 60 total
seeds were collected. Seed rain did not correspond with the seedling cover
patterns among sites (2 = 0.342 df = 1, P = 0.559; Fig. 1B). Oak seed rain
did not differ significantly among sites and was lacking or low in all sites
except HDW sites. B90 sites, which had the greatest oak seedling cover, had
no oak seed rain (Fig.1B). Longleaf pine seed rain did not significantly vary
among sites (Fig.1B).
Table 1. Percent cover of the most abundant herbaceous-layer species, total species richness,
and Shannon-Weiner diversity index value for each stand type.
Stand herbaceous Species Shannontype
Most abundant species % cover % cover richness Weiner H'
SND Vaccinium stamineum L., deerberry 31 40 30 2.19
B01 Andropogon spp., bluestem 37 52 31 2.07
B90 Gelsemium sempervirens (L.) St. Hil., 36 55 28 2.35
evening trumpet flower
B50 Cladonia leporina, lichen 32 19 22 2.14
HDW Vitis rotundifolia Michx., muscadine grape 30 14 13 2.04
528 Southeastern Naturalist Vol. 6, No. 3
Contrary to our prediction, small-mammal capture patterns did not correspond
to seed rain (Fig. 1C). Total number of animals captured were: SND = 0,
B01 = 10, B90 = 23, B50 = 6, and HDW = 6. The majority of small mammals
2007 J.L. Schnurr and B.S. Collins 529
were captured at the B90 sites (Fig. 1C), which had no oak seed rain. There
were no animals captured in the SND sites throughout the study (Fig. 1C). In
the other sites, Peromyscus gossypinus LeConte (cotton mouse) was the
most abundant animal captured (89%), followed by Ochrotomys nuttalli
Harlan (golden mouse; 8%) and Reithrodontomys humilis Audobon and
Bachman (eastern harvest mouse; 2%).
Removal of southern red oak acorns differed among sites (F = 5.814, P =
0.04; Fig. 1D), but pair-wise comparisons were not significant. B90 sites,
which had the highest acorn removal, also had the greatest number of small
mammals captured, but not the most seed rain, and there were no acorns
produced in these sites. Although some acorns were removed in the SND
sites, no small mammals were captured there.
Survival of southern red oak seedlings differed among the sites (2 =
32.96, P < 0.001; Table 2). Survival was greatest in the sites last burned
in the 1950s (B50; 2 = 22.75, P < 0.001), and lowest in sites last
burned in 2001 (B01; 2 = 22.75, P < 0.001). Seedling survival was most
variable within the HDW sites (Table 2).
Seedling survival patterns reflected environmental differences among
sites (Table 2). Specifically, soil moisture (April, May, July, and October) was
significantly different among sites, with B50 and HDW sites having consistently
higher soil moisture than the other sites (April: F = 5.986, P < 0.001;
May: F = 2.82, P = 0.034; July: F = 18.298, P < 0.001; Oct.: F = 2.559, P less than
0.049; Table 2). Canopy openness also was significantly different among sites
Table 2. Mean Quercus falcata (southern red oak) seedling survival, canopy openness (%), soil
moisture (April, May, July, and October; %) and soil temperature (April–July, °C) in southeastern
forests spanning a topographic gradient (mesic hardwoods [HDW] through upland pine
forests [B] and dry sandhills [SND]) and time-since-fire (last burned prior to 1950 [B50], in the
mid-1990s [B90], or in 2001 [B01]). Values are means ± standard error, N = 2.
Seedling Canopy Soil Soil
survival openness moisture temperature
Stand type (% ± SE) (% ± SE) (% ± SE) (°C ± SE)
SND 48 ± 8.7 30.9 ± 2.40 2.1 ± 0.14 22.9 ± 2.5
B01 20 ± 9.5* 28.2 ± 0.67 1.6 ± 0.17 23.4 ± 2.6
B90 65 ± 7.0 18.0 ± 10.80 2.3 ± 0.17 21.5 ± 2.4
B50 75 ± 9.5* 11.4 ± 0.66 3.6 ± 0.25 21.2 ± 2.3
HDW 50 ± 10.5 9.8 ± 0.16 2.9 ± 0.15 21.0 ± 2.2
* = significantly higher or lower survival.
Figure 1 (opposite page). A) Average cover of oaks (Quercus falcata, Q. laevis, and Q.
nigra) and longleaf pine (Pinus palustris) in m per each 5-m transect in five upland
forests. B) Total number of viable acorns (Q. falcata, Q. laevis, Q. nigra) and Pinus
palustris seeds collected in seed traps from September 2001–January 2002. C) Average
number of small-mammal seed predators captured from August 2001 through May
2002. D) Total percent removal of acorns at each site after the entire 60-day exposure
period. Different letters indicate significant differences based on Bonferroni pair-wise
comparisons at = 0.05. Error bars are standard deviation, N = 2.
530 Southeastern Naturalist Vol. 6, No. 3
(F = 60.518, P < 0.001), with B50 and HDW sites having the most shade in
July (Table 2). Soil temperatures mirrored the canopy-openness data, with the
B50 and HDW sites having the lowest average soil temperatures.
