ECOHYDROLOGY

Ecohydrol. (2014)
Published online in Wiley Online Library
(wileyonlinelibrary.com) DOI: 10.1002/eco.1523

Seasonal water use by deciduous and evergreen woody species

in a scrub community is based on water availability and
root distribution
Patrick Z. Ellsworth† and Leonel S. L. Sternberg*
Department of Biology, University of Miami, Coral Gables, FL, 33146, USA

ABSTRACT
In a seasonally dry plant community of central Florida, USA, that experiences water limitation in the dry season and high water
availability in the wet season, we first tested whether evergreen woody species shift from shallow water in the wet season to deep
water in the dry season. Second, we tested whether deciduous woody species restrict water uptake to the shallow soil during the
wet season and cease water uptake during the dry season. To address these questions, we measured water source use of two
deciduous and three evergreen species over 13 months using stable isotopes. As hypothesized in previous studies, we showed
that leaflessness in deciduous plants is an important source of stem water isotopic fractionation. Therefore, we compared stable
isotope ratios of stem water only when deciduous species had leaves and found that all species, except the evergreen Lyonia
ferruginea, used proportionally the same water sources. Early dry season water use was based on water availability for all species
except L. ferruginea, and deep soil (50–150 cm) was the most important water source. During the late dry and wet seasons, water
uptake from each soil layer was based on its respective proportion of fine roots. Nevertheless, deep water remained an important
water source throughout the year. This study clearly demonstrates the limitations of using stable isotopes of stem water when
comparing deciduous versus evergreen species. Further, this study is the first to directly quantify depth of water uptake via
isotope analysis and couple these findings with root distribution. Copyright © 2014 John Wiley & Sons, Ltd.

KEY WORDS stable isotopes; dry seasonal plant community; Florida sandhill; deciduous; evergreen; water source use;
rooting depth
Received 1 November 2013; Revised 21 April 2014; Accepted 9 June 2014

INTRODUCTION other seasonally dry plant communities have hypothesized

that drought-deciduous woody species fit into this category
Dry seasonal ecosystems are defined by a pronounced and
because they lose their leaves in the dry season and
consistent dry season that results in water limitation,
therefore lose their ability to transpire and photosynthesize
especially in shallow and/or sandy soils with low water
(Medina, 1984; Sarmiento et al., 1985; Olivares and
retention (Murphy and Lugo, 1986; Menges and Gallo,
Medina, 1992; Jackson et al., 1995; Givnish, 2002;
1991; Weekley et al., 2007; Saha et al., 2008). Woody
Bowman and Prior, 2005; Markesteijn and Poorter, 2009;
plant species in the Florida sandhill plant community may
Hasselquist et al., 2010).
function with high water availability during the wet season
Another strategy is the utilization of shallow water
and low water availability and decreasing soil water
during the wet season and deep water during the dry
potentials during the dry season by different strategies.
season. Studies in Mediterranean, desert savannas, and
One strategy commonly thought to occur with drought
tropical dry forests have shown that some evergreen
deciduous plants is the presence of transpiration and
species shift water uptake from shallow water to deep
photosynthetic activity only during the wet season,
water in the dry season (Walter, 1971; Scholes and Archer,
avoiding the drought conditions by losing their leaves
1997; Jackson et al., 1999a; Hasselquist et al., 2010;
and remaining dormant in the dry season. Many studies in
Esquivias et al., 2013). These trees also have been called
drought delayers because they compensate for low shallow
water availability by accessing deep water (Paz, 2003; Slot
*Correspondence to: Leonel Sternberg, Department of Biology, Univer- and Poorter, 2007; Poorter and Markesteijn, 2008;
sity of Miami, Coral Gables, FL 33146, USA. Markesteijn and Poorter, 2009).
E-mail: leo@bio.miami.edu
†
Current Address: School of Biological Sciences, Washington State The above studies, however, were usually done by
University, Pullman, WA, 99164-4236, USA sampling plants and soil conditions with low frequency and

Copyright © 2014 John Wiley & Sons, Ltd.

 P. Z. ELLSWORTH AND L. S. L. STERNBERG
did not analyse root abundance together with water uptake
measurements. Here, we intensively sampled plant and soil
conditions as well as root growth and test for differences in
water uptake between evergreen versus deciduous shrubs and
small trees in a Florida seasonal plant community. This
community, known as Florida sandhill (Menges, 1999), is
characterized by high rainfall in the summer (wet season) and
low rainfall and subsequent soil drying in the winter (dry
season) that decreases the soil moisture at all depths, but
creates a soil moisture profile characterized by an increase in
moisture with depth (Weekley et al., 2007). Water availability
changes most drastically in the shallow soil, while water
availability remains comparatively high in the deep soil
layers. During the dry season, the well-drained sand,
underlying the sandhill plant community in the Florida scrub,
exacerbates soil drying and water limitation. Fluctuation in
soil water availability requires perennial plants to adapt to low
water availability and optimally still be able to take advantage
of high water availability during the wet season.
We first tested the hypothesis that evergreen woody
species in this Florida scrub community use the strategy
mentioned above, i.e. they are drought delayers and take up
shallow soil water during the wet season and deep soil water
during the dry season when shallow soil water is no longer
available. Second, we tested whether deciduous species
were drought avoiders, i.e. they take up shallow soil water
during the wet season, but once the shallow soil dries during
the dry season, they lose their leaves and remain dormant
until shallow water becomes available again. Because it has
been suggested previously that leaflessness may change the
isotopic signature of stem water (Phillips and Ehleringer,
1995) and might confound any interpretation of water
source use by isotopic analyses, we first conducted a
defoliation experiment in evergreen trees and monitored the
isotopic signature of stem water. With an understanding of
the constraints in our ability to determine source water by
isotopic analyses from deciduous trees during their dormant
phase, we compared the changes in isotope ratio of stem
water of three evergreen and two deciduous woody species
with leaf phenology. Using an isotopic mass balance
approach, we were then able to capture inter- and intra-
seasonal differences in water source use.
METHODS AND MATERIAL
Two studies were conducted. The first established
constraints in interpreting source water in deciduous trees
by isotope analyses. It was a defoliation experiment that
determined if leaflessness can change the isotopic compo-
sition of hydrogen and oxygen in stem water. This first
study allowed us to better interpret data from our second
study. The second study was a field study on water source
use by deciduous and evergreen species.
Copyright © 2014 John Wiley & Sons, Ltd.
Defoliation experiment
The defoliation experiment was conducted in a pineland
plant community in Coral Gables, Florida, USA. Five
evergreen Quercus virginiana Mill. trees were selected
during the dry season (2010). This species was not found in
study sites where we measured water source use nor was it
one of the study species. However, we assumed that stem
water would behave similarly in a defoliated branch of this
evergreen woody species as it would behave in a leafless
branch of the deciduous study species. One large branch
having several secondary branches on each tree (treatment
branch) was completely defoliated by plucking the leaves
manually. After the initial defoliation, leaves were allowed
to regrow on the defoliated branches. The degree of leaf
regrowth on defoliated branches was recorded each time
stem samples were collected. A non-defoliated branch on
the same tree served as a control. Before defoliation, stem
water samples were collected from both treatment and the
control branches. Afterwards, stems were collected from
both defoliated and non-defoliated branches of each tree 27,
34, 57, 68, and 92 days after defoliation. At collection, stems
were de-corticated, placed in 12-mm tubes, sealed with
parafilm, and stored at
18 °C (Saha et al., 2008).
Field study of water source use
Description of site and species studied. Three sites were
selected in a sandhill plant community at Archbold
Biological Station in south-central Florida, USA (Figure 1).
The southern ridge sandhill community is located on the
southern part of the Lake Wales Ridge, which represents
the ancient beaches from the late Pliocene (Watts and
Hansen, 1994). The soils are well-drained, acid,
quartzipsamments sands (Kalisz and Stone, 1984), have
very low nutrient and organic matter content (Huck, 1987),
and experience periodic drought (Menges and Gallo, 1991;
Saha et al., 2008). The woody species that dominate the
southern ridge sandhill plant community are the evergreen
oaks—Quercus myrtifolia Willd. (myrtle oak), Q. inopina
Ashe (sandhill oak), Q. geminata Small (sand live oak), Q.
chapmanii Sarg. (Chapman oak), other evergreen species
such as Serenoa repens (Bartram) Small (saw palmetto),
Lyonia ferruginea (Walter) Nutt. (rusty staggerbush), Pinus
elliottii Engelm. var. densa Little & Dorman (Florida slash
pine), and P. clausa (Chapm. ex Engelm.) Vasey ex Sarg.
(sand pine), and deciduous species such as Carya floridana
Sarg. (scrub hickory), Q. laevis Walter (turkey oak), and
Asimina obovata (Willd.) Nash (bigflower pawpaw)
(Menges, 1999). The climate of the site is characterized
by hot, wet summers and mild, dry winters that extend
from December through May. Mean annual rainfall at
Archbold (1932–2009) was 1346 mm, with approximately
61% falling between June and September (Data provided
by Archbold Biological Station; Figure 2).
Ecohydrol. (2014)
 DECIDUOUS AND EVERGREEN WATER UPTAKE
Figure 1. Map of Florida showing the geographical range of the Lake
Wales Ridge (region highlighted in black) and the location of Archbold
Biological Station (Weekley et al., 2008).

