1983, Kozlowski et al. 1991, Vartapetian and
Jackson 1997), including Annona (Zotz et al. 1997)
and Salix (Jackson and Attwood 1996). Salix roots
and stems may also develop hypertrophied lenticels
(Pereira and Kozlowski 1977, Jackson and Attwood
1996) which further facilitate oxygen uptake and
transport in plants.
Structural adaptations can contribute to the
maintenance of stomatal conductance during flood-
ing and its recovery afterward (Kozlowski 1984,
Pezeshki and Chambers 1986, Sojka 1992, McKevlin
Figure 4.
Mean tree crown volumes (6 1 SE) under the high, low, and no flood treatments, at five sampling times, for
four representative species. ‘ High flood; e Low flood; n No flood.
Table 4.
Summary of ANOVA results showing linear and quadratic trends in treatment effects (df 5 2) on crown volume
and stomatal conductance in four species.
Species
Crown volume
Stomatal conductance
df (error)
Linear
Quadratic
df (error)
Linear
Quadratic
F
p
F
p
F
p
F
p
Annona
21
9.77
0.001
4.40
0.025
8
3.59
0.077
0.95
0.427
Bursera
11
6.76
0.012
1.81
0.209
3
2.01
0.280
9.83
0.048
Chrysobalanus
21
7.33
0.004
8.04
0.003
8
7.92
0.013
1.24
0.339
Persea
19
6.27
0.008
7.37
0.004
9
6.42
0.019
0.84
0.463
840
WETLANDS, Volume 26, No. 3, 2006
et al. 1998). This may explain the relatively higher
stomatal conductance values seen throughout the
study in some of the swamp forest species, most
notably Salix, Morella, and to some extent, Annona,
all of which formed extensive adventitious roots
under HF. The less extensive adventitious root
systems that developed in the remaining swamp
species under HF may have resulted in the relatively
lower stomatal conductance values observed in these
species. McKevlin et al. (1998) also reported di-
minished stomatal conductance in flood-tolerant
species growing in saturated soil. The early, pre-
cipitous declines in stomatal conductance shown by
the five upland forest tree species were expected,
although this contrasts with a study by Lopez and
Kursar (1999), who did not observe sharp reductions
in stomatal conductance in three upland tree species
in Panama during 90 days of inundation.
The lower survival and relatively poor growth and
physiological performance seen under HF in the
upland species tested is not surprising, given that
they are not found in regularly inundated sites.
These and many other important tropical species
occurring in upland sites of the region are adapted to
seasonally-dry conditions and commonly inhabit
thin soils that form directly on limestone (Craighead
1971, Tomlinson 1980, Armentano et al. 2002).
Consequently, they are potentially exposed to
seasonal drought, although in southern Florida,
some may be rooted in ground water. Whether the
ability to tolerate or avoid drought among upland
tree species is related to the ability to tolerate shoot
water stress induced by soil anoxia is not certain. In
a study of tropical dry forest trees, Brodribb et al.
(2003) found that Bursera simaruba, a species that
responds to drought in southern Florida by shed-
ding its leaves and avoiding drought, was especially
vulnerable to xylem cavitation (hence reduced water
conductivity). We found this species to be extremely
sensitive to flooding. Specific information on the
drought tolerance of the other upland species in our
study, however, is lacking. Our findings are in
Figure 5.
Mean stomatal conductance (6 1 SE) under the high, low, and no flood treatments, at four sampling times, for
four representative species. ‘ High flood; e Low flood; n No flood.
Jones et al., TREE RESPONSES TO HYDROLOGIC REGIMES
841
marked contrast to a similar study involving
seedlings of three upland tropical tree species
subjected to an experimental flooding regime. Lopez
and Kursar (1999) reported no mortality or visible
leaf damage after 90 days of inundation and
concluded that most tropical tree species are
relatively tolerant of flooding, yet do not become
established in inundated habitats.
The relative flood tolerances of the 12 test species
grown
under
simulated
flooding
regimes
for
25 weeks are roughly related to their observed
distribution along the natural hydrologic gradient
in tree islands of the southern Everglades. Other
studies on flood tolerance in bottomland forest tree
species of the United States have suggested a similar
relationship between flood tolerance and distribu-
tion patterns along flooding gradients (Hosner 1960,
Hosner and Boyce 1962, Dickson et al. 1965, Hook
and Brown 1973, Larson et al. 1981, McKnight et
al.1981, Mitsch and Rust 1984). In this study, flood
tolerance rankings of species under natural field
conditions were inferred by calculating mean water-
level optima for each species from vegetation surveys
in plots on three Shark Slough tree islands; using
a weighted averaging calibration and regression
procedure (Birks et al. 1990), the local hydrologic
regime was projected from long-term water-level
records. Species rankings under natural field condi-
tions differed little from those in the shadehouse
(e.g., Chrysobalanus was less flood-tolerant in the
tree island, Simarouba was more flood-sensitive in
the shadehouse). Age and size of plants, as well as
water quality, factors known to affect flood tolerance
in plants (Gill 1970, Kozlowski et al. 1991), may have
accounted for some of the discrepancies in the
rankings. The results of the few other studies of
flood tolerances in Everglades tree island tree species
(Gunderson et al. 1988, Guerra 1997) reveal similar
rankings of species as this study.
Extrapolated to the natural setting of a tree island,
the results of this study suggest that increasing water
depths and durations may have a beneficial yet
temporary effect on most hardwood hammock
species; prolonged soil surface inundation will hasten
reduction in tree growth and lead to death of these
species. The more flood-intolerant species of the
surrounding swamp forest (Persea, Magnolia, Ilex)
can be expected to respond similarly, although
a delay in the onset of reduced growth and perhaps
death would be expected. We terminated our study
before it was determined how the most flood-tolerant
swamp species (Annona, Salix, Morella) would
respond to increasingly higher and longer flood
waters. This knowledge can allow us to manage
species composition on tree islands with water level
and may allow early warning of flooding stress in tree
islands. With restoration plans under CERP antic-
ipating
modifications
in
hydrologic
conditions
throughout the Everglades, predicting responses of
tree island species to these changes becomes critical.
Knowledge of relative species tolerances, together
with ancillary information such as genotypic varia-
tion (McKevlin et al. 1998), particularly in species
distributed along a soil moisture gradient (Keeley
1979), will become important in selecting suitable
species to include in projects aimed at restoring
destroyed or degraded tree islands and creating new
ones, a CERP objective. For example, Wallace et al.
(1996) assessed flood tolerance and seedling growth
and survival under varying soil conditions and
developed guidelines for the use of nine tree species
in wetland restoration and creation in Florida.
Several of the species reported in our study have
never been evaluated for flood tolerance until now.
ACKNOWLEDGMENTS
This project received funding through the U.S.
Department of the Interior’s Critical Ecosystem
Studies Initiative (CESI). The authors gratefully
acknowledge the assistance of the following Florida
International University staff for their valuable
contributions toward the successful completion of
this study: Josh Walters, Darcy Stockman, Debbie
Nolan, Pablo Ruiz, Hillary Cooley, and Dave Reed.
This is contribution # 331 of the Southeast
Environmental Research Center at Florida Interna-
tional University.
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