We found that longleaf pine recruitment was consistent across sites
differing in slope position and fire history, while recruitment of oak species
was greatest in sites with an intermediate time-since-fire (B90). Contrary to
our expectations, the sites with the most oak seed rain (HDW) did not have
the highest numbers of small-mammal seed predators, and seed removal was
highest in sites with few small-mammal captures and low seed rain (B90).
Southern red oak seedling survival was influenced by the abiotic site conditions
found along the chronosequence, with the highest survival in the sites
with the longest time-since-fire (B50) and lowest in the most recently
burned sites (B01), corresponding with more favorable light and soil-moisture
levels (i.e., summer shade and higher summer soil moisture).
Ground-layer coverage ranged from 14% in the mesic hardwood stands
to over 50% in the most recently burned stands. Similarly, species richness
was greatest in the sites that had been recently burned along with the
sandhills sites. As expected, there was no measurable longleaf seedling
cover in the upland slope hardwood (HDW) stands. However, longleaf
seed rain was measurable over all sites, with more longleaf seeds than
acorns falling into the seed traps. Seed production by oaks in our sites was
relatively low, suggesting our study may have been conducted in a nonmast
year. Although much is known about masting in northern oak species
(Elkinton et al. 1996, Greenberg and Parresol 2000, McCracken et al. 1999,
Ostfeld et al. 1996, Schnurr et al. 2002, Wolff 1996), little is known about
southern oaks. More research on the topic is necessary.
The presence of pine and oak seedlings in all the upland longleaf pine
stands suggests that time since fire does not result in environmental or biotic
conditions that would limit regeneration. Longleaf seedling cover in recently
burned stands and in those that have not been burned since before the SRS
was created does not agree with results from other sites, which show that
longleaf regeneration declines with time since fire in upland forests in the
southeastern US (Hartnett and Krofta 1989). The relatively high cover of
oak seedlings with no recorded oak seed rain, and highest amounts of seed
removal in sites last burned in the 1990s, suggests that seedlings may have
accumulated from prior mast years or that seeds have been dispersed into the
stands by jays or woodpeckers.
Our results indicate that a complex mixture of abiotic site conditions
and biotic factors determine oak, specifically southern red oak, recruitment
with time-since-fire. Close agreement between short-term seedling-survival
patterns and environmental conditions related to evapotranspiration,
with higher survival in the cooler, wetter, and more shaded long-unburned
stands and lower survival in the hotter, drier, and more open recently
2007 J.L. Schnurr and B.S. Collins 531
burned stands, agrees with the abiotic controls on oak regeneration found
by other researchers (Garcia et al. 2002, Jacqmain et al. 1999, Jones at el.
1994, Monk 1965, Schumann et al. 2003) and suggests the potential for
oaks to increase in the absence of fire. However, we found that seed
production or survival could potentially limit oak recruitment after fire, at
least in years of low acorn production. Acorn abundance was low in all the
upland longleaf pine stands, and was not measurable in the stands burned
in 2001 and the 1990s. Over 50% of acorns were removed from all sites,
with stands burned in the 1990s (where the largest numbers of small
mammals were captured) having 98% acorn removal. Although high seed
rain during mast years may lead to episodic recruitment, the lack of seedling
cover in the stands last burned before 1950 suggests low seed rain, and
low, but consistent, rates of seedling removal, along with long-term seedling
mortality may offset short-term seedling survival and lead to low oak
regeneration in long-unburned sites.
Overall, results of our research suggest environmental conditions, including
light, associated with canopy type and soil moisture differences
related to topography have a greater influence than time since fire on
southern red oak and longleaf pine regeneration in the upland forests we
sampled. Lack of longleaf seedling cover despite seed production and low
populations of small-mammal seed predators in the hardwood stands, where
light levels were at 10% open sky and were the lowest among the sampled
stands, suggests that shade may limit longleaf regeneration. Although light
levels decreased among the upland longleaf sites with time since fire, levels
only approached those of the hardwoods forest in the longest unburned
(B50) sites. Low southern red oak seedling survival in the most recently
burned sites and the sandhills sites, where the canopy was the most open and
soil moisture was lowest in July, suggests that summer drought may limit
regeneration of this species. Although where conditions are more mesic, as
in the hardwood and longer-unburned upland longleaf sites, biotic factors
may have a relatively stronger influence on oak regeneration, relatively
consistent acorn production (J.L. Schnurr, pers. observ.) and low smallmammal
populations (Langley and Shure 1980; Loeb 1999; B. Reinhart,
Savannah River Ecology Lab, Aiken, SC, pers. comm.; Smith et al. 1984.)
may combine to produce low overall biotic effects. Results of our research
did not support our prediction that sites with greater acorn production would
have greater densities of seed predators, increased seed predation, and decreased
seedling recruitment of southern red oak. However, as predicted,
southern red oak recruitment did increase down the topographic gradient
from sandhills to mesic hardwood forest, and with time since fire in the
upland pine stands, mainly in response to soil moisture and light availability.
Field assistance was provided by Brian Hough, Chris Crawford, Nate Huber-
Fliflet, Pamela Weisenhorn, Matt Opdike, and John Dilustro. Brian Hough provided
532 Southeastern Naturalist Vol. 6, No. 3
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