1
Figure 2. Precipitation (mm month ) at Archbold research station from 1
January 2008 to 31 December 2009 (Data provided by Archbold
Biological Station).
Three sites were selected in this community: Sandhill
Long Unburned site 1 (SHLU1; N27° 11.156′; W81°
20.084′), Sandhill Long Unburned site 2 (SHLU2; N27°
09.083′; W81° 21.448′), and Sandhill Recently Burned
(SHRB; N27° 11.280′; W81° 20.181′). The water table
near Sandhill Recently Burned and Sandhill Long
Unburned 1 sites had a median depth of 24.3 ± 0.1 m (U.
S. Geological Survey). These three sites were selected to
cover the range of stand age and fire history that is found
on the southern ridge sandhill. Sandhill Recently Burned
was burned last in 2002 (6 years before this study) and was
burned five times in the 20 years prior to this study, which
is considered historically typical of the southern ridge
sandhill (Menges, 1999). Sandhill Long Unburned 1 and 2
were last burned in 1996 and 1927, respectively. As
expected because of a long fireless period (Abrahamson,
Copyright © 2014 John Wiley & Sons, Ltd.
1984; Mushinsky and Gibson, 1991; Mushinsky, 1992),
Sandhill Long Unburned 2 site had the most organic matter
accumulation on the soil surface. Sandhill Long Unburned
2 was also the tallest plant community with some C.
floridana and Q. laevis over 15 m tall and had the least
dense understory typical of sandhill that has not been
burned for a considerable time. Conversely, Sandhill
Recently Burned site was the most open and shortest site
with most woody species under 3 m tall. The soil surface at
this site had very little organic litter. The groundwater table
at Sandhill Long Unburned 2 had a yearly mean depth of
2.3 ± 0.3 m (Archbold Biological Station, unpublished
data). The elevation at Sandhill Long Unburned 1 was
58 m, and Sandhill Long Unburned 2 was 44 m and
Sandhill Recently Burned 62 m.
Two deciduous (Q. laevis and C. floridana) and three
evergreen species (Q. myrtifolia, Q. geminata, and L.
ferruginea) were selected for this study because of their
ubiquity in the southern ridge sandhill plant community
and at each of these three sites. Among angiosperms, these
species represent the vast majority of individual woody
plants at the site (Menges, 1999). In addition to their
abundance in the Florida scrub, we chose both deciduous
and evergreen Quercus because they would provide an
intra-generic comparison of water source use that differs in
leaf phenology.
Sampling methods
Phenology observations. Deciduous species were moni-
tored for leaf senescence in the early dry season and for
bud break and leafing out in the late dry season at each of
the three sites to determine when deciduous species were
with leaves.
Stem and soil water extraction for isotope analyses. We
collected stems and soils in the months of May, June,
August, September, November, and December of 2008 and
January, March, and May of 2009. At each of the three
sites, stems were collected from five individual plants of
each of five species, and the individual plants of each
species were at least 10 m from each other. Also, only one
stem was collected from a clump of the same species,
especially the clonal species Q. myrtifolia and Q. geminata
(Menges, 1999). Likely, the entire clump is the same
individual. The stems were de-corticated and stored in 12-
mm outer diameter tubes. The tubes were stoppered and
sealed with parafilm (Saha et al., 2008). The tubes were
kept in a dry, cool ice chest until they could be stored at

