Geomorphology 126 (2011) 108–120

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Geomorphology
j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / g e o m o r p h

Wood distribution along streams draining old-growth floodplain forests in Congaree

National Park, South Carolina, USA
Ellen Wohl ⁎, Lina E. Polvi, Daniel Cadol 1
Department of Geosciences, Colorado State University, Fort Collins, CO 80523-1482, USA

a r t i c l e i n f o a b s t r a c t

Article history: We measured wood volume and other characteristics along a series of instream and floodplain transects in
Received 24 August 2010 and around Congaree National Park, South Carolina. Seven transects were surveyed at most sites; one within
Received in revised form 25 October 2010 the bankfull channel, and three on each bank, at 10, 50, and 100 m lateral distances, respectively, from the
Accepted 26 October 2010
channel. Along each transect we measured piece dimensions, location along transect, orientation, and decay
Available online 9 November 2010
category. We also characterized channel width, drainage area, vegetation type, average forest basal area, and
Keywords:
inundation category for each transect. Forest transects N 100 m from a channel were also surveyed. Instream
Large woody debris wood loads for unmanaged channels draining old-growth forests were 102–178 m3/ha, which is lower than
Floodplain most instream wood loads documented for old-growth temperate forests. We attribute the lower wood loads
Subtropical to the combined effects of high transport capacity and rapid decay. Floodplain wood loads are lowest close to
Blackwater swamp the channel but increase with distance, reaching comparable values (up to 172 m3/ha) at distances N 100 m
Wood load from a channel. Channel geometry does not appear to exert the significant influence on instream wood loads
Logjams that has been documented for more laterally constrained channels. Channel-spanning jams are uncommon
Congaree
and appear to be transient features.
Coastal plain
© 2010 Elsevier B.V. All rights reserved.

1. Introduction
An extensive literature now documents the geomorphic and
ecological roles of wood in streams of varying size (Gurnell et al.,
2002; Hassan et al., 2005). Studies of the characteristics and function
of instream wood began in the U.S. Pacific Northwest during the late
1970s (e.g., Keller and Tally, 1979; Bilby and Likens, 1980), and for
more than two decades the majority of research continued to be
conducted in this region. Within the past decade, investigations of
instream wood have expanded to a wider variety of temperate and
tropical environments on several continents, but a conspicuous gap
remains in the lack of studies from the subtropical coastal plain
environments of the southeastern United States. Instream wood in
this region might reflect forest and stream characteristics intermedi-
ate between those documented for tropical (Cadol et al., 2009; Mao
and Comiti, 2009; Wohl et al., 2009) and temperate sites, although
investigations at numerous field sites will be necessary to thoroughly
evaluate this possibility.
Apart from potential regional differences, wood characteristics in
coastal plain streams are of interest because relatively few studies
have systematically compared instream and floodplain wood dis-
tributions (Young et al., 2006), despite the importance of exchanges of
⁎ Corresponding author. Tel.: + 1 970 491 5298; fax: + 1 970 491 6307.
E-mail address: ellenw@cnr.colostate.edu (E. Wohl).
1
Present address: Department of Geology, College of William and Mary, P.O.
Box 8795, Williamsburg, VA, 23187-8795, USA.
water and sediment between the channel and floodplains facilitated
by instream wood (Harwood and Brown, 1993; Piégay and Marston,
1998; Abbe and Montgomery, 2003; Jeffries et al., 2003; O'Connor
et al., 2003; Sear et al., 2010). A large proportion of instream wood
studies come from streams that are moderately to strongly laterally
confined. Streams with extensive overbank flooding likely have
different patterns and processes of wood exchange between channels
and floodplains, which could produce different values of instream
wood load and mobility. We chose streams in Congaree National Park,
South Carolina, which contains the largest remaining old-growth
bottomland forest in the southeastern U.S. (Zhao et al., 2006), to
investigate these issues. We had three specific objectives: (i) Evaluate
the relative importance of differing controls on instream and
floodplain wood loads in this field setting, which contrasts with
steep, narrow valleys in which characteristics such as channel
gradient and valley geometry can strongly influence wood load (Fox
and Bolton, 2007; Wohl and Jaeger, 2009; Wohl and Cadol, 2011). We
hypothesize that instream and floodplain wood loads correlate
strongly with forest characteristics such as basal area, rather than
with channel characteristics such as channel width, as in more
laterally confined channels. (ii) Compare wood characteristics (spatial
distribution, orientation, piece size, wood volume/channel, or flood-
plain area) between channel and floodplain environments in order to
evaluate how the extensive floodplain and frequent overbank flood-
ing of this site influence wood dynamics in and adjacent to streams.
We hypothesize that instream wood is more aggregated and aligned
relative to flow direction than floodplain wood and that instream

0169-555X/$ – see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.geomorph.2010.10.035
 E. Wohl et al. / Geomorphology 126 (2011) 108–120 109

wood loads are comparable to floodplain wood loads because of the
potential for wood transport across the floodplain during overbank
floods. (iii) Compare instream wood loads in this environment to those
in old-growth tropical and temperate forests. We hypothesize that
relatively fast wood decay and high transport capacity result in lower
wood loads in this subtropical environment than those documented for
old-growth temperate forests.
2. Study area
Congaree National Park (http://www.nps.gov/cong/) centers on an
8991-ha portion of the left bank floodplain of the Congaree River and
includes 4452 ha of old-growth forest in which the oldest trees are
N200 years old. The oldest loblolly pines (Pinus taeda) are N247 years
old and the oldest bald cypress (Taxodium distichum) are 700–
1000 years old. The river flows E–SE to its junction with the Wateree
River. Drainage area upstream from the park is 18,100 km2 and flow is
regulated. Mean monthly temperature ranges from 7.7 to 26.9 °C
(Doyle, 2009). Mean annual precipitation in Congaree National Park is
1220 mm (Zhao et al., 2006), with between 60 and 138 mm falling
each month on average (Doyle, 2009). Flooding is most frequent
during winter and early spring, but can occur at any time during the
year. Field work for this study included the first flood of the winter of
2009–10. Tropical storm Ida dropped several centimeters of rain in
the region during 10–12 November 2009, which resulted in complete
inundation of the floodplain within the national park for a few days and
made it possible to kayak over the measurement sites. Mean annual
discharge for the Congaree River during 1940–2007 is 254 m3/s (mean
annual range 98–469 m3/s) (Meitzen, 2009). Average bankfull stage
occurs at 4.5 m above the streambed, which equates to ~850 m3/s. The
average channel gradient is 0.0001 m/m, and sinuosity is 1.9 in the
vicinity of the national park. Confined to the north by uplands and
~6.5 km wide, the floodplain has a mosaic of fluvial landforms, including
natural levees, point bars, meander scroll ridge and swale complexes,
backwater swamps, and abandoned meanders (Shelley, 2007; Meitzen,
2009).
Cedar Creek and Toms Creek cross the floodplain as blackwater
tributaries to the Congaree River within the national park (Fig. 1).
Cedar Creek drains ~270 km2, and Toms drains ~ 130 km2. Although
these represent relatively large drainage areas, they are the smallest
creeks that flow through old-growth forest; the other channels in
Fig. 1 are all secondary channels rather than tributaries. Both Cedar
and Toms Creeks are sinuous sand-bed channels with pool-riffle or
dune-ripple morphology (Montgomery and Buffington, 1997) and
numerous secondary channels and sloughs. Selected pieces of
instream wood are removed once a year from the upstream portion
of Cedar Creek (sites Cedar 1, 2, and 3 in this study) by the National
Park Service in order to maintain a canoe trail.
Floodplain soils are predominantly poorly drained silty clay loams of
low permeability (Zhao et al., 2006). Tree species are distributed along

