Standing crop and aboveground biomass partitioning of a dwarf

mangrove forest in Taylor River Slough, Florida

1,3, 1 1 2
*
C. Coronado-Molina , J.W. Day Jr , E. Reyes and B.C. Perez
1
Department of Oceanography and Coastal Sciences and Coastal Ecology Institute, School of the Coast and
2
the Environment, Louisiana State University, Baton Rouge, LA 70803, USA; National Wetland Research
3
Center, US Geological Survey, Lafayette, LA 70506, USA; Current address: Everglades Division, South
*
Florida Water Management District, 3301 Gun Club Road, West Palm Beach, Florida 33406, USA; Author
for correspondence (e-mail: [email protected]; phone: 11-561-686-8800)
Received 6 April 2000; accepted in revised form 24 February 2003
Key words: Aboveground biomass, Allometric equation, Biomass allocation, Rhizophora mangle
Abstract
The structure and standing crop biomass of a dwarf mangrove forest, located in the salinity transition zone of
Taylor River Slough in the Everglades National Park, were studied. Although the four mangrove species reported
for Florida occurred at the study site, dwarf Rhizophora mangle trees dominated the forest. The structural
characteristics of the mangrove forest were relatively simple: tree height varied from 0.9 to 1.2 meters, and tree
21
density ranged from 7062 to 23 778 stems ha . An allometric relationship was developed to estimate leaf, branch,
prop root, and total aboveground biomass of dwarf Rhizophora mangle trees. Total aboveground biomass and their
components were best estimated as a power function of the crown area times number of prop roots as an
20.5083
independent variable (Y5B 3X ). The allometric equation for each tree component was highly signicant
2
(p ,0.0001), with all r values greater than 0.90. The allometric relationship was used to estimate total
21
aboveground biomass that ranged from 7.9 to 23.2 ton ha . Rhizophora mangle contributed 85% of total standing
crop biomass. Conocarpus erectus, Laguncularia racemosa, and Avicennia germinans contributed the remaining
biomass. Average aboveground biomass allocation was 69% for prop roots, 25% for stem and branches, and 6%
for leaves. This aboveground biomass partitioning pattern, which gives a major role to prop roots that have the
potential to produce an extensive root system, may be an important biological strategy in response to low
phosphorus availability and relatively reduced soils that characterize mangrove forests in South Florida.
Introduction sponses to daily, seasonal, and long-term patterns in
environmental conditions occurring in coastal tropical
Hydrology, nutrient input, soil salinity, and soil type regions. Accordingly, dwarf and hammock forests
are the most important factors that affect the structure located in the Southern Everglades are recognized as
and productivity of mangrove forests (Brown and special sub-types that respond to localized hydro-
Lugo 1982). Since the effects of these factors vary geologic and edaphic conditions (Cintron and Schaef-
over geographical regions, mangrove forests exhibit fer-Novelly 1984).
wide regional and local variation in structural charac- In ecological studies, reliable estimates of biomass
teristics. Based on differences among the above en- and growth rates are essential for estimating total net
vironmental factors, Lugo and Snedaker (1974) de- primary production (Clough and Scott 1989). A com-
veloped a mangrove classication that included six mon non-destructive method for estimating biomass
basic types of mangrove forests: riverine, fringe, of trees or shrubs is to t equations that relate tree
overwash, basin, hammock and dwarf. This classica- biomass to some dimensional characteristics of the
tion identies common patterns of mangrove re- plant. For instance, stem diameter at breast height
157
2004 Kluwer Academic Publishers. Printed in the Netherlands.
Wetlands Ecology and Management 12: 157164, 2004.
(dbh), tree height, and a combination of both variables nate the landscape before reaching Florida Bay (Fig-
have been commonly used (Causton 1985; Day et al. ure 1). Spatially, Taylor Slough can be divided into
1987). For shrubs, other dimensional variables have Upper, Middle, and Lower zones. Freshwater marsh
been used to predict biomass, including the greatest species, such as sawgrass (Cladium jamaicense) and
diameter of the canopy (Uresk et al. 1977), crown spike rush (Eleocharis cellulosa), dominate the upper
diameter axes (Rittehnhouse and Sneva 1977), and zone. The middle zone is a marsh dominated by
height times circumference (Azocar et al. 1981). willow (Salix caroliniana), sawgrass and spike rush.
Allometric relationships between aboveground In the lower zone the freshwater vegetation is re-
biomass and dbh have been reported for Rhizophora placed by mangrove forests, salt barrens, and tropical
mangle, Avicennia germinans, Laguncularia racem- hardwood hammocks. The soil in this site consists of
osa (Golley et al. 1962; Day et al. 1987; Fromard et recently developed calcareous marl overlaying peat
al. 1998), and other mangrove species in Thailand, deposits (Pool et al. 1977).