18 °C. All stems were sampled from branches that had
fully suberized bark and were far from leaves to eliminate
the effect of evaporative enrichment in the sapwood water
(Dawson and Ehleringer, 1993).
A hand auger with a 12-cm diameter bucket was used to
collect three 150-cm-deep soil cores at each site on each
Ecohydrol. (2014)
 P. Z. ELLSWORTH AND L. S. L. STERNBERG
collection date. Soil samples were collected from each core
at the following depth increments: 0–10, 10–20, 20–30,
30–50, 100–120, and 130–150 cm. From each soil depth,
approximately 50 g of soil was collected and stored in
screw-cap, glass culture tubes, and sealed with parafilm.
The soil samples were stored in a cool, dry ice chest until
they could be stored at
18 °C. Water was extracted from
stem samples by cryogenic vacuum distillation as
described by Vendramini and Sternberg (2007). Water
was extracted from the soil samples using a specially
designed distillation line where six samples could be
distilled simultaneously.
Soil water content and availability. To calculate potential
water uptake based on the distribution of available water,
soil volumetric water content (VWC) was measured from
soil samples collected at the same time and same depths
that stem and soil samples were collected for determination
of plant water source use.
As soil matric potential better reflects moisture avail-
ability to plants than VWC, we calculated soil matric
potential at each depth at each site, using a model
developed by Saxton and Rawls (2006) that incorporates
soil moisture, percent organic matter, salinity, and soil
texture. Their model uses empirically derived equations
predicting soil matric potential from the currently
available USDA soil database using the variables of soil
texture, salinity, and organic matter. Percent organic
content was calculated as the difference between soil
weight after drying for 24 h at 90 °C and soil weight after
ashing at 600 °C for 4 h. The following soil conditions
were used to calculate soil matric potential: 96% sand, no
salinity, and organic matter content depended on the soil
depth and site. Soil texture was uniform with depth at all
sites (Carter, 1989).
Water availability was calculated as the difference
between mean VWC of the soil when samples were
collected and minimum VWC from which plants can
typically extract water. Minimum VWC was calculated
from the VWC when the soil matric potential was
1.5 MP
using the above soil matric potential model. A matric
potential of
1.5 MPa was chosen as a threshold to
determine when soil water was largely unavailable
because, with the exception of extremely dry years, the
midday water potential of woody plants in the southern
ridge sandhill seldom drops below
1.5 MPa (Saha et al.,
2008). Mean VWC was averaged from soil collections
made in three time intervals representing early dry
(November and December 2008 and January 2009), late
dry (May 2008 and March and May 2009), and wet season
(June, August, and September 2008).
Root distribution. Soil cores, specific for root distribution,
were taken from 14 locations at the site SHRB. At each
Copyright © 2014 John Wiley & Sons, Ltd.
specific location at SHRB, three soil cores were collected
within 50 cm of each other: one each in the dry season
(January 2010), in the late dry season after deciduous
species had grown new leaves (March 2010), and during
the wet season (August 2010). Each core was 7 cm in
diameter and 150 cm long and was divided into three
sections: 0–20, 20–50, and 50–150 cm. The soil samples
were stored at 4 °C until roots were removed from the soil.
All roots were scanned using a high resolution flat-bed
scanner (1200-dpi resolution, 19-μm pixel size, 48-bit
color, TIFF format; Epson Scanner Perfection V500 Photo,
USA). We categorized root branches into terminal order
classes based on the morphometric approach developed by
Fitter (1982). All root branches with a meristematic end
were considered first terminal order roots. Second, terminal
order roots were those that had a junction with a first
terminal order root. This method was used to define third
and fourth order roots as well.
Root distribution was calculated as the proportion of the
total number of roots of the first four terminal orders found
in the top 150 cm that are found in each soil layer: 0–20,
20–50, and 50–150 cm. Root count from all 14 cores were
summed together and were not calculated separately. The
total number of roots was calculated as the total number
root branches from the first four terminal orders in each soil
layer divided by the total number of root branches found in
the top 150 cm.
Isotope analysis
Soil and stem water were analysed in a Multiflow system
connected to an Isoprime mass spectrometer (Elementar,
Hanau, Germany). The following modification of Prosser
and Scimgeour (1995) was used to analyse the hydrogen
isotope ratio of water. Each water sample (0.5 ml) and a
cuvette containing ~1 mg of platinum black powder (Sigma-
Aldrich, St. Louis, Missouri, USA) were placed in 5.9-ml
vials (Exetainer vials; Labco, High Wycombe, UK) and
sealed with screw caps that had a pierceable rubber septum
(Exetainer cap; Labco). A 10% H2/He gas mixture was
injected into the vials and equilibrated with water vapor for
24 h. The isotopic analysis of the equilibrated hydrogen gas
occurred as described by Vendramini and Sternberg (2007).
After hydrogen isotope analysis, a 5% CO2/He gas mixture
was flushed through the vials and was equilibrated with
water for a period of 48 h. The equilibrated CO2 gas was
analysed to derive the oxygen isotope ratios of the water as
in Vendramini and Sternberg (2007). All isotope ratios were
expressed in terms of per mil (‰):
 