81°W

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6 km

Fig. 1. Location map of study reaches. Inset map shows the State of South Carolina. Dot at center of state is Columbia, the capitol. Paler gray squares are study areas; two sites along
the Lynches River, one along Congaree Creek in the city of Columbia, four at Beidler Forest south of the Santee River, and the remainder in Congaree National Park. The larger map
shows the location of sites (asterisks) in Congaree National Park, along the Congaree River, Cedar and Toms Creeks, and forest transects away from channelized flow. Regional flow is
from upper left to lower right. Primary channels are shown in thicker lines; the network of secondary channels and sloughs shown here is greatly simplified and most of these
channels are not longitudinally connected during periods of low flow.
110 E. Wohl et al. / Geomorphology 126 (2011) 108–120

topographic and hydrologic gradients (Sharitz et al., 1986). Common
species include bald cypress (Taxodium distichum), water tupelo
(Nyssa aquatic), loblolly pine (Pinus taeda), sweetgum (Liquidambar
styraciflua), green ash (Fraxinus pennsylvanica), and red maple (Acer
rubrum). Basal area of old-growth forests ranges from 24–54 m2/ha in
bottomland hardwoods to 56–75 m2/ha in sloughs (Allen and Sharitz,
1999), and diameter at breast height of the largest trees is N80 cm.
In addition to the sites in Congaree National Park, we also surveyed
transects along the bank of the Lynches River and Congaree Creek and in
the Beidler Forest Preserve of the National Audubon Society. The Lynches
River at the study sites drains 1875 km2. The channel is ~20 m wide,
and flow is unregulated. Forests along stream bank sections in the Lee
State Natural Area (http://www.southcarolinaparks.com/park-finder/
state-park/891.aspx) and a Nature Conservancy preserve, although not
cut within the past few decades, do not qualify as old-growth, which
is N200 years old (Devall, 1998). Congaree Creek drains about 350 km2 at
the study area in the Congaree Creek Heritage Preserve (https://www.dnr.
sc.gov/mlands/managedland?p_id=30) near the city of Columbia, SC. The
channel is 11 m wide, and flow is unregulated. As with the Lynches River
sites, the riparian forest along the creek does not qualify as old-growth,
although it has not been cut within the past few decades. The Beidler
Forest (http://sc.audubon.org/Centers_FBF.html) is a blackwater
swamp with limited development of channels in the vicinity of the
study sites and inundation via rising groundwater rather than out-of-
channel flooding.
3. Methods
3.1. Field methods
We chose study sites to provide diversity of forest age and
composition, drainage area, and channel–floodplain interactions
(Table 1). We distinguished four vegetation types based on vegetation
mapping and communities delineated in Congaree National Park by
the National Park Service: water tupelo dominant, mixed bottomland
hardwoods, riverbank and levee forests, and clearcut 20–30 years ago.
The forest sites at Congaree National Park and the sites at Beidler
Forest represent old-growth floodplain forests hundreds of meters
away from any channel.
At each site, we surveyed transects parallel to flow and to one
another. Transect length varied in relation to channel width; 100-m-
long transects for average channel width ≤15 m, 200-m-long transects
for channel width ≤20 m, and 500-m-long transects for channel width
≤100 m. Because of their large channels and turbid water, instream
wood was not surveyed along the Lynches and Congaree Rivers. At all
of the other stream sites, one transect included all wood within
the bankfull channel. Bankfull was defined by a pronounced break in
slope that was well developed on all the channels and differentiated
channelized from overbank flows. On each side of the channel,
additional transects were measured at 0–10, 50–60, and 100–110 m
from the bankfull edge. In other words, a full measurement set at a
stream site included seven transects: one within the channel and three
on each bank, all parallel to one another and to the predominant
direction of flow in the channel. We measured single transects at each of
the six forest sites. Within the channel transects, we measured the
length within the bankfull channel, diameter, and center point (with
respect to distance along the transect) of each piece of wood ≥10 cm
wide and ≥1 m long. The orientation of each piece was measured with a
hand-held compass, as was the predominant direction of flow. Pieces
within the channel were categorized as being a bridge (both ends above
the bankfull channel), ramp (one end above the bankfull channel),
pinned (wedged beneath other wood), buried (partially buried in the
stream bed), or unattached. On transects outside the channel, we
measured piece length within a band 10-m wide and perpendicular to
the transect, as well as diameter, center point distance, and piece
orientation. Pieces on all transects were assigned to a decay category of 1
(leaves or needles still attached), 2 (at least some bark still attached, as
well as small branches), or 3 (bark and branches gone, wood partly soft
to the touch). Pieces touching two or more additional wood pieces were
categorized as being part of a jam. Basal area of the surrounding forest
was measured at the start and end of each transect using a gage (Panama
Angle gauge).
The endpoints of each transect were also located with a hand-held
GPS unit so that the transect could be assigned an inundation category
from the map developed by Kimberly Meitzen. This map relates
topography and measured flood depths across the floodplain to flood
frequency using an interactive GIS model of floodplain inundation and
discharges from the Congaree River at Columbia gage (#02169500)
and the US Army Corps of Engineers Hydrologic Engineer Center
Regime Prescription Tools (USACE HEC-RPT) (Graf and Meitzen, 2006;
Kupfer et al., 2010). The Cedar Creek gage (02169672) within
Congaree National Park was used to validate modeled water-surface
elevations within the floodplain. The inundation map for the floodplain
was classified into six categories that distinguished major geomorphic
features (e.g., channels, abandoned meanders, levees, and point bars)
without creating too many divisions to be mapped (Graf and Meitzen,
2006). Category 1 represents discharge up to ~280 m3/s on the Congaree,
which equates to no out-of-bank floodplain inundation. Category 2,
when discharge is ≤560 m3/s, includes hydrologic connectivity between
the mainstem, tributaries, and secondary channels, with no out-of-bank

Table 1
Characteristics of study sites.

Site Aa (km2) Wb (m) Vegetationc Transect types Floodd BAe (m2/ha) Wood loads2 (m3/ha)