Malaysia, and Australia (Ong et al. 1982; Putz and
Chan 1986; Clough and Scott 1989; Tam et al. 1995;
Clough et al. 1997). For dwarf R. mangle, the domi- Methods
nant species in our study site, there is no literature on
the optimal dimensional variables that can be used to Forest structure
estimate its biomass. In this paper, we focus on the
structure and standing crop biomass of a dwarf man- Trees with a diameter greater than 2 cm, measured at
grove forest and develop an allometric equation to 60 cm from the ground (D60), were sampled using the
estimate aboveground biomass of a red dwarf man- point centered quarter method (Cottam and Curtis
grove community located in the salinity transition 1956; Pool et al. 1977). Four 75 m long transects
zone of Taylor Slough in the Everglades National perpendicular to the channel edge, and with points 5
Park, South Florida. m apart, were randomly chosen. At each transect,
oristic composition, diameter (D60), total height,
crown diameter and number of prop roots were mea-
Study site sured. Data from each transect were used to estimate
standing crop biomass using the allometric equation
The Everglades salinity transition zone is located in obtained with the methodology described below.
the southern part of Everglades National Park at
258 109 N and 808 459 W. It borders Florida Bay, a Aboveground biomass
large, shallow, sub-tropical embayment, bounded on
the south and east by the Florida Keys (Figure 1). The To estimate biomass of the dwarf red mangroves, ten
region is characterized as sub-tropical savanna with trees ranging from 0.7 to 1.6 m in total height were
distinctive rainy (MayOctober) and dry (November harvested. From each tree, height, diameter (D60),
April) seasons (Hela 1952). During the rainy season, number of prop roots, and crown diameter were
winds are dominated by a southerly sea breeze system recorded, and then all leaves were stripped. Fresh
that promotes the formation of daily convection thun- weights of all tree components were measured in the
derstorms (Chen and Gerber 1990). Tropical storms eld, and a subsample was taken to the laboratory.
and hurricanes also contribute signicant amounts of Subsamples were oven-dried at 70 8C to constant
rainfall during the rainy season. Low rainfall and mild weight and then re-weighed. A dry to wet ratio of
temperatures, associated with the passage of winter each component was obtained, and dry weights of all
cold fronts, characterize the dry season (Chen and tree components were calculated. Because most
Gerber 1990). In NE Florida Bay, tides are semi- shrubs, including dwarf Rhizophora mangle trees,
diurnal with an average range of 30 cm. However, the have several stems at the ground level and no recog-
mean tidal range in the Taylor Slough area is approxi- nizable main stem from which dbh can be measured,
mately 5 cm due to restricted water circulation (Wang biomass can not be calculated using dbh as an in-
et al. 1994). dependent variable. Thus, other variables have been
The study site is located at Taylor Slough, which used including crown diameter axes (Rittehnhouse
drains into an inland basin where several shallow and Sneva 1977), crown volume (Crow 1978), and
ponds and an extensive dwarf mangrove forest domi- height times circumference (Azocar et al. 1981; Et-
158
Figure 1. Everglades Salinity Transition Zone showing Taylor Slough and the basin area dominated by dwarf red R. mangle.
159
tiene 1989). We developed an allometric equation to against the natural logarithm of the variable crown
estimate dwarf Rhizophora mangle biomass using area 3 number of prop roots. The least square line of
several dimensional characteristics of the plant such best t is described as:
as height, crown area, number of prop roots, and a
LnBiomass51.252210.5083
combination of crown area 3 number of prop roots.
3Ln crown area s
Data were natural log transformed and a multiple
3number of prop rootsd
linear analysis was performed using JMP 4.0 (SAS
Institute 2000). A signicance level of p 50.05 was
While the natural log-log equation provides a con-
used to interpret results of all statistical tests.
venient method for estimating the values of the inter-
cept and slope, it is more useful to display the al-
lometric relationship as the power function of the
Results
independent variable. Thus, the allometric equation
that best explained the biomass variability of dwarf
Forest structure
Rhizophora mangle trees was:
21
20.5083 2
Tree density ranged from 7062 to 23 778 stems ha ,
Y5B3X R
2 21
basal area ranged from 2.35 to 6.44 m ha , and tree
5 0.96 p , 0.0001 s d
height varied from 0.9 to 1.2 meters. Rhizophora
where: Y5total above-ground biomass; B53.4980
mangle was the dominant species and accounted for
(natural antilog of intercept in Equation 1); X5
99% of the dominance and 97% of the density.
crown area 3 number of prop roots.
Although L. racemosa and A. germinans individuals
The regression constants (B and X), the correlation
occurred in two of the transects, they were few in
2
coefcient (R ), and p values for each component
number and were dwarfed as well.