Rsample
δ H or δ O ¼
2 18

1 1000
Rstandard
where Rsample and Rstandard are the 2H/1H or 18O/16O ratios
of the sample and the standard, respectively. The standard
Ecohydrol. (2014)
 DECIDUOUS AND EVERGREEN WATER UPTAKE
used was Vienna Standard Mean Ocean Water, and the
precision of analysis for hydrogen and oxygen isotope ratios
was ±3‰ and ±0.1‰, respectively.
Data analysis
In the defoliation experiment, the difference between
control and defoliated branches was reported as isotopic
separation (Δ), which is the defoliated stem water δ value
minus the control stem water δ value.
In the water source use study, IsoSource (EPA v. 1.3.1)
was used to calculate the most likely proportion of water
used by plants at each soil depth (Phillips and Gregg,
2003). IsoSource is more effective than standard mixing
models at calculating water source use because it calculates
the mixture of water sources that are possible to produce
the isotopic signature of stem water instead of assuming
that the plant is only taking water from one soil layer.
Isotopic composition of oxygen was used to calculate the
possible distribution of water uptake along the soil profile
because oxygen provided a more accurate and reliable
tracer of water source use than hydrogen. Plants do not
fractionate oxygen isotopes, whereas some halophytic and
xerophytic plants have been found to fractionate hydrogen
isotopes (Lin and Sternberg, 1993; Ellsworth and Williams,
2007). The proportional water uptake at each depth was
calculated from the isotopic ratios of soil water and stem
water for each individual plant collected in this study,
except when deciduous plants were leafless. The total
number of individual plants that were sampled over the
study was 675. Soil water sources and their characteristic
oxygen isotope ratio were categorized into three depth
classes (0–20, 20–50, and 50–150 cm) based on their
similar isotopic signature. Proportional use of the three
depth classes was calculated using IsoSource for each
individual plant sampled in the study by comparing oxygen
isotope ratios of plant water to those of soil water at each
depth class. The mean proportional water uptake from each
of the three soil depths produced by IsoSource was used in
the statistical analyses. For analyzing seasonal water source
use, water source use data was divided into three periods
(early and late dry season and wet season). The early dry
season was from November to February, the late dry season
was from March to May, and the wet season was from June
to October.
We considered two potential drivers of plant water
uptake along the soil profile: (1) that imposed by water
availability along the soil profile and (2) that imposed by
root distribution along the soil profile. These two drivers
likely interact with each other as the presence of roots in a
particular soil layer may lower water availability at that
depth, especially in the shallow soil where roots are more
abundant. Also, the presence of moisture in a particular soil
depth may induce root proliferation. However, we were not
Copyright © 2014 John Wiley & Sons, Ltd.
able to examine these interactions explicitly here. Consid-
ering the soil depth classes (0–20, 20–50, and 50–150 cm),
the distribution of water availability and root abundance
with soil depth had contrasting profiles. We designated the
potential water source use based on water availability
simply as the proportion of plant-available water in each
soil depth. While the potential water source use based on
root distribution was the proportional distribution of live
roots along the soil profile.
Statistical analysis
All statistical tests were conducted in R (R Core Team,
2013). In the defoliation experiment, paired t-tests of δ2H
and δ18O values of stem water were performed between the
control (non-defoliated) branches and defoliated branches
from the same tree for each collection time.
For the field water source use studies, Shapiro–Wilk test
of normality and Levene’s test of homogeneity of variance
were performed to test whether the data was normal and
homogenous. When the data were not normal and
homogeneous, the data were rank transformed. Two sets
of two-way analyses of variance (ANOVAs) compared
proportional water uptake in each soil depth to determine
how proportional water uptake differed with species, sites,
and seasons. The first set of two-way ANOVAs was
performed on each soil depth separately and compared
season and species to test if the proportional water use of
each species changed between the seasons (early dry, late
dry, and wet seasons). In this set of ANOVAs, the three
sites were combined to look at seasonal differences across
the southern ridge sandhill region. In the second set of two-
way ANOVAs was performed in each season on each soil
depth and compared site and species to test whether the five
species differed in water uptake within each site and if each
of the five species differed in water uptake between
different seasons at the same site.
Chi-squared tests were performed to compare observed
proportional water uptake with expected proportional water
uptake based on the distribution of available water in the
soil profile or based on fine root distribution with depth.
Yate’s correction for continuity was used for all Χ2 tests
(Zar, 1999). Separate Χ2 tests were performed for each
species during the early dry, late dry, and wet seasons.
RESULTS
Defoliation experiment
Artificial defoliation of branches resulted in the isotopic
enrichment of stem water. The isotopic separations (Δ18O
and Δ2H ± standard error), 27 days after defoliation, were
3.6 ± 0.3 and 21 ± 4‰, respectively (P < 0.001; Figure 3).
Twenty-seven days after defoliating Q. virginiana
branches, buds were swelling and some small new leaves
Ecohydrol. (2014)
 P. Z. ELLSWORTH AND L. S. L. STERNBERG
Figure 3. The effect of artificial defoliation of Quercus virginiana branches
on isotopic separation. Branches were defoliated on day 0. Isotopic
separation is the mean of the δ values of water from defoliated stem minus
those of the control stems from the same tree. Error bars are standard errors
of the means. Paired t-tests were conducted to calculate if the δ values of
stem water from control branches were significantly different from those of
defoliated branches on the same trees. One, two, or three asterisks show
significance level at P < 0.05, 0.01, and 0.001, respectively.
had emerged from buds. New leaf growth after 34 days was
still slight with new leaves barely emerging, and Δ18O and
Δ2H were slightly lower than the previous week (2.9 ± 0.3
and 16 ± 3‰, respectively; P < 0.01; Figure 3). On day 57,
the new leaves were about two thirds of the mature leaf
size, and the mean Δ18O and Δ2H were greater than zero,
though only mean Δ18O was significantly different from
zero (P < 0.05). The new leaves on the defoliated branches
were approximately full size in the last two collections at
68 and 92 days, and Δ18O and Δ2H were not significantly
different from zero (Figure 3). When the δ2H values were
regressed on the δ18O values of the stem water from
defoliated branches that differed significantly in δ18O and
δ2H from those of the control branches, the slope was 5.6,
which is less than the slope of 8 of meteoric waters and
typical of evaporative enrichment.
Leaf phenology and stem water isotopic composition
Isotopic variation in water through time was relatively
similar among species at each site, except that the
deciduous species (C. floridana and Q. laevis) had higher
δ18O and δ2H values during leafless periods of the year
(December–March; Figure 4). C. floridana dropped its
leaves in late November, and Q. laevis dropped its leaves in
mid-December. Although the timing of leaf drop did not
Copyright © 2014 John Wiley & Sons, Ltd.
differ much within each site, the timing of deciduous leaf
flush differed among sites because of the location of
individual plants with respect to topography. Low areas
were subject to lower temperatures from cold air drainage
than the crest of a hill, so deciduous plants growing at the
crest of the hill leafed out earlier than those at the base of
the hill. Sandhill Long Unburned 1 and Sandhill Recently
Burned, located at a high point in the sandhill, grew new
leaves in late January through early February for Q. laevis
and late February through mid-March for C. floridana.
Located at the base of the hill, Sandhill Long Unburned 2
did not begin to leaf out until early March for Q. laevis and
late March to early April for C. floridana. Leaf lifespan
depended on the species and location of the individual
plants with respect to topography, and was 9.5–10.5 and
7.5–8.5 months for Q. laevis and C. floridana, respectively.
Water source use
In light of the results from our defoliation experiment, stem
water δ2H and δ18O values of leafless deciduous woody
plants were not included in the analysis of water source
use. When proportional water uptake was compared
between species and sites in each season, site main effect
was significant in the early and late dry season for all soil
depths (Table I). In the early dry season, SHLU 2 had the
lowest deep water use and conversely the highest shallow
water use. Also, in the early dry season, SHLU 2 had
proportionally the least amount of available deep water of
the three sites. In the late dry season, the two most recently
burned sites, SHRB and SHLU 2, had shifted to
proportionally shallow water source use, while SHLU 1
was still taking about 50% of its water from the deep soil
(Table III). Although some differences existed between
sites, each species did not differ in its water use of each soil
depth among the three sites.
The species main effect was significant at all depths in
both the early and late seasons except 20–50 cm in the late
dry season (Table I). Within each site, the five species did
not differ in the proportional water use at each soil depth,
except at SHLU 1. At SHLU 1, L. ferruginea proportion-
ally took up more water from the shallow soil and less
water from 50 to 150 cm in the early and late dry season
than Q. geminata (Table II). L. ferruginea at SHLU 1 was
driving the significant species main effect throughout
the early and late dry season and the interaction between
species and site in the shallow soil (0–20 cm) in the
late dry season. Considering that the only two species
(L. ferruginea and Q. geminata) that differed in water
source use were two evergreen species, leaf phenology was
not a differentiating factor in water source use.
Depth of water uptake shifted to deeper soil water during
the early dry season (Table III). The interaction between
season and species at 20–50 and 50–150 cm were
Ecohydrol. (2014)
 DECIDUOUS AND EVERGREEN WATER UPTAKE

18 2
Figure 4. Mean δ O of stem water (a, b, c) and mean δ H of stem water (d, e, f) from May 2008 through May 2009. The three sites are (a, d) Sandhill
Recently Burned, (b, e) Sandhill Long Unburned 1, and (c, f) Sandhill Long Unburned 2. Error bars represent the standard error of the mean. Dotted lines
represent deciduous species, and solid lines represent evergreen species. Solid bar above the x-axis represents the duration of the dry season. The shaded
region of each graph represents the period of time when deciduous woody plants were leafless. Note: Leafless period is less than the entire dry season.