Cedar 1 186 9.3 Mixed hardwoods* All 1–6 30.2 3–42

Cedar 2 210 14 Tupelo* All 1–6 30.9 23–111
Cedar 3 250 12 Tupelo* All 1–6 40.8 7–71
Cedar 4 270 13 Mixed hardwoods* All 1–5 30.2 12–131
Toms 1 110 10 Tupelo* All 1–6 26.2 1–102
Toms 2 120 8 Tupelo* All 1–6 44.1 14–121
Toms 3 130 10.4 Mixed hardwoods* All 1–3 38.2 54–178
Congaree forest 1 n/a n/a Mixed hardwoods* Forest only 5 30.4 132
Congaree forest 2 n/a n/a Mixed hardwoods* Forest only 3 34.9 147
Congaree River 18,100 77 Riverbank Forest only 6 23.0 54–71
Congaree Creek 350 11 Riverbank All 1–5 23.6 19–52
Lynches River 1 1,875 20 Riverbank Forest only 3–4 23.5 28–84
Lynches River 2 1,875 20 Riverbank/mixed hardwoods Forest only 3–4 23.2 27–54
Beidler Forest n/a n/a Tupelo* Forest only 4 44.2 53–172
a
A is drainage area.
b
W is bankfull channel width.
c
Asterisk indicates old-growth forest.
d
Flood indicates inundation category (see text for explanation of categories); range of values are for different transects.
e
BA is average basal area of adjacent forest at each study site.
 E. Wohl et al. / Geomorphology 126 (2011) 108–120
floodplain inundation. Category 3, with discharge ≤1120 m3/s, repre-
sents minimal overbank flooding in low-lying areas. Category 4, with
discharge ≤1685 m3/s, represents the start of substantial overbank
flooding, with only the levees remaining dry. Category 5 (≤2245 m3/s)
and category 6 (≤2808 m3/s) represent progressively greater inundation
across the study area (K. Meitzen, University of South Carolina, personal
communication, 2010). Therefore, a transect categorized as category 1 has
the most frequent and deepest inundation, and a transect in category 6 is
least flooded.
3.2. Statistical analyses
Multiple linear regression was used to determine which channel and
floodplain characteristics explain the most variability in wood char-
acteristics within channel reaches and transects. Wood characteristics,
tested as response variables, are wood volume per area (m/ha), mean
length, and mean diameter. Predictor variables are drainage area,
channel width, average basal area, inundation category, and the
categorical variable of vegetation type. For wood in floodplain and
forest transects, distance from channel was included as a predictor
variable. An ANOVA was used to test differences in wood characteristics
for the four vegetation categories. For response variables with a
significant model from continuous variables and significantly different
vegetation categories, an overall model was created using dummy
variables. Unless specified otherwise, all statistical analyses were
performed using SAS Statistical analysis software Version 9.2 (SAS
Institute, 2002–2008). To reduce collinearity in models, Spearman rank
correlation coefficients were analyzed, and one variable was removed if
the absolute value of R exceeded 0.80. Models were chosen based on
Akaike's Information Criteria (AIC) (Akaike, 1973; Simonoff and Tsai,
1999) and adjusted coefficient of multiple determination (rs) values.
Orientation data were analyzed to determine whether wood in a
channel, floodplain transect, or forest transect followed a uniform
distribution. The Watson's U2 test (Mardia and Jupp, 2000), which
performs a goodness-of-fit test between the distribution of wood
orientations from the sample of interest and a uniform distribution,
was applied using the circular statistics program Oriana (Kovach
Computing Services, 1994–2010). The mean square deviation be-
tween the tested and uniform distributions is calculated, and a high
deviation results in a low probability and thus rejection of the null
hypothesis that the data fit a uniform distribution. Rose diagrams
were plotted for those distributions that were significantly different
than a normal distribution.
Following Kraft and Warren (2003), we used a one-dimensional
version of Ripley's K, a second-order statistic that evaluates the spatial
pattern of points within a landscape (Ripley, 1977), to assess the
uniformity of longitudinal distribution of wood along each transect.
Individual wood pieces were characterized as points distributed along
each one-dimensional transect. The K statistic is equivalent to the
average number of pieces within a given distance t of each piece,
noted K(t), where t is the length scale being considered. Individual
pieces were analyzed at length scales from 1 to 150 m, and jams were
analyzed at length scales from 1 to 25 m. To evaluate the significance
of aggregation (points closer to one another than random, i.e.,
clustered) and segregation (points farther from one another than
random, i.e., regularly spaced) found in our neighbor K analyses, we
compared the values of K(t) from each transect with values of K(t)
calculated from 100 simulated wood distributions known to be
spatially random. Because K(t) depends on the average spatial density
of pieces, the simulations were done using the same transect length
and the same number of pieces as the transect of concern. Observed
values of K(t) outside the envelope defined by the 5th and 95th
percentiles of the 100 simulations provide evidence that the pieces in
the study reach are not randomly distributed. Values of K(t) higher
than the envelope indicate aggregation at that scale; i.e., pieces are
clustered, causing more pieces within distance t than expected in a
111
random distribution. Values of K(t) lower than the envelope indicate
segregation at that scale; i.e., pieces are more evenly spaced than they
would be in a random distribution.
In calculating K(t), an edge effect must be considered. If a piece is
closer than distance t to the start or end of the transect, then there is a
chance that the number of pieces within distance t of it will be
undercounted, artificially lowering K(t). Therefore, in calculating K(t),
we did not include associated piece counts for those pieces that were
closer than t to the start or end of the transect, but we did continue to
consider those pieces in the associated piece counts for all other pieces.
Thus, we had larger sample sizes for smaller t values because, as t
increased, more pieces had to be excluded. In some cases this led
to sudden changes in K(t) with increasing t, as significant clusters
of pieces became too close to the edge and were excluded from
contributing to K(t).
With respect to the objectives and hypotheses in the introduction,
we used multiple linear regression and ANOVA to statistically evaluate
the effects of (i) forest characteristics by comparing wood loads in
relation to forest basal area and vegetation type and (ii) channel
characteristics by comparing wood loads in relation to drainage area
and channel width (first objective). We used Watson's U test and
Ripley's K to statistically evaluate differences in spatial distribution,
orientation, piece size, and wood load between channel and floodplain
transects (second objective). We qualitatively compared wood loads
reported in the literature for streams draining old-growth forests in
other environments to the wood loads that we measured at the
Congaree old-growth study sites (third objective).
4. Results
4.1. Controls on instream and floodplain wood loads
Table 2 summarizes the data for each surveyed transect. Multiple
linear regressions did not produce any significant models for wood
load in the channel transects. There was no significant effect of
vegetation category on the volume per area, mean length, or mean
diameter of the wood. Using the continuous variables (drainage area,
channel width, average basal area, and inundation category), no
significant model was found. However, when examining Spearman
rank correlation coefficients, average basal area is positively correlat-
ed to volume per area, mean length, and mean diameter, with rs
values of 0.25, 0.46, and 0.61, respectively. Drainage area is negatively
correlated to the three response variables, with rs values of − 0.40,
−0.45, and − 0.24, respectively.
For floodplain transects, no significant model was found for mean
length of wood. There is an effect of vegetation category on mean
diameter of wood; all other forest types are significantly different
than areas clearcut 20–30 years ago (Table 3). The only significant
multiple linear regression model was found for volume of wood per
unit area of floodplain. The ANOVA model found a significant effect of
vegetation category on volume per area; vegetation categories 1, 2,
and 3 were not significantly different, and categories 3 and 4 were
not significantly different. The multiple linear regression includes
the distance from the channel and the dummy variable of Veg2,
representing mixed bottomland hardwoods (Table 3). The results
thus partially support our hypothesis that wood load correlates more
strongly with forest characteristics (vegetation type, basal area) than
with channel geometry; forest characteristics influence floodplain
wood loads, although hydrologic characteristics (distance from the
channel) also exert an influence.
4.2. Wood characteristics in channel and floodplain environments
Wood loads differ significantly between channels and floodplains
in old-growth forests (Fig. 2A) and this difference appears to reflect
fluvial processes rather than forest characteristics. Comparing wood
112 E. Wohl et al. / Geomorphology 126 (2011) 108–120

Table 2
Results of transect surveys and K statistics analyses.

