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may contribute to the mechanism of canopy decline. al. 1986). The soils are derived from Kula series lava flows of
The chemical environment of waterlogged soils can affect approximately 410,000 years (Macdonald et al. 1983) and are
the physiological functioning of plants and overall plant struc- classified as Inceptisols (Aquertic Humitropept; Schuur
ture (Zimmermann and Brown 1971, Kozlowski 1976, Chapin 1999). Waterlogging has been a major factor in the develop-
1991). Oxygen deficiency associated with soil waterlogging ment of these soils, resulting in an organic horizon in the top
may cause plants to exhibit physiological stress responses 20 cm subtended by a waterlogged mineral horizon where
including reduced transpiration and stomatal closure (Koz- plant roots are absent or very rare (Kitayama and
lowski and Pallardy 1984, Chapin 1991, Sojka 1992). Water- Mueller-Dombois 1994a). These soils are high in extractable
logged soil may also be associated with thicker leaves and less aluminum, acidic and hypoxic (Kitayama and Mueller-
negative δ13C values signifying stomatal closure and greater Dombois 1994a, Kitayama et al. 1997). The topography con-
water-use efficiency (Meinzer et al. 1992). Avoidance of an- sists of narrow, flat to moderately sloped ridges formed by re-
oxic horizons through the production of aboveground roots is sistant portions of the shield volcano, dissected by steep
a common strategy of plants adapted to waterlogged soils ravines. Soil waterlogging varies with topography and is most
(Armstrong et al. 1991) and has been reported in high precipi- evident in level areas creating a mosaic of depressions and
tation montane forests (Gill 1969). However, aboveground slopes on broad ridges.
and shallow rooting may increase susceptibility to drought Moderately sloped areas support communities dominated
(Holbrook and Putz 1996), especially in cloud forest vegeta- by a closed canopy of M. polymorpha, which is 11–13 m tall,
tion adapted to humid conditions (Jane and Green 1985). and an open sub-canopy shrub and tree fern layer, which is
Most estimates of transpiration from tropical montane for- 1–3 m tall. Level, poorly drained areas support open canopy
ests are derived from top-down approaches such as water bal- M. polymorpha stands of approximately the same height, but
ance calculations (Bruijnzeel and Proctor 1993). Such results with smaller individual tree crowns, and an understory domi-
are difficult to extrapolate to smaller scales because they do nated by the sedge, Carex alligata Boott, an indicator of wet-
not account for the contributions of individual plant species or land conditions (Kitayama and Mueller-Dombois 1994a,
functional groups that can be important for comparing the ef- 1994b). Six 10 × 10 m plots were chosen on the basis of slope
fects of vegetation on hydrological cycles (McNaughton and angle (Table 1). Level and sloped sites had canopy coverage of
Jarvis 1983). Alternatively, bottom-up models are determinis- approximately 60 and 90%, respectively, as determined with a
tic, process-based approaches that predict how systems may spherical densiometer.
function at large scales, based on investigations of small spa-
tial and short temporal scales (Jarvis 1993). Hawaiian mont- Micrometeorological measurements
ane forests are optimum systems for investigating the utility of Climatic conditions at the study sites were monitored at can-
bottom-up models because the single canopy species simpli- opy height from September 16, 1996 until February 10, 1997.
fies scaling from leaves or whole-plant measures to the canopy Net radiation was measured with a net radiometer (Model Q7,
level. We have examined the effects of waterlogging on tran- Radiation and Energy Balance Systems, Seattle, WA). Rela-
spiration and forest structure by comparing stands growing in tive humidity and air temperature were measured with a
topographically level, waterlogged soils with stands on mod- shielded temperature/humidity sensor (Model HMP35C,
erately sloped, drained soils, to obtain physiological informa- Vaisala, Helsinki, Finland). Windspeed was measured with a
tion on M. polymorpha canopy decline in waterlogged soils. cup anemometer (Model 014A, Met One Instruments, Grants
Specific objectives were to: (1) investigate physiological limi- Pass, OR). Photosynthetic photon flux density (PPFD) was
tation of water use by trees in waterlogged soil; (2) determine measured with a quantum sensor (Model Quantum, Li-Cor,
how canopy water fluxes are affected by differences in LAI
and stand structure; (3) develop a model that predicts M.
polymorpha canopy transpiration based on climatic parame-
ters; and (4) examine the effects of tree size and rooting me- Table 1. Stand characteristics of pairs of level and sloped study sites.
dium on stomatal conductance. Soil water is expressed as the temporal mean of three measurement
periods.