significant, and only L. ferruginea of the five species
differed in seasonal water uptake pattern from the other
species. All species except L. ferruginea took up 9 and
72% of their water from 0 to 20 and 50 to 150 cm,
respectively, but L. ferruginea used more shallow soil
water and less deep water, taking up 20 and 53% of its
water from 0 to 20 and 50 to 150 cm, respectively
(Table III). In the late dry and wet seasons, the proportion
of deep soil water uptake decreased and only provided 30–
32% of all water uptake, while the shallow soil water
uptake increased to 34–40% of water uptake. This shift to
the top 0–20 cm was most dramatic in L. ferruginea during
the late dry season when 50% of water uptake was in the
top 20 cm. The deep soil over the entire year for all species
contributed a mean 39% for all species compared to 33%
taken up from the shallow soil.
The pool size of available water at each soil depth was
inversely proportional to the distribution of fine roots with
soil depth. In the early dry season, 0 to 20 cm had
proportionally the least amount of water available in the top
150 cm (only ~8% of the total water available at the top
150 cm; Figure 5). This was followed by a gradual
proportional increase during the late dry season (~10%) and
reaching the highest amount during the wet season (~13%).
The bulk of water availability, however, regardless of season
occurred between 50 and 100 cm (61–78%). The mid soil
depth proportional water content (20–50 cm) was intermedi-
ate between the shallow and deep soil layers (15–28%). In
contrast to proportional water availability at each depth, the
bulk of the roots were found at the top 20 cm of the soil profile
(Figure 5) with the lower depth intervals (20–50 cm and
50–150 cm) having similar root abundance.
During the early dry season, observed water source use
of all species except L. ferruginea was not significantly
different than the expected water source use based on the
distribution of available water in the soil profile, but

Copyright © 2014 John Wiley & Sons, Ltd. Ecohydrol. (2014)

 P. Z. ELLSWORTH AND L. S. L. STERNBERG

Table I. Two-factor ANOVA table with main effects, site and availability proportional to the other soil depths (Table IV).
species, comparing the use of each water source: 0–20, 20–50, During the wet season, clearly plant water uptake along the
and 50–150 cm. These ANOVAs were conducted to determine if
the five species differed from each other within each season and soil profile was dictated by root distribution and not the
among sites. Separate two-factor ANOVAs were calculated at distribution of available soil moisture (Table IV; Figure 5).
each depth and in each season for a total of 9 ANOVAs. The
seasons were early dry (November–February), late dry (March–
May), and wet season (June–October). Evergreen species are Q. DISCUSSION
geminata, Q. myrtifolia, and Lyonia ferruginea, and the deciduous
species are Carya floridana and Quercus laevis. Level of Effect of leaflessness on stem water δ2H and δ18O
significance is represented by ‘ns’ or one, two, or three
asterisks, where ns, *, **, and *** represent P values of >0.05, The defoliation experiment revealed that evaporation as the
<0.05, <0.01, and <0.001, respectively. result of leaflessness enriched stem water isotopically.
Although stems were suberized and evaporation from the
Site Species Site × species
stem surface is minimal (Schönherr, 1982), evaporation
Early dry season over a period of 1 month was sufficient to isotopically
0–20 cm ** *** ns enrich stem water (Figure 3). The presence of newly
20–50 cm ** ** ns formed leaves did not increase transpiration sufficiently to
50–150 cm * *** ns flush the isotopically enriched water from the sapwood.
Late dry season Only when leaves were nearly at their maximum size,
0–20 cm *** ** **
20–50 cm *** ns ns
transpiration appears to have been sufficient to maintain
50–150 cm *** *** ns high sapflow such that the residence time of water in the
Wet season sapwood was insufficient to allow for evaporative
0–20 cm ns ns ns enrichment of the stem water pool. This was evidenced
20–50 cm ns ns ns by Δ2H and Δ18O being indistinguishable from zero when
50–150 cm * ns * leaves were nearly fully formed.
Consequently, this experiment explained why δ2H and
δ18O values of deciduous woody plants in our field studies
were much higher than those of evergreen shrubs during
Table II. Two-factor ANOVA table with main effects, season and the deciduous species leafless period (Figure 4). The early
species, comparing the use of each water source: 0–20, 20–50,
and 50–150 cm. These ANOVAs were calculated to determine if dry season difference in δ2H and δ18O between deciduous
the five species differed in water source use from each other and and evergreen species cannot represent water uptake from
across seasons. To be able to calculate differences in water uptake an enriched soil water source because leafless deciduous
for the southern ridge sandhill, all three sites were analysed stem water δ2H and δ18O values were often higher than the
together. The seasons were early dry (November–February), late
dry (March–May), and wet season (June–October). Evergreen highest soil water δ2H and δ18O values at the 0- to 20-cm
species are Q. geminata, Q. myrtifolia, and Lyonia ferruginea, depth. We, therefore, did not use the stem water δ2H and
and the deciduous species are Carya floridana and Quercus δ18O values of deciduous plants during their leafless period
laevis. Level of significance is represented by ‘ns’ or one, two, or in the determination of water source use along the soil
three asterisks, where ns, *, **, and *** represent P values of
>0.05, <0.05, <0.01, and <0.001, respectively. profile. Future protocols of studies of this nature must
avoid comparing water source use between deciduous and
Main effects Interactions evergreen plants during the deciduous leafless state,
Depth because the stem water will be isotopically enriched by
Season Species Season × species
evaporation and will not reflect the isotopic composition of
0–20 cm *** *** ns the water source.
20–50 cm ns *** ns
50–150 cm *** *** * Leaf phenology and water source use
Water uptake from the soil profile in any of the three time
periods was not significantly different between deciduous
differed from the expected water source use based on early and evergreen species when we excluded the leafless
dry season root distribution (Figure 5). During the late dry period of deciduous species, except L. ferruginea (Table II).
season, water source use shifted towards a pattern of water However, L. ferruginea like the other four species accessed
source use similar to that based on root distribution and deeper soil water during the early dry season than the other
was significantly different than the expected water source two periods of the year and shallower soil water during the
use based on the distribution of available water (Figure 5). late dry and wet seasons (Table III). Donovan et al. (2000)
During the late dry season, deep water was at its lowest found that the deciduous Q. laevis had greater access to soil

Copyright © 2014 John Wiley & Sons, Ltd. Ecohydrol. (2014)

 DECIDUOUS AND EVERGREEN WATER UPTAKE

Table III. Mean water uptake ± standard deviation of each species at three different soil depths (0–20, 20–50, and 50–150 cm) at three
sites and during three seasons. Each soil depth in each season represents a two-way ANOVA for a total of 9 two-way ANOVAs. Means
of water uptake within each ANOVA followed by the same letter are not significantly different. The species codes are Lyonia
ferruginea (LF), Quercus geminata (QG), Q. myrtifolia (QM), Q. laevis (QL), and Carya floridana (CF). The seasons were early dry
(November–February), late dry (March–May), and wet season (June–October). At soil depths 0–20 and 20–50 cm in wet season, neither
species and season main effects nor interaction were significant. No post hoc test of pairwise comparisons was conducted on these
ANOVAs. The species main effect was not significant at 20–50 cm in the late dry season and at 50–150 cm in the wet season. In these
ANOVAs, Tukey post hoc test was only conducted on the site main effect, so only the mean water uptake for all species were evaluated
for significance. The standard deviation given here is the cumulative standard deviation of the IsoSource calculations and the standard
deviation among the replicates.