Reach Flooda Vegb BAc (m2/ha) # pieces Ld (cm) De (cm) Loadf (m3/ha) Transect Longitudinal patterng

Cedar 1 1 2 n/a 19 206.0 18.5 12.6 Channel 1–7 m random; 8–25 m agg
3 2 32.6 12 240.0 11.7 3.5 10 m rt bank Aggregated
4 2 24.8 8 342.5 19.4 42.1 50 m rt bank Random
5 4 30.4 20 318.0 19.4 26.7 100 m rt bank Random
3 2 23.6 18 293.9 19.0 21.0 10 m left bank Random
5 2 30.4 23 483.0 13.4 18.7 50 m left bank 1–12 m random; 13–25 m agg
6 4 39.4 49 433.5 13.1 33.5 100 m left bank Random
Cedar 2 1 1 n/a 26 355.4 19.2 42.9 channel Random
2 1 51.8 14 358.6 30.5 72.7 10 m rt bank Random
2 1 31.5 22 271.4 19.8 45.2 50 m rt bank 0–8 m agg; 9–25 m random
2 1 22.5 33 374.8 22.0 111.2 100 m rt bank Aggregated
2 1 20.2 14 335.0 22.4 23.3 10 m left bank Random
3 2 28.1 24 477.1 23.6 107.2 50 m left bank 1–7 m agg; 8–25 m random
5 2 31.5 27 306.3 24.8 75.0 100 m left bank 1–9 m agg; 10–25 m random
Cedar 3 (200 m transects) 1 1 n/a 88 295.3 24.5 71.6 Channel Random
3 1 50.2 23 301.3 31.7 43.1 10 m rt bank 1–26 m random; 27–50 m seg
3 1 55.6 81 305.9 16.5 41.6 50 m rt bank Aggregated
4 1 36.7 30 394.7 22.7 33.9 100 m rt bank Aggregated
3 1 n/a 28 202.1 14.9 7.1 10 m left bank 1–33 m agg; 34–50 m random
6 4 20.7 24 415.8 15.5 13.5 50 m left bank 1–35 m agg; 36–50 m random
Cedar 4 1 2 n/a 127 252.0 21.1 131.5 Channel 1–15 m agg; 16–25 m random
4 2 23.6 31 184.5 14.6 12.5 10 m rt bank 1–14 m agg; 15–25 m random
4 2 30.4 27 424.8 19.0 84.4 50 m rt bank 1–7 m random; 8–25 m agg
5 2 32.6 45 272.0 21.2 80.2 100 m rt bank 1–12 m random; 13–25 m seg
4 2 29.2 18 338.9 19.4 23.6 10 m left bank 1–4 m random; 5–25 m agg
4 2 37.1 41 423.7 21.9 91.4 50 m left bank 1–10 m random; 11–25 m agg
5 2 28.1 35 349.1 24.0 108.6 100 m left bank Random
Toms 1 1 1 n/a 66 401.5 18.6 102.2 Channel 1–15 m random; 16–25 m seg
3 1 29.2 30 347.0 21.1 49.8 10 m rt bank Aggregated
4 2 10.1 27 349.3 25.1 56.4 50 m rt bank 1–13 m agg; 14–25 m random
6 4 20.2 11 207.3 14.9 4.1 100 m rt bank Random
3 1 38.2 23 343.0 19.2 29.8 10 m left bank Random
4 2 30.4 37 387.0 18.5 62.5 50 m left bank Random
6 4 29.2 3 450.0 11.7 1.4 100 m left bank Insufficient no. logs to test
Toms 2 1 1 n/a 47 310.0 22.4 121.8 Channel 1–18 m random; 19–25 m seg
1 2 39.4 17 292.4 17.2 15.0 10 m rt bank Random
1 2 52.9 23 266.1 22.9 39.0 50 m rt bank 1–20 m random; 21–25 m seg
1 5 63.0 25 370.8 24.6 55.9 100 m rt bank Random
1 2 47.2 17 364.7 19.5 23.5 10 m left bank Random
1 2 45.0 21 476.7 20.6 58.5 50 m left bank Random
2 6 16.9 45 214.0 20.1 42.3 100 m left bank Random
Toms 3 1 2 n/a 50 372.4 24.8 178.5 Channel 1–19 m random; 20–25 m seg
3 2 38.2 13 420.0 28.7 54.3 10 m left bank Random
Congaree forest 5 2 30.4 46 353.0 20.5 131.9 Forest 1 Random
3 2 34.9 36 379.2 29.5 147.0 Forest 2 1–20 m random; 21–25 m seg
Congaree River (500 m transects) 6 3 19.6 215 257.7 20.1 54.3 10 m left bank Aggregated
6 3 23.2 192 377.1 17.7 54.9 50 m left bank Aggregated
6 3 26.3 147 432.2 21.3 71.2 100 m left bank 1–14 m agg; 15–100 m random
Congaree Creek 1 3 n/a 23 350.9 17.9 21.0 Channel Random
2 3 29.2 31 295.8 17.1 19.6 10 m rt bank Random
4 3 30.4 26 383.5 16.6 41.5 50 m rt bank Random
5 3 20.2 23 469.1 20.1 46.7 100 m rt bank 1–5 m random; 6–25 m agg
2 3 27.0 13 510.0 17.8 27.2 10 m left bank Random
4 3 14.6 34 314.7 20.1 52.0 50 m left bank Random
5 3 20.2 23 463.0 18.9 44.6 100 m left bank Random
Lynches 1 (200 m transects) 3 3 23.2 76 338.3 22.0 84.4 10 m rt bank Aggregated
3 3 27.0 42 309.5 20.5 27.8 50 m rt bank Random
4 3 20.2 44 372.3 19.4 29.6 100 m rt bank Random
Lynches 2 (200 m transects) 3 3 21.4 52 270.8 18.2 26.6 10 m left bank Random
4 2 24.6 55 272.0 18.7 35.5 50 m left bank 1–20 agg; 21–50 random
4 2 23.7 54 368.9 18.0 54.1 100 m left bank Random
Beidler 4 1 46.1 72 374.3 21.4 172.3 Forest 1 Random
4 1 36.0 67 363.7 21.7 155.1 Forest 2 Aggregated
4 1 51.8 50 314.2 18.4 53.6 Forest 3 Random
4 1 42.8 74 345.1 19.8 140.2 Forest 4 Aggregated
a
Flood = inundation category; 1 = greatest inundation, 6 = least inundation.
b
Veg = vegetation category; 1 = water tupelo dominant, 2 = mixed bottomland hardwoods, 3 = riverbank and levee forests, 4 = clearcut 20–30 years ago.
c
BA = average basal area of forest surrounding transect.
d
L = average length of pieces measured.
e
d = average diameter of pieces measured.
f
Load = wood load.
g
agg = aggregated; seg = segregated.
 E. Wohl et al. / Geomorphology 126 (2011) 108–120
Table 3
Results of ANOVA and multiple linear regression on channel and floodplain transects for wood load response variables.
Wood load response Vegetation category Multiple linear regression model
variable effect
Parameter Parameter estimate
Volume per area YES Distance from channel 0.01903
Mean length NO – –
Mean diameter YES – –
loads in relation to distance from the channel at the various sites
indicates the lowest wood loads immediately adjacent to the channel
(Fig. 2B). As might be expected, floodplain wood loads 50–100 m
distant from the channel are higher than instream wood loads along
managed channels, but lower than instream wood loads in unman-
aged channels. These results are not explained by differences in forest
basal area, which are statistically similar at transects 10, 50, and 100 m
from the channel (Fig. 2C). Wood loads in the forest transects N100 m
distant from a channel are comparable to those in unmanaged, old-
growth channels (Table 2).
With respect to piece dimensions, average instream piece length
and diameter do not correlate with either channel width or drainage
area. Average piece length differs little between channel (mean
318 cm) and floodplain (mean 349 cm) environments, as does
average piece diameter (channel mean 20.9 cm, floodplain mean
20 cm). Average piece length for floodplain transects differs little by
vegetation type (means 337, 344, 369, and 365 cm for vegetation
types 1–4, respectively). Average piece diameter for floodplain
transects varies little for vegetation categories 1–3 (means 21.4,
20.6, and 19.2 cm, respectively), but is lower (mean 14.9 cm) for
vegetation category 4, as might be expected given that this category
represents recent regrowth following clearcutting. This is supported
by the ANOVA testing for a significant effect of vegetation category on
mean diameter, which found that vegetation category 4 is different
than the three other categories. The multiple linear regressions did
not produce a significant model for piece length.
With respect to spacing of wood pieces along each transect, the
results from the K statistics are quite mixed (Table 2). Wood tends to
be randomly spaced along most of the channel transects, particularly
at lengths b20 m. The uncleared, old-growth channels (Cedar 4, Tom's
1–3) are more likely to have segregated wood at greater length scales
(Table 2). There is no apparent pattern of spacing along floodplain
transects as a function of distance from the channel; approximately
half of the transects display a random pattern. Spacing along
floodplain transects also shows no pattern with respect to inundation
or vegetation category or to basal area. Forest transects also fail to
display any consistent characteristics of piece spacing.
With respect to orientation of wood pieces, of the 70 channel,
floodplain, and forest transects that were surveyed, 26 transects had
distributions of wood orientations that were significantly different
from a uniform distribution (α = 0.1) (Table 4). Over half (5) of the
8 channel transects surveyed had nonuniform distributions of wood
orientation. Rose diagrams of the nonuniform distributions elucidate
differences between preferred orientation in the channel and
floodplain (Fig. 3). For example, in reach Cedar3, the wood in the
channel has a preferred orientation of NW–SE (which is also the
direction of channelized flow), and the nonuniform distributions on
the floodplain (transect 4, transect 5, and the entire floodplain) have a
preferred orientation of NE–SW.
The results thus partially support our second set of hypotheses.
There is no consistent difference in aggregation of wood between
instream and floodplain transects, contrary to our hypothesis,
although wood in channels is more likely to be preferentially aligned
than wood on the floodplain. Instream wood loads are significantly
higher than floodplain wood loads in unmanaged channels, although
113
Parameter Categorical variable Categorical parameter Model
p-value estimate p-value
b.0001 Veg2 22.047 b.0001
– – – –
– – – –
floodplain wood loads increase with distance from the channel,
suggesting a spatially limited process of concentrating wood in the
channel, perhaps during the falling limb of overbank floods.
4.3. Instream wood loads in old-growth forests
Table 5 summarizes the limited studies that document wood loads
for unmanaged streams in old-growth forests of diverse regions. The
mean value of wood volume per unit area of channel for the Congaree
study sites falls at the lower end of this range, with only the
subantarctic site from Tierra del Fuego, Argentina, having a lower
mean value, although the range of instream wood loads at Congaree
fall within the range of most other sites except those in the Pacific
Northwest temperate rainforests, which are much higher. Compar-
isons based on Table 5 are limited by the fact that the drainage areas at
the Congaree sites are much larger than the other drainage areas
except for the upper range from the Fox and Bolton (2007) study.
Multiple studies indicate that wood load in most regions is inversely
proportional to drainage area (Keller and Tally, 1979; Bilby and Ward,
1989; Marcus et al., 2002), but we are limited in our ability to hold
drainage area constant by the lack of published data for larger
drainages in old-growth from regions outside of Congaree, and the
lack of very small streams in old-growth forests at Congaree.
Evaluations of downstream trend in wood load typically use channel
width rather than drainage area (Gurnell, 2003; Wohl and Jaeger,
2009), however, because the ratio of wood piece length to channel
width is a useful indicator of relative transport capacity; pieces
shorter than channel bankfull width are more likely to be mobile
(Gurnell, 2003). Using the criteria of channel width, the sites along
Cedar and Toms Creeks are more comparable to other old-growth
sites. Therefore, the results appear to support our hypothesis of lower
wood loads in this subtropical old-growth site than in comparable
old-growth sites of the temperate zones.
5. Discussion
One of the most notable features of the South Carolina study sites
is the relative lack of large logjams, defined as channel-spanning jams
that elevate the local water-surface elevation and create substantial
backwater effects. This type of jam is not present even in unmanaged
channels draining old-growth forest sites. (Logjam here does not
connote cut pieces, but rather refers to accumulations of naturally
recruited and uncut wood.) Small instream logjams tend to form in
association with large ramp or bridge pieces recruited to the channel
via bank erosion or tree fall (Fig. 4A). Small jams also occur at the
mouths of secondary and abandoned channels (Fig. 4B), and very
small wood accumulations can be present on the upstream side of
trees growing on the floodplain. These features, along with the higher
wood loads in channels than on stream banks, indicate that wood
transport downstream and into the channel occurs in the study sites,
but wood does not concentrate sufficiently to form channel-spanning
jams that rise a meter or more above the stream bed and help to
increase stage during large flows. The removal of some instream wood
at Cedar Creek sites 1–3 could reduce the occurrence of channel-
spanning jams at these sites and at Cedar Creek site 4, which is
114 E. Wohl et al. / Geomorphology 126 (2011) 108–120