Inc., Lincoln, NE). Data were recorded every 5 s and averaged opy at these sites is sparse, individual shoots of clustered
every 10 min with a data logger (Model 21X, Campbell Scien- leaves could be identified consistently, and thus errors in leaf
tific, Logan, UT). Potential evapotranspiration (PET) was cal- area determination are likely to be small. The LAI of the
culated by the Penman equation (Penman 1948). understory sedge C. alligata was estimated with a portable
area meter (Model 3100, Li-Cor, Inc.) by destructively har-
Redox potential and soil water vesting four randomly selected 0.5 × 0.5 m plots along a 20-m
transect in an area of continuous C. alligata groundcover.
Soil redox potentials were measured with platinum electrodes,
Canopy sun leaves were obtained from the three tallest mea-
a calomel reference electrode and a portable mV meter (Model
surement trees in each plot with a telescopic pruning hook.
290A, Orion, Inc., Boston, MA). Electrodes were inserted to
Leaf area was determined with a portable area meter (Li-Cor,
10- and 40-cm depths at 11 randomly selected points within
Inc.). Leaves were then dried in an oven at 70 °C to constant
each 10 × 10 m plot and allowed to equilibrate for 45 min be-
weight for the determination of leaf mass per area (LMA). The
fore measurements. Values were corrected to a standard hy-
dried leaves were finely ground and a subsample was sent to
drogen electrode (+244 mV) and to a pH of 7 (–59 mV per pH
the Agricultural Diagnostic Service Center, University of Ha-
point below 7; Bohn et al. 1985) from a soil pH of 3.7
waii at Manoa for chemical analysis of leaf nutrient concentra-
(Kitayama and Mueller-Dombois 1994a). Soil water was mea-
tions. An additional subsample was sent to the University of
sured gravimetrically three times throughout the measurement
California at Berkeley where the relative abundances of 13C
period. In each 10 × 10 m plot, six soil cores were taken to a
and 12C were analyzed by mass spectrometry.
depth of 10 cm, weighed, then dried in an oven at 70 °C to con-
stant weight. Water content of the soil was expressed in grams
of water per gram of oven-dried soil (g g –1). Transpiration and stomatal conductance
Between September 1996 and February 1997, transpiration
Stand structure, leaf area index and leaf chemistry was measured by a paired plot technique where simultaneous
The diameters at 1.4 m height (DBH) of all M. polymorpha measurements of the four measurement trees in one level plot
stems ≥ 2 cm were measured in each 10 × 10 m plot. To evalu- and one sloped plot were allowed to run for 15–20 days. The
ate the degree of soil avoidance by roots, the rooting habits of 5 days with highest evaporative demand from each measure-
M. polymorpha individuals were assessed by observing ment period were used to compare both sets of measurement
where, in relation to the soil surface, rooting initiated. A plant trees. Transpiration was estimated from sap flow measured
was considered to be rooted in the soil if there were adventi- with the constant heating method (Granier 1987). One pair of
tious roots and the shoot to root boundary occurred below 30-mm long probes (Dynamax Inc., Houston, TX) was in-
ground. If the shoot to root boundary occurred above ground, serted into the north-facing side of each measurement tree at
the plant was considered aerially rooted. A four-class system 1.4 m height. Measurements of mean temperature differences
was used to distinguish rooting habit: (1) soil (stem entered along the lengths of the upper, heated probe, and the lower,
soil intact with little or no aboveground rooting); (2) nurse log reference probe were taken every 5 s and averaged every 10
(roots penetrated an aboveground horizontal log); (3) min by a data logger (Model CR10X, Campbell Scientific).
epiphytic (roots were dependent on the bole of another tree for Sap flow velocity was calculated from the temperature differ-
support); and (4) aerial (shoot to root boundary occurred ence between probes by the empirical equations developed by
above the soil surface, either exposed or in aboveground, Granier (1987). Transpiration (E) was determined by multi-
moss-covered root mats). plying sap flow velocity by sapwood area at the height of mea-
In each plot, four trees with a diameter greater than 6 cm surement.