Site LF QG QM QL CF All species

Early dry season

Depth 0–20 cm
SHLU 1 0.25 ± 0.14a 0.04 ± 0.2c 0.11 ± 0.05abc 0.12 ± 0.16abc 0.08 ± 0.1abc 0.13 ± 0.12
SHLU 2 0.22 ± 0.13ab 0.14 ± 0.19abc 0.22 ± 0.13ab 0.05 ± 0.14bc 0.15 ± 0.17abc 0.18 ± 0.16
SHRB 0.13 ± 0.18abc 0.08 ± 0.11abc 0.08 ± 0.09bc 0.01 ± 0.13c 0.07 ± 0.09bc 0.09 ± 0.07
Depth 20–50 cm
SHLU 1 0.32 ± 0.12a 0.10 ± 0.29b 0.16 ± 0.11ab 0.19 ± 0.21ab 0.17 ± 0.16ab 0.19 ± 0.19
SHLU 2 0.29 ± 0.18a 0.22 ± 0.27ab 0.26 ± 0.20a 0.12 ± 0.15ab 0.27 ± 0.24ab 0.24 ± 0.23
SHRB 0.22 ± 0.15ab 0.22 ± 0.21ab 0.21 ± 0.20ab 0.04 ± 0.08b 0.15 ± 0.20ab 0.19 ± 0.18
Depth 50–150 cm
SHLU 1 0.44 ± 0.21b 0.85 ± 0.30a 0.73 ± 0.21ab 0.70 ± 0.28ab 0.76 ± 0.28ab 0.68 ± 0.28
SHLU 2 0.50 ± 0.19b 0.65 ± 0.32ab 0.52 ± 0.30ab 0.83 ± 0.21ab 0.58 ± 0.21ab 0.58 ± 0.30
SHRB 0.66 ± 0.22ab 0.70 ± 0.38ab 0.71 ± 0.41ab 0.95 ± 0.14a 0.78 ± 0.34ab 0.72 ± 0.36
Late dry season
Depth 0–20 cm
SHLU 1 0.61 ± 0.21a 0.22 ± 0.17c 0.19 ± 0.19abc 0.36 ± 0.25abc 0.25 ± 0.19abc 0.32 ± 0.20
SHLU 2 0.54 ± 0.20ab 0.39 ± 0.34abc 0.50 ± 0.27ab 0.39 ± 0.29bc 0.56 ± 0.24abc 0.48 ± 0.27
SHRB 0.36 ± 0.20abc 0.33 ± 0.19abc 0.38 ± 0.22bc 0.28 ± 0.20c 0.28 ± 0.26bc 0.32 ± 0.22
Depth 20–50 cm
SHLU 1 0.20 ± 0.26 0.27 ± 0.25 0.29 ± 0.27 0.30 ± 0.28 0.28 ± 0.26 0.26 ± 0.24b
SHLU 2 0.31 ± 0.23 0.34 ± 0.30 0.29 ± 0.27 0.33 ± 0.31 0.25 ± 0.23 0.30 ± 0.28b
SHRB 0.52 ± 0.24 0.48 ± 0.25 0.44 ± 0.27 0.53 ± 0.24 0.52 ± 0.24 0.50 ± 0.23a
Depth 50–150 cm
SHLU 1 0.20 ± 0.28b 0.52 ± 0.13a 0.52 ± 0.29ab 0.34 ± 0.24ab 0.48 ± 0.29ab 0.41 ± 0.25
SHLU 2 0.16 ± 0.16b 0.26 ± 0.16ab 0.21 ± 0.21b 0.28 ± 0.23ab 0.21 ± 0.18b 0.22 ± 0.18
SHRB 0.12 ± 0.16b 0.19 ± 0.12ab 0.19 ± 0.17b 0.19 ± 0.18ab 0.19 ± 0.17b 0.18 ± 0.16
Wet season
Depth 0–20 cm
SHLU 1 0.48 ± 0.22 0.30 ± 0.16 0.29 ± 0.18 0.45 ± 019 0.45 ± 0.20 0.39 ± 0.19
SHLU 2 0.47 ± 0.26 0.38 ± 0.20 0.36 ± 0.25 0.32 ± 0.24 0.47 ± 0.21 0.39 ± 0.23
SHRB 0.36 ± 0.27 0.31 ± 0.25 0.29 ± 0.26 0.27 ± 0.22 0.30 ± 0.23 0.35 ± 0.25
Depth 20–50 cm
SHLU 1 0.23 ± 0.22 0.30 ± 0.18 0.45 ± 0.19 0.31 ± 0.16 0.24 ± 0.18 0.31 ± 0.18
SHLU 2 0.31 ± 0.27 0.40 ± 0.20 0.38 ± 0.29 0.25 ± 0.23 0.23 ± 0.21 0.32 ± 0.24
SHRB 0.27 ± 0.25 0.33 ± 0.33 0.27 ± 0.25 0.28 ± 0.26 0.29 ± 0.26 0.29 ± 0.27
Depth 50–150 cm
SHLU 1 0.30 ± 0.23 0.41 ± 0.26 0.26 ± 0.20 0.24 ± 0.29 0.30 ± 0.27 0.30 ± 0.21b
SHLU 2 0.26 ± 0.22 0.22 ± 0.18 0.27 ± 0.32 0.43 ± 0.36 0.30 ± 0.21 0.37 ± 0.23a
SHRB 0.38 ± 0.24 0.36 ± 0.26 0.25 ± 0.20 0.46 ± 0.29 0.41 ± 0.27 0.30 ± 0.25ab

water than evergreen Quercus species because of less
negative pre-dawn water potentials. In this study, depth of
water uptake did not differ between deciduous and
evergreen species, so differences in adaptation to drought
conditions between deciduous and evergreen oaks were not
because they were accessing different water sources.
According to Markesteijn and Poorter (2009) and
consistent with our first hypothesis, the evergreen species
should be considered ‘drought delayers’ because they
delayed the effects of the drought by accessing deep water
when shallow water was depleted. Deciduous species lost
their leaves in the early dry season, but leafed out in the
late dry season when the drought conditions can be most
severe due to low water moisture and high vapor pressure
deficit (Weekley et al., 2007; Saha et al., 2008). Contrary
to our second hypothesis, deciduous species only avoided
the first half of the dry season and used a ‘drought delayer’
strategy for the last half of the dry season. In other words,

Copyright © 2014 John Wiley & Sons, Ltd. Ecohydrol. (2014)

 P. Z. ELLSWORTH AND L. S. L. STERNBERG

Table IV. Chi-squared (Χ2) tests comparing water source use of each species to expected water source use based on the proportion of
available water in the soil profile that is found in each soil layer or based on distribution of roots with depth. Χ2 tests were performed
separately on each species in each season. The seasons were early dry (November–February), late dry (March–May), and wet season
(June–October). Level of significance is represented by ‘ns’ or one, two, or three asterisks, where ns, *, **, and *** represent P values
of >0.05, <0.05, <0.01, and <0.001, respectively.