A Table 4
200 Results of tests of uniform distribution of wood orientation.a
180 a b a Reach Transect Significance level
160 Cedar1 Channel –
133
140 Trans2 –
Wood load (m3/ha)

133
Trans3 –
120 Trans4 –
Trans5 0.025
100 old-growth old-growth
uncleared 50-100 m Trans6 –
with no
80 old-growth from channel adjacent Trans7 –
channel
channel Floodplain –
60
Cedar2 Channel –
40 36 Trans2 –
Trans3 –
20 Trans4 –
0 Trans5 –
Trans6 –
Toms 1-3 Toms 1-3 Congaree forest Trans7 0.025
Cedar 4 Cedar 1-4 Beidler forest Floodplain –
Cedar3 Channel 0.1
Cedar 4 Trans2 –
uncleared Trans3 –
B Toms 1
Toms 2
old-growth
Trans4 0.01
200
Toms 3 Trans5 0.05
Congaree Cr
180 2nd growth Trans6 –
Cedar 1
Cedar 2 cleared Floodplain 0.01
160
Cedar 3 old-growth Cedar4 Channel 0.005
140 Trans2 –
Wood load (m3/ha)

Trans3 –
120
Trans4 0.025
100 Trans5 0.005
Trans6 0.05
80 Trans7 –
60 Floodplain 0.005
Lynches1 Trans1 0.005
40 Trans2 –
20 Trans3 0.01
Floodplain 0.1
0 Lynches2 Trans1 0.005
Trans2 –
0 20 40 60 80 100 120 Trans3 0.025
Floodplain 0.025
Distance from channel (m)
Toms1 Channel 0.005
Trans2 –
C Trans3 –
70
a a a Trans4 n/a
Trans5 0.1
60 Trans6 0.01
Average basal area (m2/ha)