were randomly selected for leaf area and transpiration mea- Sapwood area was determined by dye injection and the dif-
surements. The LAI was estimated by a new technique con- ference in wood color. Toluidine blue dye (Sigma Chemicals,
sisting of counting the number of leaf clumps in photographs St. Louis, MO) was inserted into the bole of the tree at mea-
of individual tree crowns. Because leaf arrangement is tightly surement height. A core was taken with an increment borer
clumped in M. polymorpha, mean leaf area per clump was de- 2 cm above the point of insertion several hours later and sap-
termined and the numbers of clumps observed in photographs wood thickness was observed as the distance from the inner
were used to estimate individual tree leaf area. The relation- bark toward the center of the trunk that was stained blue. Be-
ships between DBH and individual tree leaf area were then cause long periods of low evaporative demand made this tech-
used to predict leaf area for other trees. Leaf area index was nique difficult, sapwood thickness was also determined by the
calculated as the sum of predicted leaf areas (m 2) for trees in a change in color between the lighter, conducting sapwood and
10 × 10 m plot as: LAI = Σ(predicted tree leaf area)/100. the darker heartwood. On trees where both methods were suc-
The canopy photo method for estimating LAI has an advan- cessful, the border of color change between the sapwood and
tage over other methods because it considers leaf clumping. heartwood corresponded well to the extent of stained tissue.
Methods based on gap-fraction analysis (Welles and Norman The relationship between basal area and sapwood area of trees
1990) underestimate LAI when leaf area is spatially clumped in all plots fell on the same line, so a single log–log regression
nonrandomly. In M. polymorpha, this error may be quite large (Sprugel 1983) was used to predict sapwood area from basal
(Herbert and Fownes 1997). Because the M. polymorpha can- area in cm (SA = e–0.18 BA0.92, r 2 = 0.98, n = 22). In all but four
Table 2. Summary of transpiration (E) for 5-day measurement periods comparing level, waterlogged, open-canopy sites with sloped, closed-can-
opy sites. Values of E are actual measurements from four trees per plot for 5 days and were normalized by basal area (EBA ) and leaf area (ELA ) of
measurement trees for comparison. Stand transpiration (Estand) was estimated by multiplying EBA by total stand basal area. Potential
evapotranspiration (PET) was used to normalize Estand for climatic conditions.
sloped topographies. During cloudy conditions, the mean gs of tioning of this ecosystem. At the whole-plant and stand scales,
juveniles on level and sloped sites was 177 ± 24 and 187 ± leaf surface area was the best predictor of plant water use (Fig-
25 mmol m –2 s –1, respectively. The corresponding values dur- ures 5 and 6). This finding supports the idea that LAI is the
ing clear conditions were 233 ± 17 and 219 ± 32 mmol m –2 s –1, most important variable for relating vegetation structure to en-
respectively. Mean gs in larger trees during clear weather ergy and mass exchange because it is a measure of the evapo-
showed a linear decrease with increasing tree height from a rative surface area (Running and Coughlan 1988). In addition,
mean of 339 mmol m –2 s –1 at 0.9 m to a mean of 54 mmol m –2 leaf area appears to be an integrator of processes between the
s –1 at 6.4 m (Figure 7). leaf and stand scales. The success of bottom-up models de-
pends on integrators between different scales, and such rela-
tionships are often revealed by observations at several levels
Discussion of organization. Observations at a single scale can lead to in-
High precipitation inputs, low evaporative demand and local compatible conclusions. For example, differences in ELA
topography combined to form patches of saturated soil that within pairs of sites were much smaller than differences in
were associated with a low proportion of soil rooting, low Estand (Table 2). Without investigating transpiration processes
basal area and low LAI. Differences in canopy water fluxes at the leaf, whole-plant and stand scales, it would be difficult
appeared to be mediated by LAI, and to a lesser degree by to determine that leaf area is the best structural predictor of
crown exposure. Although there were structural differences transpiration at different scales.
between trees on level and sloped sites, such as lower basal One limitation to using leaves as an integrator is that leaves
area and more soil avoidance by roots in level sites, differ-
ences in LAI dominated in controlling the evaporative func-
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