Season Species

Evergreen Deciduous

Lyonia ferruginea Quercus geminata Quercus myrtifolia Quercus laevis Carya floridana

Expected water use based on proportional distribution of available water in the soil profile
Early dry *** ns ns ns ns
Late dry *** *** *** *** ***
Wet *** *** *** *** ***
Expected water use based on proportional root distribution with depth
Early dry ** *** *** *** ***
Late dry ns ns ns ns ns
Wet ns ns ns ns ns

when deciduous woody species had leaves, they accessed
principally deep water similar to that of evergreen plants.
Deciduousness may improve water status, so that they can
flush new leaves in late dry season as has been found in the
New World dry tropical forests (Reich and Borchert, 1982;
Reich and Borchert, 1984; Borchert et al., 2002; Borchert
et al., 2005).
Differences in water uptake based on site characteristics
Considering that climatic conditions and water inputs were
the same for all sites, time since fire was the dominant
driver for inter-site differences. Time since fire affects plant
community composition, increases plant height, decreases
understory plant density, and increases water retention in
the shallow soil because of the litter accumulation (Gholz
and Fisher, 1982; Menges et al., 1993; Abrahamson and
Abrahamson, 1996). All of which can change soil water
availability or plant water demands and can potentially
change water source use. At SHRB, the soil was the most
exposed of the three sites and had very little litter layer. Not
surprisingly, shallow soil water availability and shallow
water uptake were the lowest at SHRB in the early dry
season. Similarly, SHLU 2 had the most developed litter
layer and retained a greater proportion of available water in
the top 50 cm. Consequently, plant uptake was higher in
the top 50 cm than at the other two sites.
In the late dry season, water uptake behavior and water
availability no longer were closely coupled as in the early
dry season, with exception of shallow soil use at SHLU 2.
Like in the early dry season, SHLU 2 continued to have
more water available in the shallow soil proportionally and
uptake from the shallow soil was the highest of the three
sites. However, use of the mid and deep soil layers did not
reflect a shift in water availability. Instead plants at all sites
shifted to shallower soils for water uptake. At this time,
evergreen and deciduous species flushed new leaves and
increased production of fine roots (Ellsworth, 2012).
Possibly, fine root production and new leaves were in
preparation for increased water and nutrient availability in
the shallow soil that comes with the wet season rains
(Bucci et al., 2009).
L. ferruginea utilized the most shallow water and
conversely least deep water of all the species at SHLU 1
only. SHLU 1 had the longest time since fire of the three
sites. As stated above, litter accumulates and the organic
matter content of the shallow soil increases as fire
frequency and time since fire increases. This increased
reliance on the shallow soil for water uptake may be
because L. ferruginea is an ericoid mycorrhizal species.
Ericoid mycorrhiza have saprophytic qualities such as
producing enzymes that break down organic compounds,
which allows for increased nutrient uptake from organic
matter (Cairney and Burke, 1998; Cairney and Ashford,
2002; Read et al., 2004). Also, ericoid plants form ‘hair
roots’ being less than 50 μm in the first terminal order and
several times smaller than the ectomycorrhizal roots of the
other study species (Wells and Eissenstat, 2001; Guo et al.,
2008; Valenzuela-Estrada et al., 2008). Possibly, having hair
roots and the reliance on the litter layer for nutrients
increased water uptake from the shallow soil as well, leading
a greater use of the shallow soil for water uptake throughout
the year than the other ectomycorrhizal study species.
Shifts in water source use
Dry season water source use along the soil profile followed
the distribution of available water in the soil profile and not
the root distribution (Figure 5). High evaporation and low
rainfall quickly dry the shallow soil, leaving the deep soil
as the source of most available soil water (Weekley et al.,
2007). In contrast, the deep soil (50–150 cm) had only 33%

Copyright © 2014 John Wiley & Sons, Ltd. Ecohydrol. (2014)

 DECIDUOUS AND EVERGREEN WATER UPTAKE
Figure 5. Water source use in the early dry (A), late dry (B), and wet
seasons (C). The axes represent the proportion of water uptake from each
of the three soil depths. Large black squares represent expected water
source use based on proportion of fine roots located in each depth. Large
black triangles represent expected water source use based on proportion of
available soil water in each depth. Small open circles represent water
source use of each species at each site. Water source use of the five study
species was measured three different times during each of the three
seasons for a total of 15 points for each season.
of the total roots observed at the top 150 cm of the soil
profile and a root density 6.25 times less than the root
density at 0–20 cm (Ellsworth, 2012), yet it provided most
of the water transpired (Figure 5). As has been shown also
Copyright © 2014 John Wiley & Sons, Ltd.
in other studies, the presence of roots does not indicate
their function (Jackson et al., 1999b). Deep roots contribute
to drought tolerance by increasing the total soil volume that
plants can exploit, making the contribution of the relatively
few deep roots large when shallow soil water availability is
low (Garwood and Sinclair, 1979; Paz, 2003; Slot and
Poorter, 2007; Poorter and Markesteijn, 2008; Markesteijn
and Poorter, 2009).
During the late dry and wet seasons, water source use
followed the root distribution profile and not the availabil-
ity of soil water along the profile (Figure 5). In the wet
season, water was plentiful and more evenly distributed
along the soil profile, so water uptake could be at
maximum rate and limited by root distribution and not
water availability (Yadav et al., 2009). This water uptake
pattern along the soil profile being proportional to the root
distribution was also observed in another study of Malus
domestica where 70% of the roots and water uptake were in
the top 40 cm (Green and Clothier, 1999). Although the top
20 cm of the sandy soil in this site does not contain a large
storage of available water to meet transpiration demand, the
frequent wet season rains are more important to maintain
high water availability than water storage capacity (Guswa,
2010). What is not clear is why woody plants shift water
uptake towards the shallow soil in the late dry season when
water availability in the shallow soil is still low. We would
suggest that as plant water uptake dries the deeper soil
layers, the distribution of water along the soil profile
becomes more even. This even distribution of soil moisture
along the soil profile, regardless of quantity, would favor
root distribution as a determinant of water uptake.
Although plants shifted their water uptake along the soil
profile, deep water was the predominant water source
throughout the year, contributing 39% of annual water
uptake from 50 to 150 cm and 67% from 20 to 150 cm. The
nearly pure sand substrate may have low water retention,
but soil volume compensates in part for low water content
by providing a relatively permanent deep soil water
reservoir. This deep water reservoir is important in the
wet season because low water retention and short rainless
periods lead to periods of shallow soil water unavailability.
Deep roots must take up water from the deep soil to
supplement the intermittently available shallow soil water,
so that woody plants have a greater and more continual
supply of water for transpiration.
CONCLUSIONS
We observed here that water source use changes inter- and
intra-seasonally. Therefore, single measurements cannot
capture the natural variation found within and between
seasons. Most water source use studies measured water
source use either a single time or in one case twice in
Ecohydrol. (2014)
 P. Z. ELLSWORTH AND L. S. L. STERNBERG