Trans7 0.05
50 Floodplain 0.05
Toms2 Channel –
Trans2 –
40 Trans3 –
34 Trans4 –
32
30 30 Trans5 –
Trans6 –
Trans7 –
20
Floodplain –
Toms3 Channel 0.005
10 Floodplain –
Congaree Cr Channel 0.005
Trans1 –
0
10 50 100 Trans2 –
Trans3 –
Distance from channel (m)
Trans4 –
Trans5 –
Fig. 2. (A) Box plots of wood loads in old-growth forest sites for transects representing Trans6 –
unmanaged channels, floodplain transects within 100 m of a channel, and bottomland Floodplain –
transects ≫100 m from a channel. The horizontal line within each box indicates the median Beidler Forest TransA –
value, which is also written above the line. Box ends are the 25th and 75th percentiles, TransB –
whiskers are the 10th and 90th percentiles, and solid dots are outliers. Significant pairwise TransC 0.05
differences in means are indicated with contrasting letters above each box. (B) Scatter plots of TransD –
wood loads for selected channel and floodplain transects. ‘Cleared’ and ‘uncleared’ refer to Forest 0.1
channels from which wood is selectively removed and those which are not managed,
a
respectively. A dashed line between transects on Cedar 4 is provided for reference. Congaree Significance level shows p-values, where low p-values lead to rejection of null
Creek, a site with second-growth forest, is included for comparison with the old-growth sites. hypothesis that the orientations of the wood are in a uniform distribution. The test
Multiple symbols for floodplain transects at a given site represent right- and left-bank cannot be conducted with small samples sizes (n/a).
transects. (C) Box plots of average basal area versus distance from channel for sites in Fig. 2B.
As in Fig. 2A, the median value is written above the line within each box and the lack of
significant pairwise differences in means are indicated with the same letter above each box.
 E. Wohl et al. / Geomorphology 126 (2011) 108–120 115

Cedar1-Transect 5 Cedar2-Transect 7 Cedar3-Channel Cedar3-Transect 4 Cedar3-Transect 5

270 90 270 90 270 90 270 90 270 90

180 180 180 180 180

Cedar3-Floodplain Cedar4-Channel Cedar4-Transect 4 Cedar4-Transect 5 Cedar4-Transect 6

270 90 270 90 270 90 270 90 270 90

180 180 180 180 180

Cedar4-Floodplain Toms1-Channel Toms1-Transect 5 Toms1-Transect 6 Toms1-Transect 7

270 90 270 90 270 90 270 90 270 90

180 180 180 180 180

Toms1-Floodplain Toms3-Channel Congaree Cr.-Channel Beidler Forest-Transect C Beidler Forest

270 90 270 90 270 90 270 90 270 90

180 180 180 180 180

Lynches1-Transect 1 Lynches1-Transect 3 Lynches1-Floodplain Lynches2-Transect 1 Lynches2-Transect 3

270 90 270 90 270 90 270 90 270 90

180 180 180 180 180

Lynches2-Floodplain

270 90

180

Fig. 3. Rose diagrams of wood orientations for transects in which wood orientation did not follow random distribution. All transects that are not labeled as ‘channel’ or ‘forest’ are in the
floodplain. Diagrams labeled ‘floodplain’ represent all of the floodplain transects at that site grouped together. The arrow in each diagram represents the primary direction of stream flow.

downstream. Instream wood removal, however, does not occur at the
Toms Creek sites. Large jams do occur on other, larger rivers in the
area, including the Lynches River (K. Meitzen, University of South
Carolina, personal communication, 2010), but these jams are at least
partially removed by periodic mechanical channel clearing. Conse-
quently, we confine our discussion of large jams to those in the old-
growth forest sites within Congaree National Park.
Large, channel-spanning jams are commonly reported from
laterally confined channels in colder temperate environments (e.g.,
Kaczka, 2003; May and Gresswell, 2003; Oswald and Wohl, 2008;
116 E. Wohl et al. / Geomorphology 126 (2011) 108–120

Table 5
Wood loads reported for old-growth forests.

Location Forest type Valley Drainage area Channel V/L V/A Reference
geometrya (km2) width (m) (m3/100 m)b (m3/ha)c

Northwestern North America Coniferous temperate Both types 0.4–325 3–50 0–305 Fox and Bolton, 2007
rainforest Confined 0.1–60.5 3–24 478 (230–750) Lienkaemper and Swanson,
1987
Unconfined 0.7–55 5–26 24 (7–62) Robison and Beschta, 1990
Eastern & central USA Temperate deciduous Both types 40 15 25 (7–62) 291 (75–622) Morris et al., 2007
Confined 3.5–11 8–11 21.7 Hedman et al., 1996
New Zealand Subantarctic to subtropical Confined 0.6–5.8 3–6 18 (4–49) 206 (85–470) Meleason et al., 2005
rainforest
Chilean Andes Temperate Nothofagus & Confined 5.5–11 5–16 39 (3.8–236) 495 (4.8–4000) Andreoli et al., 2007
Araucaria
Tierra del Fuego, Argentina Subantarctic Nothofagus Confined 5–12.9 4–10 7 (1–24) 121 (20–395) Mao et al., 2008
Costa Rica Tropical wet Confined 0.1–8.5 3–15 12.3 (3–35) 189d (41–612) Cadol et al., 2009
Southern Rocky Subalpine conifer Confined 2.4–29 3–10 6.6–27.1 182 (92–254) Richmond and Fausch, 1995
Mountains, USA Confined 7.8–19.8 5–20 7 (0.2–32) 161 (0–708) Wohl and Cadol, 2011
South Carolina, USA Bottomland hardwood Unconfined 110–270 8–14 13.8 (9.4–18.6) 134 (102–178) This study
a
Valley geometry distinguishes channels that are relatively laterally confined (limited to no floodplain) from those that are relatively unconfined (extensive floodplain).
b
V/L is volume per length of channel; average value, followed by range in parentheses.
c
V/A is volume per area of channel; average value, followed by range in parentheses.
d
The average drops to 174 if the outlier (612 m3/ha) is excluded; the remaining 29 stream segments have values ≤428 m3/ha.