(leaves, stem and branches, prop roots, and total
biomass) are summarized in Table 1. Even though the
Above-ground biomass
Ln (crown diameter) variable did not enter the model
when the multiple-linear regression analysis was run,
The allometric relationship that best t our data was
a simple linear model was obtained explaining 89% of
based on two different types of multivariate analysis:
the total aboveground variability. The allometric
stepwise and backward regression. Both models con-
equation was:
tained the same subset of variables: Ln (height), Ln
(crown diameter), Ln (number of prop roots), and Ln
20.8166 2
Y5B3X R
(crown area 3 number of prop roots). This analysis
5 0.89 p , 0.0001 s d
was useful to determine the relative importance of the
variables in explaining the data variance. The newly where: Y5total above-ground biomass; B5
created variable Ln (crown area 3 number of prop 0.84126; X5crown area.
roots) was the rst and only variable to enter both This allometric model can also be used to estimate
models, and it explained the majority of the total the aboveground biomass of dwarf R. mangle. How-
2
biomass variability (R 5 0.96). Ln (height), Ln ever, biomass calculated by this equation was under-
(crown diameter), and Ln (number of prop roots) estimated by 22% compared to that of the former
explained very little of the mangrove biomass vari- equation. Similarly, we estimated the partitioning of
ability, and did not remain in the model. A linear aboveground biomass using the allometric equations
relationship was obtained by plotting dry weight developed for each component (Table 1). According-
Table 1. Linear regression equations used to estimate aboveground biomass.
2
Plant component Intercept (A) Slope (B) R Signicance of F value
Leaf 22.0045 0.5568 0.97 p , 0.0000
Branch 20.6769 0.5751 0.96 p , 0.0001
Prop root 1.1809 0.4806 0.91 p , 0.0000
Total 1.2522 0.5083 0.96 p , 0.0000
The equations are in the form log e Y 5 A 1 B long e X, where Y 5 dry biomass (g), and X 5 log e (crown area 3 number of prop roots).
160
ly, dwarf R. mangle allocated biomass as follows: velopment in South Florida is highest, soil salinity,
prop roots (69%), stem-branches (25%), and leaves total soil nitrogen and phosphorus values averaged 20
21 23 23
(6%). g kg , 2.2 mg cm , and 0.24 mg cm , respectively
(Chen and Twilley 1999). Those results suggest that
soil salinity and soil nitrogen were not limiting forest
Discussion development at our study site. However, low P availa-
bility might be a limiting factor as has been shown for
Few studies have estimated biomass of mangrove other red dwarf mangrove communities in Belize and
species and most of the available estimates are for the Florida where tree growth and development are the
genus Rhizophora (Brown and Lugo 1982; Day et al. poorest among Caribbean mangrove forests (Feller
1987). Ross et al. (2001) developed an allometric 1995; Koch 1997; Chen and Twilley 1999; Feller et
equation for mixed dwarf and fringe red mangroves al. 1999). Additionally, Koch (1997) working in two
from a site at Biscayne Bay, Florida, using DBH 3 red dwarf mangrove forests similar to our study site,
Crown Volume as an independent variable to estimate showed that soil redox potentials were highly reduced
biomass for trees located at fringe and dwarf sites. (2100 to 2200 mV). Thus, it is possible that both low
Average aboveground biomass, estimated by Ross et phosphorus availability and highly reduced soils are
21
al. (2001), was 18.2 t ha . Using our allometric important factors limiting mangrove tree growth and,
equation, aboveground biomass estimation ranged therefore, aboveground biomass accumulation.