consecutive dry seasons (Jackson et al., 1995; Schulze Table AI. (Continued )
et al., 1996; Jackson et al., 1999a; Stratton et al., 2000; Parameters ss df ms F P
Querejeta et al., 2007; Hasselquist et al., 2010). These
studies provide information on water source use at one Late dry season
point in time, but it is difficult to infer how depth of water 0–20 cm Site 15.55 2 7.773 8.602 <0.001
uptake changes with the natural seasonal variability of Species 14.84 4 3.711 4.106 <0.01
Site × species 19.93 8 2.492 2.757 <0.01
water availability and leaf phenology.
Residuals 170.8 189 0.904
Analyzing the δ2H and δ18O values of leafless or newly Total
leafed deciduous species can lead to errors in water source 20–50 cm Site 43.63 2 21.677 18.059 <0.001
use calculations because of evaporative enrichment of stem Species 4.63 4 1.157 0.964 0.43
water. Our intensive measurements of deciduous (when Site × species 4.39 8 0.549 0.458 0.89
fully leafed out) and evergreen species indicate that at this Residuals 226.87 189 1.2
Total
site evergreen species are drought delayers, while decid- 50–150 cm Site 31.43 2 15.715 22.21 <0.001
uous species shift from early dry season drought avoidance Species 14.46 4 3.616 5.11 <0.001
to late dry season drought delayers. In the early dry season, Site × species 9.45 8 1.181 1.67 0.11
edaphic and climatic factors controlling shallow soil water Residuals 133.73 189 0.708
availability were the principal drivers determining water Total
source use, resulting in the deep soil being an important Wet season
0–20 cm Site 0.55 2 0.2739 0.27 0.76
water source throughout the year. During the late dry Species 5.46 4 1.3643 1.344 0.26
season and the wet season, the principal driver in Site × species 12.7 8 1.5869 1.563 0.14
determining water source use shifted from the distribution Residuals 186.84 184 1.0154
of available water to the distribution of fine roots. In other Total
words, the abundance and evenness of water available 20–50 cm Site 0.97 2 0.4866 0.439 0.65
throughout the soil profile lead to water uptake from all Species 9.94 4 2.4842 2.239 0.07
Site × species 7.91 8 0.9885 0.891 0.53
depths being proportional to root abundance. Residuals 204.13 184 1.1094
Total
50–150 cm Site 7.32 2 3.66 4.178 <0.05
APPENDIX Species 6.75 4 1.686 1.925 0.11
Site × species 16.31 8 2.039 2.327 <0.05
Table AI. Two-factor ANOVA table with main effects, site and Residuals 161.2 184 0.876
species, comparing the use of each water source: 0–20, 20–50, Total
and 50–150 cm. These ANOVAs were conducted to determine if
the five species differed from each other within each season and
among sites. Separate two-factor ANOVAs were calculated at
each depth and in each season for a total of 9 ANOVAs. The
seasons were early dry (November–February), late dry (March– Table AII. Two-factor ANOVA table with main effects, season
May), and wet season (June–October). Evergreen species are Q. and species, comparing the use of each water source: 0–20,
geminata, Q. myrtifolia, and Lyonia ferruginea, and the deciduous 20–50, and 50–150 cm. These ANOVAs were calculated to
species are Carya floridana and Quercus laevis. P values in bold determine if the five species differed in water source use from each
are significant. other and across seasons. To be able to calculate differences in water
uptake for the southern ridge sandhill, all three sites were analysed
Parameters ss df ms F P together. The seasons were early dry (November–February), late dry
(March–May), and wet season (June–October). Evergreen species
Early dry season are Q. geminata, Q. myrtifolia, and Lyonia ferruginea, and the
0–20 cm Site 9.88 2 4.938 6.739 <0.01 deciduous species are Carya floridana and Quercus laevis. P values
Species 18.22 4 4.556 6.218 <0.001 in bold are significant.
Site × species 11.34 8 1.418 1.935 0.06
Residuals 109.91 150 0.733 Parameters ss df ms F P
Total
20–50 cm Site 14.15 2 7.075 6.392 <0.01 0–20 cm Species 21.6 4 5.41 6.627 <0.001
Species 18.12 4 4.53 4.092 <0.01 Season 118.5 2 59.25 72.556 <0.001
Site × species 16.63 8 2.079 1.878 0.07 Species × season 3.9 8 0.48 0.590 0.79
Residuals 166.04 150 1.107 Residuals 451.6 553 0.82
Total Total 595.6 599
50–150 cm Site 6.3 2 3.15 3.832 <0.05 20–50 cm Species 1.1 4 0.27 0.252 0.91
Species 20.68 4 5.17 6.29 <0.001 Season 64.1 2 32.04 29.42 <0.001
Site × species 12.01 8 1.501 1.826 0.08 Species × season 27.4 8 3.43 3.146 <0.01
Residuals 123.29 150 0.822 Residuals 602.2 553 1.09
Total Total 694.8 599
(Continues) (Continues)

Copyright © 2014 John Wiley & Sons, Ltd. Ecohydrol. (2014)

 DECIDUOUS AND EVERGREEN WATER UPTAKE

Table AII. (Continued ) Borchert R, Robertson K, Schwartz MD, Williams-Linera G. 2005.

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of Biometeorology 50: 57–65.
Bowman DMJS, Prior LD. 2005. Turner Review No. 10 Why do
50–150 cm Species 13.6 4 3.39 5.452 <0.001 evergreen trees dominate the Australian seasonal tropics? Australian
Season 177.2 2 88.62 142.327 <0.001 Journal of Botany 53: 379–399.
Species × season 11.0 8 1.38 2.213 <0.05 Bucci SJ, Scholz FG, Goldstein G, Meinzer FC, Arce ME. 2009. Soil
Residuals 344.3 553 0.62 water availability and rooting depth as determinants of hydraulic
Total 546.1 599 architecture of Patagonian woody species. Oecologia 160: 631–641.
Cairney JWG, Ashford AE. 2002. Tansley Review No. 135. Biology of
Mycorrhizal Associations of Epacrids (Ericaceae). New Phytologist
154: 305–326.
Cairney JWG, Burke R. 1998. Extracellular enzyme activities of the
Table AIII. Chi-squared (Χ2) tests comparing water source use of ericoid mycorrhizal endophyte Hymenoscyphus ericae (Read) Korf &
each species to expected water source use based on the proportion Kernan: their likely roles in decomposition of dead plant tissue in soil.
of available water in the soil profile that is found in each soil layer Plant and Soil 205: 181–192.
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