Wohl and Cadol, 2011), where wood recruited to the channel cannot
readily disperse laterally and where wood transport can be inter-
rupted by changes in channel geometry. In laterally confined streams
draining old-growth forest in the Colorado Front Range, for example,
51% of logs are incorporated in a jam (defined as three or more pieces
touching one another), whereas 29% of logs are included in a jam in
the unmanaged Congaree stream sites draining old-growth forest.
Jams may be less common in laterally confined tropical streams,
although this inference is based on very limited field data. Stream
surveys at La Selva Biological Station in Costa Rica indicate that 42% of
logs are included in a jam, and only seven channel-spanning jams
were present among 30 channel segments surveyed (each channel
segment 50 m long) (Cadol et al., 2009). Although channel dimen-
sions and log sizes are broadly comparable at the three sites, the ratios
of log length/channel width and log diameter/flow depth display
some important differences (Table 6). These ratios provide an indirect
indicator of the likelihood that logs will undergo fluvial transport;
lower values of these ratios indicate greater likelihood of transport
because long pieces are more likely to wedge against the channel
banks or other obstructions and depth is less likely to reach sufficient
values to float thicker pieces (Abbe et al., 1993; Braudrick and Grant,
2000, 2001; Haga et al., 2002). The low ratios in Table 6 suggest that
instream wood at the Congaree sites should be highly mobile and this,
combined with the fairly uniform channel geometry when compared
to sites such as those in the Front Range where large boulders
protrude above the maximum flow depth and channel and valley width
show substantial longitudinal variations (Wohl and Cadol, 2011), may
explain the relative lack of jams at the Congaree sites. Large, channel-
spanning jams have also been reported from laterally unconfined
channels in the temperate zone; the jams facilitate overbank flows and
sediment exchange between the channel and floodplain (O'Connor et
al., 2003; Brummer et al., 2006; Curran, 2010; Sear et al., 2010).
Although data of the type in Table 6 are not available for these sites, the
lack of large jams at the Congaree sites may reflect the more rapid wood
decay and much greater flow depths than are present at laterally
unconfined sites described thus far in the temperate zone.
The lack of jams at the Congaree sites is also reflected in the results
from the K statistics. First used by Kraft and Warren (2003) to assess
aggregation and segregation of wood along 700–1000 m lengths of
eight streams in the Adirondack Mountains of New York, USA, this
type of statistical analysis has thus far been applied to only a few sites.
Kraft and Warren (2003) found that individual pieces were
aggregated at spatial extents of 0 to 40 m and segregated at spatial
extents of 80 to 100 m following extensive wood deposition during an
ice storm. This suggests that pieces cluster into jams at shorter length
scales, with jams more uniformly spaced than random at greater
length scales. Surveying 12 laterally confined stream segments each
1.25 km long in the Colorado Front Range, Wohl and Jaeger (2009)
found wood pieces to be nonrandomly distributed at lengths of tens to
hundreds of meters. Jams tend to cluster in stream segments at scales
of a few hundred to a few thousand meters in streams of northern
Michigan, largely as a result of longitudinal variations in valley
geometry (Morris et al., 2009). In four laterally confined channel
segments 3–9 km long in the Colorado Front Range, wood pieces were
highly aggregated for all sites at all length scales analyzed, indicating a
strong tendency to cluster into jams (Wohl and Cadol, 2011).
Differences in wood dynamics (recruitment, transport, and decay)
between various sites provide some insight into the lower instream
wood loads relative to other old-growth sites and the lack of large
logjams at the Congaree sites. Recruitment at Congaree reflects
individual tree mortality, bank erosion, transport from the floodplain,
and mass mortality associated with hurricanes. Numerous examples
of individual tree fall, in some cases induced by bank erosion, were
observed during a month of field work at the site. Wood pieces larger
than the minimum dimensions for this study were also commonly
produced by falling branches. Sharitz et al. (1993) estimated a
recurrence interval circa 25 years for hurricanes during the past
century at Congaree National Park and documented 37% of trees in
bottomland hardwood forest study plots as being moderately to
severely damaged by Hurricane Hugo in 1989 (although only 10% of
trees were damaged in slough study plots). Phillips and Park (2009)
propose that, although bank erosion is the primary wood recruitment
mechanism for sinuous, sand-bed channels in coastal plain rivers of
the southeastern U.S. between hurricanes, historic megajams
reported from these rivers could represent massive blowdown from
hurricanes. Based on all of these observations, lower wood loads at the
Congaree sites are unlikely to represent rates of recruitment
substantially lower than other old-growth forest sites.
Wood transport at the Congaree study sites reflects interactions
among hydrology, hydraulics, and channel geometry. Potential
transport events during which flow depth greatly exceeds average
wood piece diameter occur multiple times each year during overbank
floods. Although it is difficult to quantify frequency and duration of
overbank flooding at a specific site because of backwater effects and
secondary channels that complicate stage-discharge relationships,
gage records from Cedar Creek during the period of March 2009 to
 E. Wohl et al. / Geomorphology 126 (2011) 108–120 117

Table 6
Parameters indicative of relative wood transport capacity at three field sites.

Site Wa Wood Lavgb Wood davgc Lavg/Wd davg/flow

(m) (cm) (cm) depthe

Colorado Front 5–17 250 19.5 0.31 0.28

Range (0.09–1.27) (0.08–1.32)
La Selva, Costa 3–15 395 18.8 0.62 0.35
Rica (0.23–1.85) (0.13–1.13)
Congaree National 8–13 334 21.7 0.30 0.14f
Park (0.19–0.40) (0.05–0.26)
0.07
(0.06–0.08)
a
W is bankfull channel width; range of values for different channel segments surveyed.
b
Wood Lavg is average wood piece length.
c
Wood davg is average wood piece diameter.
d
Lavg/W is the ratio of average wood piece length to channel width; average for all
channel segments surveyed, followed by range in parentheses.
e
davg/flow depth is ratio of average wood piece diameter to bankfull flow depth;
average for all channel segments surveyed, followed by range in parentheses.
f
These values represent a minimum because bankfull depth was used; depth greatly
exceeds bankfull during overbank flooding, so the second set of values assumed a flow
depth of 3 m.