21
from 7.9 to 21.2 t ha which is similar to the The partitioning of dry matter between different
biomass of dwarf red mangroves located at Biscayne components of the tree is affected by factors such as
Bay. Thus, we are condent of the biomass estima- age, environmental conditions, forest structure, and
tions obtained using our allometric relationship; how- competitive interactions (Landsberg 1986; Koslowski
ever, care should be taken in applying the allometric et al. 1991). Rhizophora apiculata and R. mangle
equation developed in this study to other mangrove individuals growing in optimal conditions of light and
stands. nutrients allocate more dry matter to stem and branch-
Our aboveground biomass estimation (7.9 to 23.2
t
es at expense of the prop roots (Clough 1992; Lugo
21
ha ) is lower than that of Riverine, Fringe, or Basin and Snedaker 1974). Working with multi-stemmed
mangrove forests located in the Caribbean and Atlan- Rhizophora stylosa trees, located in northwestern
tic regions (Table 2). However, our biomass estima- Australia, Clough et al. (1997) showed that the contri-
tion was slightly higher than that of a similar red bution to stem-branches increased from 25% to 50%
21
mangrove dwarf forest (7.9 t ha ) located in Turkey when mangrove trees increased their dbh from 5 to 25
Point, Florida (Lugo and Snedaker 1974) and rela- cm. In contrast, prop roots contribution decreased
tively higher than that of a dwarf forest dominated by from 40% to 30% when trees increased from 5 to 25
21
Avicennia marina (6.8 t ha ) located in Tuff Crater, cm in dbh. At our study site, dwarf R. mangle showed
New Zealand (Woodroffe 1985). The low above- a similar pattern in which stem-branch dry matter
ground biomass, observed at Tuff Crater, New contribution increased and prop root dry matter con-
Zealand, was directly related to ecological factors tribution decreased (Figure 2). However, the average
such as low nutrient availability and high-reduced soil prop root contribution was higher in dwarf R. mangle
conditions (Woodroffe 1985). Additionally, low (69.2%) than in multi-stemmed R. stylosa (35%)
aboveground biomass has been linked to long hy- (Clough et al. 1997). In contrast, average stem-branch
droperiods (Pool et al. 1977), high soil salinity (Ball contribution was higher in R. stylosa (53%) than in
1988; Lin et al. 1992), sulde accumulation (McKee dwarf R. mangle (25.2%). Clough et al. (1997) argued
1993), reduced soils (Pezeshki et al. 1997) and nu- that the dry matter partitioning pattern in R. stylosa
trient limitation (Boto and Wellington 1983). At our was due to extreme dry conditions that characterized
study site, low aboveground biomass (Table 2) might their study area and that more dry matter was invested
be related to low nutrient resources particularly nitro- in developing an extensive root system to cope with
gen and phosphorus. Accordingly, total soil nitrogen high soil gradient salinities around the roots. At our
23
ranged from 1.9 to 2.9 mg cm , total soil phosphorus study site, soil salinity is not considered a constraint
23 21
ranged from 0.07 to 0.09 mg cm , and interstitial since it ranged from 10 to 25 g kg , falling within
21
soil salinity ranged from 10 to 15 g kg . In contrast, the range in which R. mangle grows optimally (Day et
at Shark River, Florida, where mangrove forest de- al. 1987, 1996; Twilley et al. 1996). In contrast,
161
Table 2. Aboveground biomass, and biomass partition among mangrove forests in North and South America.
Above-ground biomass partitioning
Mangrove Forest Above-ground % Leaves % Prop % Stems and Source
location type biomass roots branches
21
(ton ha )
Florida
Ten Thousand Islands Riverine 135.5 4.7 8.3 87.0 Lugo and Snedaker (1974)
Ten Thousand Islands Fringe 102.5 5.8 18.5 75.8 Lugo and Snedaker (1974)
Ten Thousand Islands Overwash 124.0 5.7 37.6 56.6 Lugo and Snedaker (1974)
Turkey Point Dwarf 7.9 9.1 40.6 50.3 Lugo and Snedaker (1974)
Biscayne Bay Fringe 57.8 13.6 22.1 64.3 Ross et al. (2001)
Byscaine Bay Dwarf 8.8 16.4 17.7 65.9 Ross et al. (2001)
Taylor Slough Dwarf 12.5 5.6 69.2 25.2 This study
Caribbean
Puerto Rico Fringe 63.0 Golley et al. (1962)
Mexico
Terminos Lagoon
Boca Chica Riverine 135.0 29.1 70.9 Day et al. (1987)
Estero Pargo Fringe 120.0 22.4 77.6 Day et al. (1996)
Estero Pargo Basin 49.4 Day et al. (1996)
Central America
Panama Riverine 280.0 Golley et al. (1975)
South America
French Guiana Riverine 188.6 2.8 97.2 Fromard et al. (1998)
French Guiana Riverine 122.2 2.8 97.2 Fromard et al. (1998)
French Guiana Fringe 31.5 9.1 91.8 Fromard et al. (1998)
French Guiana Fringe 71.8 6.0 94.0 Fromard et al. (1998)
French Guiana Fringe 35.1 10.2 89.8 Fromard et al. (1998)
French Guiana Basin 247.5 2.9 97.2 Fromard et al. (1998)
phosphorus availability and highly reduced soils play Feller et al. 1999). Our results suggest that both
a major role in determining tree growth and develop- factors may be important in explaining the partition-
ment (Twilley et al. 1996; Koch 1997; Feller 1995; ing pattern of dwarf R. mangle in South Florida.
Figure 2. Aboveground biomass partitioning at Taylor Slough. X-axis shows the number of prop roots in each tree that was used as a proxy for
increase in tree biomass.
162
Clough B.F., Dixon P. and Dalhaus O. 1997. Allometric relation-
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