Fig. 4. (A) View across Cedar Creek Reach 4 at very low flow, showing small jam
forming around ramp from right bank (in foreground; flow is left to right). Channel is
~ 15 m wide here and is not appreciably wider at the jam; the apparent widening at the
right rear is associated with a channel bend. (B) Logjam at the mouth of a small,
secondary channel along the Congaree River. Channel is ~ 15 m wide and flow is right to
left when it occurs during out-of-bank flooding from the mainstem. No appreciable
increase in sediment storage or bed scour is apparent in association with the jam.
(C) Large floating log trapped among closely spaced tree trunks on the floodplain of the
Congaree River during flooding associated with tropical storm Ida in November 2009.
Flow depth was ~ 3 m and flow velocity ≫1 m/s.
March 2010 record eight floods with stage at least 0.7 m above the
threshold for overbank flow, each of which lasted a few days.
Observations during the November 2009 flood at Congaree indicate
that longer pieces of wood have limited transport distances across the
floodplain because they are trapped by the trunks of standing trees
(Fig. 4C), but shorter pieces are presumably able to move more readily
between the channel and floodplain. Working on the Ogeechee River,
a coastal plain stream with tributary swamps, Benke and Wallace
(1990) found that wood load increases during moderate floods as
mobile pieces are transported from upstream, broken branches from
the riparian area enter the channel, and pieces come in from overbank
areas, even though the floodplain buffers stream velocity and large
wood is not highly mobile. They also found that floodplain wood was
more mobile (21–84% of tagged pieces moved during moderate
flooding) than instream wood (17% of pieces moved). The prevalence
of wood transport at Congaree is also suggested by the proportion of
piece types in channel transects; only 13% of the pieces surveyed were
ramps or bridges (1% bridges) that have not experienced significant
fluvial transport, whereas 46% of all pieces surveyed were buried,
suggesting substantial scour and fill in these sand-bed streams during
high flows. By comparison, 26% of the pieces surveyed in laterally
confined tropical streams in Costa Rica were ramps or bridges (Cadol
et al., 2009) and 18–24% of wood in laterally confined channels of the
Colorado Front Range were ramps or bridges (Wohl and Cadol, 2011).
Based on these observations and comparisons, we infer that
floodplain and instream wood pieces and jams are likely to be quite
mobile at the Congaree sites because of the frequent occurrence of
flows exceeding the minimum depth necessary to float most wood
pieces.
The limited evidence available for the Congaree field sites suggests
rapid decay relative to other sites in the temperate zone (M. Duever,
South Florida Water Management District, personal communication,
2009). In this study, 95% of the channel wood and 80% of the
floodplain wood was in decay category 3, the most advanced category
assigned. Smaller pieces of floodplain wood that could be lifted by
hand were commonly weakly attached to the underlying soil by the
roots of plants growing on the decaying log. Benke and Wallace
(1990) noted that floodplain wood loads along the Ogeechee River are
low although swamp forest productivity is high relative to other
mixed deciduous forests. They attribute this discrepancy to rapid
decay and fragmentation. Rates of wood turnover on the forest floor
typically increase with increasing mean annual temperature and
precipitation (Zabel and Morrell, 1992). Although rates of wood decay
in the subtropical climate at the Congaree sites are not likely to be as
high as those documented for tropical rainforests (Clark et al., 2002),
they are likely to exceed rates of decay in other temperate environments
where drier conditions and/or seasons with much lower temperatures
retard wood decay.
118 E. Wohl et al. / Geomorphology 126 (2011) 108–120
In summary, the lower wood loads at the Congaree study sites
relative to other unmanaged streams flowing through old-growth
forests could reflect some combination of slower wood recruitment,
greater mobility and lesser instream retention, and more rapid decay.
Of these, the combination of high mobility and rapid decay likely
explains the lower values of wood load relative to most temperate
environments.
The streams surveyed in tropical Costa Rica (Table 5) are the most
similar to those at the Congaree sites in terms of rapid wood decay and
high mobility associated with frequent large floods. Channel-
spanning jams are also notably uncommon in the Costa Rican streams,
although smaller jams are more common than along the Congaree
stream sites. The mean tree diameter and basal area are comparable
between the sites (Cadol et al., 2009). The primary difference between
the Costa Rican and Congaree sites is the unconfined valley geometry
and potential for wood exchange between the channel and floodplain
at the Congaree sites. As noted previously, near-channel transects at
Congaree have a dearth of wood, suggesting primarily transport into
the channel rather than onto the floodplain, but transects at greater
distances from the channel have comparable wood loads, probably
partly because long pieces cannot effectively move great distances
into channels. Thus, the evidence does not suggest that lack of lateral
confinement would cause noticeably lower wood loads at the
Congaree sites, although one-third of the 30 sites in Costa Rica have
greater wood loads than the highest values at the Congaree sites.
Greater insight into the mechanisms underlying these differences
requires further field work at both tropical and subtropical sites.
The observed absence of channel-spanning jams at the Congaree
sites may also partly reflect temporal variation in relation to storm
sequence. The field work was conducted during Oct–Nov 2009, at the
start of the winter–spring flood season. Channel-spanning jams were
subsequently observed at Congaree following a series of unusually
large and frequent floods during winter 2010 (T. Thom, Congaree
National Park, personal communication, 2010). By autumn 2010, most
of the instream jams had dispersed or become substantially smaller.
The floodplain jams were mostly still present, but degrading from
wood decomposition and dispersion during feeding by feral hogs
(T. Thom, Congaree National Park, personal communication, 2010). It
is not clear how long these jams will persist, but their absence during
the 2009 field work suggests that they are relatively transient
features. Channel-spanning jams are also more likely to form fol-
lowing major hurricanes such as Hugo in 1989, when large amounts of
wood were introduced to channel and floodplain environments. The
absence of such jams, assuming that they did form, is in contrast to
channel–floodplain systems in the temperate zone, where long-term
studies document persistence of individual jams for as much as
30 years (D. Sear, University of Southampton, personal communica-
tion, 2010).
The emphasis in this study was on spatial patterns of wood
distribution and potential controls on those patterns, and we did not
measure geomorphic or ecological effects of instream or floodplain
wood. Field observations suggest that the smaller accumulations and
more transient character of individual wood pieces and jams in the
Congaree sites creates less bed and bank scour, channel widening,
sediment storage, and overbank flow than in comparably sized
channels in temperate environments. Several studies of sand-bed
streams in the southeastern U.S. indicate that instream wood effectively
increases hydraulic resistance (Dudley et al., 1998), traps sediment and
enhances channel stability in incised channels (Wallerstein and Thorne,
2004), and provides high quality invertebrate habitat. Despite forming
b20% of the area colonizable by macroinvertebrates in Congaree's
Cedar Creek, instream wood has N90% of the standing crop at a given
time, as well as the highest species richness and density of available
substrates (Smock and Gilinsky, 1982). Studies of other bottomland
hardwood wetlands and blackwater rivers also indicate that instream
and floodplain wood is a hot spot for invertebrate richness and
arthropod biomass (Benke, 2001; Braccia and Batzer, 2001), although
macroinvertebrate abundance and diversity are also influenced by the
location of the wood (swamp versus tributary and outflow stream sites;
Thorp et al., 1985) and potentially by the degree to which the wood is
decayed (Kaller and Kelso, 2006). Submerged wood is especially
important in sand-bed streams, where it provides stable attachment
points for filter-feeding macroinvertebrates (Benke, 1998). Floodplain
wood provides invertebrate habitat (Benke, 2001; Braccia and Batzer,
2001) and germination sites for plant propagules (Schowalter et al.,
1998), enhances nutrient cycling and soil formation (Zalamea et al.,
2007), and creates habitat diversity for various species of plants and
animals (Harmon et al., 1986). Hydrochory is important to bald cypress
and water tupelo, and seeds of these species are preferentially deposited
against logs, trees, cypress knees, and other emergent substrates on the
floodplain (Schneider and Sharitz, 1988). Given the geomorphic and
ecological significance of instream and floodplain wood, the relatively
low wood loads and transience of wood suggests that maintenance of
these functions requires continuing wood recruitment and thus the
presence of mature riparian forests and both wood and hydrologic
connectivity across floodplain bottomlands. Wood loads are demon-
strably lower outside of old-growth forests and in otherwise unman-
aged channels from which wood is annually removed. These lower
wood loads may compromise the geomorphic and ecological influence
of instream and floodplain wood, although further research is needed to
demonstrate whether wood load has a linear or threshold relationship
to geomorphic and ecological function.
6. Conclusions
Instream wood loads observed in sand-bed channels draining old-
growth floodplain forests at Congaree National Park, South Carolina, are
lower than values documented for most other old-growth environ-
ments, which we interpret to reflect the combined effects of relatively
high transport capacity during frequent and prolonged out-of-bank
flooding and relative rapid decay in this warm, moist environment.
Although none of the measured variables correlate with instream
wood loads, floodplain wood loads reflect distance from the channel and
riparian forest characteristics, with the greatest wood loads at the
farthest distance from the channel and in old-growth bottomland
hardwood forests. Channel geometry does not appear to exert the sig-
nificant influence on instream wood loads that has been documented for
more laterally constrained channels. Instream wood loads are signifi-
cantly higher than floodplain wood loads within 100 m of the channel,
but show little difference in aggregation with respect to floodplain wood.
Instream wood is also more likely to be preferentially oriented than
wood on the floodplain. The absence of channel-spanning jams during
field work at the start of the winter–spring flood season may reflect
temporal variability, with channel-spanning jams forming during larger
and more prolonged flooding, but not persisting for more than a year.
Acknowledgements
The Warner mini-grant program of the Warner College of Natural
Resources at Colorado State University funded this research. Will Graf
suggested Congaree National Park as an appropriate site to study wood
and helped with logistical arrangements for field work. The staff at
Congaree National Park went out of their way to facilitate field work and
to provide logistical support; particular thanks to Theresa Thom and
David Shelley. Kimberly Meitzen provided valuable insights into
hydrology and floodplain dynamics, as well as a guided kayak trip
during the November 2009 flooding. Thanks to all for an enjoyable
sojourn in the swamp. EW also thanks the Department of Geography at
Durham University, United Kingdom for a Distinguished International
Fellowship that facilitated work on this paper during a sabbatical leave.
Comments by David Shelley, Kimberly Meitzen, and two anonymous
reviewers helped to improve the manuscript.
 E. Wohl et al. / Geomorphology 126 (2011) 108–120
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